Ki-Duk Song

Chonbuk National University, Tsiuentcheou, Jeollabuk-do, South Korea

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Publications (48)113.42 Total impact


  • No preview · Article · Dec 2015 · Journal of Microbiology and Biotechnology
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    ABSTRACT: Natural and artificial selection following domestication has led to the existence of more than a hundred pig breeds, as well as incredible variation in phenotypic traits. Berkshire pigs are regarded as having superior meat quality compared to other breeds. As the meat production industry seeks selective breeding approaches to improve profitable traits such as meat quality, information about genetic determinants of these traits is in high demand. However, most of the studies have been performed using trained sensory panel analysis without investigating the underlying genetic factors. Here we investigate the relationship between genomic composition and this phenotypic trait by scanning for signatures of positive selection in whole-genome sequencing data. We generated genomes of 10 Berkshire pigs at a total of 100.6 coverage depth, using the Illumina Hiseq2000 platform. Along with the genomes of 11 Landrace and 13 Yorkshire pigs, we identified genomic variants of 18.9 million SNVs and 3.4 million Indels in the mapped regions. We identified several associated genes related to lipid metabolism, intramuscular fatty acid deposition, and muscle fiber type which attribute to pork quality (TG, FABP1, AKIRIN2, GLP2R, TGFBR3, JPH3, ICAM2, and ERN1) by applying between population statistical tests (XP-EHH and XP-CLR). A statistical enrichment test was also conducted to detect breed specific genetic variation. In addition, de novo short sequence read assembly strategy identified several candidate genes (SLC25A14, IGF1, PI4KA, CACNA1A) as also contributing to lipid metabolism. Results revealed several candidate genes involved in Berkshire meat quality; most of these genes are involved in lipid metabolism and intramuscular fat deposition. These results can provide a basis for future research on the genomic characteristics of Berkshire pigs.
    Full-text · Article · Dec 2015 · BMC Genetics
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    J D Oh · Ki-Duk Song · E S Kim · H K Lee
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    ABSTRACT: c-MYC (v-myelocytomatosis viral oncogene homologue) is a transcription factor that plays important role in many biological process including cell growth and differentiation, such as myogenesis and adipogenesis. In this study, we aimed to detect MYC gene polymorphisms, their genotype frequencies and to determine associations between these polymorphisms and meat quality traits in Berkshire pigs. We identified a single nucleotide polymorphism (SNP) in intron 2 of MYC gene by Sanger sequencing, i.e., g.3350G>C (rs321898326), that is only found in Berkshire pigs, but not in other breeds including Duroc, Landrace, and Yorkshire pigs that were used in this study. Genotypes of total 378 Berkshire pigs (138 sows and 240 boars) were determined using Hha I restriction enzyme digestion after polymerase chain reaction. Observed allele frequencies of GG, GC, and CC genotypes were 0.399, 0.508, and 0.093 respectively. Statistical analysis indicated that the g.3350G>C polymorphism was significantly associated with pH45min and cooking loss (p<0.05), suggesting that g.3350G>C SNP can be used for pre-selection of pH45min and cooking loss traits in Berkshire pigs.
    Preview · Article · Sep 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: Embryonic stem cells (ESCs) have been used as a powerful tool for research including gene manipulated animal models and the study of developmental gene regulation. Among the critical regulatory factors that maintain the pluripotency and self-renewal of undifferentiated ESCs, NANOG plays a very important role. Nevertheless, because pluripotency maintaining factors and specific markers for livestock ESCs have not yet been probed, few studies of the NANOG gene from domestic animals including bovine have been reported. Therefore, we chose mouse ESCs in order to understand and compare NANOG expression between bovine, human, and mouse during ESCs differentiation. We cloned a 600 bp (-420/+181) bovine NANOG 5'-flanking region, and tagged it with humanized recombinant green fluorescent protein (hrGFP) as a tracing reporter. Very high GFP expression for bovine NANOG promoter was observed in the mouse ESC line. GFP expression was monitored upon ESC differentiation and was gradually reduced along with differentiation toward neurons and adipocyte cells. Activity of bovine NANOG (-420/+181) promoter was compared with already known mouse and human NANOG promoters in mouse ESC and they were likely to show a similar pattern of regulation. In conclusion, bovine NANOG 5-flanking region functions in mouse ES cells and has characteristics similar to those of mouse and human. These results suggest that bovine gene function studied in mouse ES cells should be evaluated and extrapolated for application to characterization of bovine ES cells.
