D. DEPARTMENT OF POPULATION GENETICS
D-a. Division of Population Genetics - Naruya Saitou Group

RESEARCH ACTIVITIES

Saitou Naruya, Professor

(1) DNA sequence and comparative analysis of chimpanzee chromosome 22

The International Chimpanzee Chromosome 22 Consortium [Hidemi Watanabe1, 2, Choong-Gon Kim, Satoshi Oota, Takashi Kitano, Yuji Kohara, Naruya Saitou and Yoshiyuki Sakaki2] ( 1Nara Institute of Science, and Technology,2RIKEN ,Genomic Sciences Center)

--Human-chimpanzee comparative genome research is essential for narrowing down genetic changes involved in the acquisition of unique human features, such as highly developed cognitive functions, bipedalism or the use of complex language. Here, we report the high-quality DNA sequence of 33.3 megabases of chimpanzee chromosome 22. By comparing the whole sequence with the human counterpart, chromosome 21, we found that 1.44% of the chromosome consists of single-base substitutions in addition to nearly 68,000 insertions or deletions. These differences are sufficient to generate changes in most of the proteins. Indeed, 83% of the 231 coding sequences, including functionally important genes, show differences at the amino acid sequence level.
--Furthermore, we demonstrate different expansion of particular subfamilies of retrotransposons between the lineages, suggesting different impacts of retrotranspositions on human and chimpanzee evolution. The genomic changes after speciation and their biological consequences seem more complex than originally hypothesized. For details, see ref. (1)

(2) Human specific amino acid changes found in 103 protein coding genes. Molecular Biology and Evolution

Takashi Kitano, Yuhua Liu1, Shintaroh Ueda2 and Naruya Saitou (1The Jackson Laoratory, USA,2Department of Biological Sciences, Graduate School of Science, University of Tokyo)

--We humans have many characteristics that are different from those of the great apes. These human-specific characters must have arisen through mutations accumulated in the genome of our direct ancestor after the divergence of the last common ancestor with chimpanzee. Gene trees of human and great apes are necessary for extracting these human-specific genetic changes. We conducted a systematic analysis of 103 protein-coding genes for human, chimpanzee, gorilla, and orangutan. Nucleotide sequences for 18 genes were newly determined for this study, and those for the remaining genes were retrieved from the DDBJ/EMBL/GenBank database. The total number of amino acid changes in the human lineage was 147 for 26,199 codons (0.56%). The total number of amino acid changes in the human genome was, thus, estimated to be about 80,000. We applied the acceleration index test and Fisher's synonymous/nonsynonymous exact test for each gene tree to detect any human-specific enhancement of amino acid changes compared with ape branches. Six and two genes were shown to have significantly higher nonsynonymous changes at the human lineage from the acceleration index and exact tests, respectively. We also compared the distribution of the differences of the nonsynonymous substitutions on the human lineage and those on the great ape lineage. Two genes were more conserved in the ape lineage, whereas one gene was more conserved in the human lineage. These results suggest that a small proportion of protein-coding genes started to evolve differently in the human lineage after it diverged from the ape lineage. For details, see ref. (2)

(3) Genetic variation versus recombination rate in a structured population of mice

Aya Takahashi, Yuhua Liu1 and Naruya Saitou (1The Jackson Laoratory, USA)

--The correlation between genetic variation and recombination rate was investigated in a structured mouse population. Nucleotide sequence data from 19 autosomal DNA loci from eight inbred strains of mouse (Mus musculus) sampled from three major subspecies were analyzed. The recombination rate was estimated from the comparison of genetic and physical map distances between markers flanking a 10-cM region of each locus. The strains were categorized into four groups (subpopulations) based on geography. By partitioning the genetic diversity into within-group and among-group variation, we detected a positive correlation between the recombination rate and nucleotide diversity within groups. The level of nucleotide differentiation among groups (G(ST)) showed a negative correlation with the rate of recombination. There was no significant correlation between recombination rate and nucleotide diversity when data from different subpopulations were pooled. No correlation was detected between recombination rate and nucleotide divergence of M. musculus and M. spicilegus. These patterns deviate from the strict neutral expectation under the constant nucleotide substitution rate, and they are likely to have been formed either by a hitchhiking effect of positively selected mutants or by background selection of deleterious mutants occurring in a subdivided population. Our series of comparisons show that because a real population always has some structure, incorporation of its information is important in detecting non-neutral evolution. For details, see ref. (3)