    Preview · Article · Sep 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: Since ancient days, domestic horses have been closely associated with human civilization. Today, horse racing is an important industry. Various genes involved in energy production and muscle contraction are differentially regulated during a race. Among them, creatine kinase (CK) is well known for its regulation of energy preservation in animal cells. CK is an iso-enzyme, encoded by different genes and expressed in skeletal muscle, heart, brain and leucocytes. We confirmed that the expression of CK-M significantly increased in the blood after a 30 minute exercise period, while no considerable change was observed in skeletal muscle. Analysis of various tissues showed an ubiquitous expression of the CK-M gene in the horse; CK-M mRNA expression was predominant in the skeletal muscle and the cardiac muscle compared to other tissues. An evolutionary study by synonymous and non-synonymous single nucleotide polymorphism ratio of CK-M gene revealed a positive selection that was conserved in the horse. More studies are warranted in order to develop the expression of CK-M gene as a biomarker in blood of thoroughbred horses.
    Full-text · Article · Sep 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: Genetic parameters of Berkshire pigs for reproduction, carcass and meat quality traits were estimated using the records from a breeding farm in Korea. For reproduction traits, 2,457 records of the total number of piglets born (TNB) and the number of piglets born alive (NBA) from 781 sows and 53 sires were used. For two carcass traits which are carcass weight (CW) and backfat thickness (BF) and for 10 meat quality traits which are pH value after 45 minutes (pH45m), pH value after 24 hours (pH24h), lightness in meat color (LMC), redness in meat color (RMC), yellowness in meat color (YMC), moisture holding capacity (MHC), drip loss (DL), cooking loss (CL), fat content (FC), and shear force value (SH), 1,942 pig records were used to estimate genetic parameters. The genetic parameters for each trait were estimated using VCE program with animal model. Heritability estimates for reproduction traits TNB and NBA were 0.07 and 0.06, respectively, for carcass traits CW and BF were 0.37 and 0.57, respectively and for meat traits pH45m, pH24h, LMC, RMC, YMC, MHC, DL, CL, FC, and SH were 0.48, 0.15, 0.19, 0.36, 0.28, 0.21, 0.33, 0.45, 0.43, and 0.39, respectively. The estimate for genetic correlation coefficient between CW and BF was 0.27. The Genetic correlation between pH24h and meat color traits were in the range of -0.51 to -0.33 and between pH24h and DL and SH were -0.41 and -0.32, respectively. The estimates for genetic correlation coefficients between reproductive and meat quality traits were very low or zero. However, the estimates for genetic correlation coefficients between reproductive traits and drip and cooking loss were in the range of 0.12 to 0.17 and -0.14 to -0.12, respectively. As the estimated heritability of meat quality traits showed medium to high heritability, these traits may be applicable for the genetic improvement by continuous measurement. However, since some of the meat quality traits showed negative genetic correlations with carcass traits, an appropriate breeding scheme is required that carefully considers the complexity of genetic parameters and applicability of data.
    Preview · Article · Sep 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: Epoxide hydrolase 1 (EPHX1) plays an important role in both the activation and detoxification of exogenous chemicals. Reverse transcription-polymerase chain reaction (RT-PCR) analysis showed that the highest level of EPHX1 expression occurred in Berkshire liver, which is an organ that plays a key role in detoxification. We examined EPHX1 SNPs to analyze effect on increased expression of EPHX1 gene in Berkshire liver by total of 192 pigs of a pure Berkshire line (males = 97; females = 95). As a result, two nonsynonymous SNPs (nsSNPs) of EPHX1 were found from c.685T>G and c.776C > T, and located in 5th and 6th exons, respectively, which constitute the A/b hydrolase 1 domain of epoxide hydrolase. The nsSNP c.685T > G was significant differences in meat color, protein content, collagen content, and pH24 hr. Especially, T and G alleles of the nsSNP c.685T > G were significantly associated with CIE a*/CIE b* and protein content/pH24 hr, respectively. The nsSNP c.776C > T was significant differences in drip loss and protein content. Among meat quality traits to associate with SNPs, the protein content was only significantly associated with sex. Therefore, it is suggested that nsSNP c.685T > G in EPHX1 gene is a potential to apply as appropriate DNA markers for improvement of porcine economic traits.
    No preview · Article · Jul 2015 · Animal Biotechnology

  • No preview · Article · Jun 2015
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    ABSTRACT: The T-cell antigen receptor (TCR) complex contains 10 copies of a di-tyrosine Immunoreceptor-Tyrosine-based-Activation-Motif (ITAM) that initiates TCR signalling by recruiting protein tyrosine kinases. ITAM multiplicity amplifies TCR signals, but the importance of this capability for T-cell responses remains undefined. Most TCR ITAMs (6 of 10) are contributed by the CD3z subunits. We generated 'knock-in' mice that express non-signalling CD3z chains in lieu of wild-type CD3z. Here we demonstrate that ITAM multiplicity is important for the development of innate-like T cells and follicular helper T cells, events that are known to require strong/sustained TCR–ligand interactions, but is not essential for 'general' T-cell responses including proliferation and cytokine production or for the generation of a diverse antigen-reactive TCR repertoire.
    Full-text · Article · Jun 2015 · Nature Communications
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    ABSTRACT: Ginsenosides, the major active component of ginseng, are traditionally used to treat various diseases, including cancer, inflammation, and obesity. Among these, compound K (CK), an intestinal bacterial metabolite of the ginsenosides Rb1, Rb2, and Rc from Bacteroides JY-6, is reported to inhibit cancer cell growth by inducing cell-cycle arrest or cell death, including apoptosis and necrosis. However, the precise effect of CK on breast cancer cells remains unclear. MCF-7 cells were treated with CK (0-70 micrometer) for 24 or 48 h. Cell proliferation and death were evaluated by 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide (MTT) and flow cytometry assays, respectively. Changes in downstream signaling molecules involved in cell death, including phospho-glycogen synthase kinase 3β (GSK3β), β-catenin, and cyclin D1 were analyzed by Western blot. To block GSK3β signaling, MCF-7 cells were pretreated with GSK3β inhibitors 1 h prior to CK treatment. Cell death and the expression of β-catenin and cyclin D1 were then examined. CK dose- and time-dependently inhibited MCF-7 cell proliferation. Interestingly, CK induced programmed necrosis, but not apoptosis, via the GSK3β signaling pathway in MCF-7 cells. CK inhibited GSK3β phosphorylation, thereby suppressing the expression of β-catenin and cyclin D1. Our results suggest that CK induces programmed necrosis in MCF-7 breast cancer cells via the GSK3β signaling pathway.
    Full-text · Article · Jun 2015 · Journal of Microbiology and Biotechnology
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    ABSTRACT: The purpose of this study was to investigate the alternative splicing in equine cordon-bleu WH2 repeat protein-like 1 (COBLL1) gene that was identified in horse muscle and blood leukocytes, and to predict functional consequences of alternative splicing by bioinformatics analysis. In a previous study, RNA-seq analysis predicted the presence of alternative spliced isoforms of equine COBLL1, namely COBLL1a as a long form and COBLL1b as a short form. In this study, we validated two isoforms of COBLL1 transcripts in horse tissues by the real-time polymerase chain reaction, and cloned them for Sanger sequencing. The sequencing results showed that the alternative splicing occurs at exon 9. Prediction of protein structure of these isoforms revealed three putative phosphorylation sites at the amino acid sequences encoded in exon 9, which is deleted in COBLL1b. In expression analysis, it was found that COBLL1b was expressed ubiquitously and equivalently in all the analyzed tissues, whereas COBLL1a showed strong expression in kidney, spinal cord and lung, moderate expression in heart and skeletal muscle, and low expression in thyroid and colon. In muscle, both COBLL1a and COBLL1b expression decreased after exercise. It is assumed that the regulation of COBLL1 expression may be important for regulating glucose level or switching of energy source, possibly through an insulin signaling pathway, in muscle after exercise. Further study is warranted to reveal the functional importance of COBLL1 on athletic performance in race horses.
    Preview · Article · Jun 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: The adrenergic receptor beta 2 (ADRB2) plays a role in various physiological responses of the muscle to exercise, such as contraction and relaxation. Given its important role in muscle function, we investigated the structure of the horse ADRB2 gene and its expression pattern after exercise to determine if it can serve as a putative biomarker for recovery. Evolutionary analyses using synonymous and non-synonymous mutation ratios, were compared with other species (human, chimpanzee, mouse, rat, cow, pig, chicken, dog, and cat), and revealed the occurrence of positive selection in the horse ADRB2 gene. In addition, expression analyses by quantitative polymerase chain reaction exhibited ubiquitous distribution of horse ADRB2 in various tissues including lung, skeletal muscle, kidney, thyroid, appendix, colon, spinal cord and heart, with the highest expression observed in the lung. The expression of ADRB2 in skeletal muscle was significantly up-regulated about four folds 30 minutes post-exercise compared to pre-exercise. The expression level of ADRB2 in leukocytes, which could be collected with convenience compared with other tissues in horse, increased until 60 min after exercise but decreased afterward until 120 min, suggesting the ADRB2 expression levels in leukocytes could be a useful biomarker to check the early recovery status of horse after exercise. In conclusion, we identified horse ADRB2 gene and analyzed expression profiles in various tissues. Additionally, analysis of ADBR2 gene expression in leukocytes could be a useful biomarker useful for evaluation of early recovery status after exercise in racing horses.
    Preview · Article · May 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: While athletic abilities such as speed, endurance and recovery are important in the horse, genes related to these abilities have not been extensively investigated. Here, we characterized the horse peroxisome proliferator-activated receptor delta (PPARδ) gene and analyzed the expression of PPARδ during exercise. PPARδ is a known regulator of β-oxidation, muscle fiber transformation, and running endurance. Through evolutionary analysis using the synonymous and non-synonymous mutation ratio, it was revealed that positive selection occurred in the horse PPARδ gene. Two important domains related to nuclear hormone receptors, C4 zinc finger and ligand binding domain, were also found to be conserved well in horse PPARδ. Horse PPARδ was expressed ubiquitously in many tissues, but the expression level was various depending on the tissues. In the skeletal muscle, PPARδ increased about 2.5 folds after 30 min of exercise. Unlike in muscle, the increase of PPARδ expression was observed at 60 min but not 30 min of exercise in leukocytes. This finding might be useful for testing the endurance of horse using blood samples. Conclusively, the horse PPARδ gene is evolutionarily conserved well and can be used as a biomarker of endurance in horse.
    Preview · Article · May 2015 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: The influenza A virus infects a broad range of species and spreads easily through the respiratory tract. Because of these characteristics, the influenza A virus has caused pandemic disease in humans and livestock. To investigate the early molecular responses after influenza A virus infection in chickens, we infected tracheal epithelial cells derived from 20-day-old chick embryos with influenza A virus (H1N1). The gene expression patterns of the infected tracheal epithelial cells were analyzed via DNA microarray at different time points (0, 6, 12, 24, and 36 hr) after viral infection. Differentially expressed genes were identified in 6, 12, 24, and 36 hours post infection. A total of 1,936, 2,168, 3,670 and 2,894 genes were upregulated (≥2-fold, P < 0.05), whereas 884, 592, 1,503 and 1,925 genes were downregulated at the respective time points (≤0.5-fold, P < 0.05). When the differentially expressed genes were functionally categorized, immune-related and defense response gene ontology terms were detected in 12, 24, or 36 hours post infection. Interestingly, in the defense response, most of the gallinacin (GAL) genes were rapidly induced within 24 hr post infection. Subsequently, we predicted transcription factor binding sites within promoters of the GAL gene family, and analyzed the gene expression pattern for the common GAL gene regulatory factors to identify the viral infection-induced immune mechanism. Our results might contribute to an understanding of early host responses and regulatory mechanisms for host defense peptide induction against viral infections in chicken. © 2015 Poultry Science Association Inc.
    Preview · Article · Apr 2015 · Poultry Science
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    ABSTRACT: The Korean Native Chicken (KNC) is an important endemic biological resource in Korea. While numerous studies have been conducted exploring this breed, none have used next-generation sequencing to identify its specific genomic features. We sequenced five strains of KNC and identified 10.9 million SNVs and 1.3 million InDels. Through the analysis, we found that the highly variable region common to all 5 strains had genes like PCHD15, CISD1, PIK3C2A, and NUCB2 that might be related to the phenotypic traits of the chicken such as auditory sense, growth rate and egg traits. In addition, we assembled unaligned reads that could not be mapped to the reference genome. By assembling the unaligned reads, we were able to present genomic sequences characteristic to the KNC. Based on this, we also identified genes related to the olfactory receptors and antigen that are common to all 5 strains. Finally, through the reconstructed mitochondrial genome sequences, we performed phylogenomic analysis and elucidated the maternal origin of the artificially restored KNC. Our results revealed that the KNC has multiple maternal origins which are in agreement with Korea's history of chicken breed imports. The results presented here provide a valuable basis for future research on genomic features of KNC and further understanding of KNC's origin.
    Full-text · Article · Dec 2014 · PLoS ONE
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    ABSTRACT: There are five native chicken lines in Korea, which are mainly classified by plumage colors (black, white, red, yellow, gray). These five lines are very important genetic resources in the Korean poultry industry. Based on a next generation sequencing technology, whole genome sequence and reference assemblies were performed using Gallus_gallus_4.0 (NCBI) with whole genome sequences from these lines to identify common and novel single nucleotide polymorphisms (SNPs). We obtained 36,660,731,136 ± 1,257,159,120 bp of raw sequence and average 26.6-fold of 25-29 billion reference assembly sequences representing 97.288 % coverage. Also, 4,006,068 ± 97,534 SNPs were observed from 29 autosomes and the Z chromosome and, of these, 752,309 SNPs are the common SNPs across lines. Among the identified SNPs, the number of novel- and known-location assigned SNPs was 1,047,951 ± 14,956 and 2,948,648 ± 81,414, respectively. The number of unassigned known SNPs was 1,181 ± 150 and unassigned novel SNPs was 8,238 ± 1,019. Synonymous SNPs, non-synonymous SNPs, and SNPs having character changes were 26,266 ± 1,456, 11,467 ± 604, 8,180 ± 458, respectively. Overall, 443,048 ± 26,389 SNPs in each bird were identified by comparing with dbSNP in NCBI. The presently obtained genome sequence and SNP information in Korean native chickens have wide applications for further genome studies such as genetic diversity studies to detect causative mutations for economic and disease related traits.
    Full-text · Article · Oct 2014 · Molecular Biology Reports
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    ABSTRACT: Copy number variations (CNVs), important genetic factors for study of human diseases, may have as large of an effect on phenotype as do single nucleotide polymorphisms. Indeed, it is widely accepted that CNVs are associated with differential disease susceptibility. However, the relationships between CNVs and gene expression have not been characterized in the horse. In this study, we investigated the effects of copy number deletion in the blood and muscle transcriptomes of Thoroughbred racing horses. We identified a total of 1,246 CNVs of deletion polymorphisms using DNA re-sequencing data from 18 Thoroughbred racing horses. To discover the tendencies between CNV status and gene expression levels, we extracted CNVs of four Thoroughbred racing horses of which RNA sequencing was available. We found that 252 pairs of CNVs and genes were associated in the four horse samples. We did not observe a clear and consistent relationship between the deletion status of CNVs and gene expression levels before and after exercise in blood and muscle. However, we found some pairs of CNVs and associated genes that indicated relationships with gene expression levels: a positive relationship with genes responsible for membrane structure or cytoskeleton and a negative relationship with genes involved in disease. This study will lead to conceptual advances in understanding the relationship between CNVs and global gene expression in the horse.
    Full-text · Article · Sep 2014 · Asian Australasian Journal of Animal Sciences
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    ABSTRACT: The transcription factor ThPOK promotes CD4(+) T cell differentiation in the thymus. Here, using a mouse strain that allows post-thymic gene deletion, we show that ThPOK maintains CD4(+) T lineage integrity and couples effector differentiation to environmental cues after antigenic stimulation. ThPOK preserved the integrity and amplitude of effector responses and was required for proper differentiation of types 1 and 2 helper T cells in vivo by restraining the expression and function of Runx3, a nuclear factor crucial for cytotoxic T cell differentiation. The transcription factor LRF acts redundantly with ThPOK to prevent the transdifferentiation of mature CD4(+) T cells into CD8(+) T cells. As such, the ThPOK-LRF transcriptional module was essential for CD4(+) T cell integrity and responses.
    Full-text · Article · Aug 2014 · Nature Immunology
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    ABSTRACT: Background Ginseng, the root of Panax ginseng, has been extensively used in traditional oriental medicine and is a modern pharmaceutical reagent for the prevention of various human diseases, including cancer. Ginsenosides are the major active component of ginseng and exhibit immunomodulatory effects. However, the mechanism and function underlying such effects have not been fully elucidated, especially in human monocytes and dendritic cells (DCs). Methods We investigated the immunomodulatory effect of ginsenosides from the root of Panax ginseng on CD14+ monocytes purified from human adult peripheral blood mononuclear cells (PBMC) and differentiation into DCs that affect CD4+ T cell activity. Results The results showed that tumor necrosis factor (TNF)-α, interleukin (IL)-6 and IL-10 increased in monocytes upon treatment with ginsenoside fractions through phosphorylation of ERK1/2 (pERK1/2) and JNK, but not p38 MAP kinase. Interestingly, TNF-α production and phosphorylation of ERK1/2 and JNK decreased in lipopolysaccharide (LPS)-sensitized monocytes upon treatment with ginsenoside fractions. Next, we confirmed that DCs derived from CD14+ monocytes in the presence of ginsenoside fractions (Gin-DCs) contained decreased levels of the co-stimulatory molecules, CD80 and CD86. In the presence of ginsenoside fractions, expression of these co-stimulatory molecules decreased in LPS-treated DCs compared with LPS-treated DCs in the absence of ginsenoside fractions. Furthermore, Gin-DCs treated with LPS could not induce proliferation and IFN-γ production of CD4+ T cells at co-culture of Gin-DCs with CD4+ T cells. Conclusion These results suggest that ginsenoside fractions from the ginseng root suppress cytokine production and maturation of DCs treated with LPS, resulting in the down-regulation of CD4+ T cells.
    Full-text · Article · Aug 2014 · Journal of ginseng research
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    ABSTRACT: Pork is a major source of animal protein for humans. The subcutaneous, intermuscular and the intramuscular fat are the factors responsible for meat quality. RNA-seq is rapidly adopted for the profiling of the transcriptomes in the studies related to gene regulation. The discovery of differentially expressed genes (DEGs) between adult animals of Jeju Native Pig (JNP) and Berkshire breeds are of particular interest for the current study. RNA-seq was used to investigate the transcriptome profiling in the fat tissue. Sequence reads were obtained from Ilumina HiSeq2000 and mapped to the pig genome using Tophat2. Total 153 DEGs were identified and 71 among the annotated genes, have BLAST matches in the non- redundant database. Metabolic, immune response and protein binding are enriched pathways in the fat tissue. In our study, biological adhesion, cellular, developmental and multicellular organismal processes in fat were up-regulated in JNP as compare to Berkshire. Multicellular organismal process, developmental process, embryonic morphogenesis and skeletal system development were the most significantly enriched terms in fat of JNP and Berkshire breeds (p = 1.17E-04, 0.044, 3.47E-04 and 4.48E-04 respectively). COL10A1, COL11A2, PDK4 and PNPLA3 genes responsible for skeletal system morphogenesis and body growth were down regulated in JNP. This study is the first statistical analysis for the detection of DEGs from RNA-seq data generated from fat tissue sample. This analysis can be used as stepping stone to understand the difference in the genetic mechanisms that might influence the identification of novel transcripts, sequence polymorphisms, isoforms and noncoding RNAs.
    No preview · Article · Jul 2014 · Molecular Biology Reports

Publication Stats

139 Citations
113.42 Total Impact Points

Institutions

  • 2015
    • Chonbuk National University
      Tsiuentcheou, Jeollabuk-do, South Korea
    • Iowa State University
      Ames, Iowa, United States
  • 2013-2015
    • Hankyong National University
      • Genomic Information Center
      Anjŏ, Gyeonggi-do, South Korea
    • North Carolina State University
      • Department of Animal Science
      Raleigh, North Carolina, United States
  • 2011-2015
    • Seoul National University
      • • Department of Agricultural Biotechnology
      • • Research Institute for Agriculture and Life Science
      Sŏul, Seoul, South Korea
  • 2010-2014
    • Eunice Kennedy Shriver National Institute of Child Health and Human Development
      Роквилл, Maryland, United States