(4) Phylogenetic analysis of proteins associated in four major energy metabolism systems: photosynthesis, oxidative phosphorylation, nitrogen metabolism and sulfur metabolism

Takeshi Tomiki and Naruya Saitou

--The four electron transfer energy metabolism systems, photosynthesis, aerobic respiration, denitrification, and sulfur respiration, are thought to be evolutionarily related because of the similarity of electron transfer patterns and the existence of some homologous proteins. How these systems have evolved is elusive. We therefore conducted a comprehensive homology search using PSI-BLAST, and phylogenetic analyses were conducted for the three homologous groups (groups 1-3) based on multiple alignments of domains defined in the Pfam database. There are five electron transfer types important for catalytic reaction in group 1, and many proteins bind molybdenum. Deletions of two domains led to loss of the function of binding molybdenum and ferredoxin, and these deletions seem to be critical for the electron transfer pattern changes in group 1. Two types of electron transfer were found in group 2, and all its member proteins bind siroheme and ferredoxin. Insertion of the pyridine nucleotide disulfide oxidoreductase domain seemed to be the critical point for the electron transfer pattern change in this group. The proteins belonging to group 3 are all flavin enzymes, and they bind flavin adenine dinucleotide (FAD) or flavin mononucleotide (FMN). Types of electron transfer in this group are divergent, but there are two common characteristics. NAD(P)H works as an electron donor or acceptor, and FAD or FMN transfers electrons from/to NAD(P)H. Electron transfer functions might be added to these common characteristics by the addition of functional domains through the evolution of group 3 proteins. Based on the phylogenetic analyses in this study and previous studies, we inferred the phylogeny of the energy metabolism systems as follows: photosynthesis (and possibly aerobic respiration) and the sulfur/nitrogen assimilation system first diverged, then the sulfur/nitrogen dissimilation system was produced from the latter system. For details, see ref. (4)

(5) Mitochondrial DNA Genealogy of Chimpanzees in Nimba Mountains and Bossou, West Africa

Makoto Shimada, Sachiko Hayakawa1, Tatyana Humle2, Shiho Fujita3, Satoshi Hirata4, Yukimaru Sugiyama5 and Naruya Saitou (1Primate Research Institute, Kyoto University,2University of Stirling United Kingdom,3Faculty of Agriculture, Gifu University, 4Hayashibara Biochemical Laboratories Inc., 5Tokai-Gakuen University)

--The chimpanzee populations of the Bossou and Nimba regions in West Africa were genetically surveyed to 1) reveal the genetic relationship between the Bossou and Nimba populations, and 2) elucidate the evolutionary relationship between the Bossou-Nimba and other West African populations. The chimpanzee group at Bossou is characterized by its small population size, no evidence of contact with neighboring populations, and no female immigration. It is believed that most females and adolescent males emigrate from this population. To reveal the genetic signature of these characteristics, we examined the genetic diversity of Bossou and two neighboring populations (Seringbara and Yeale) in the Nimba Mountains by sequencing approximately 605 bp of the mitochondrial DNA (mtDNA) control region. A total of 20 distinct mtDNA variants were observed from 56 sequences of noninvasively collected, anonymous samples. Nucleotide diversity in the Nimba Mountain populations was 0.03-0.04, and did not differ significantly from that in the Bossou population. Very few mitochondrial variants are shared among the sites sampled, which suggests that there is little gene flow involving mtDNA. Nevertheless, no clear population structures were revealed in either population. A comparison with published sequences from West African chimpanzees (Pan troglodytes verus) indicates that the variants observed in the Bossou and Nimba regions are scattered throughout the subspecies, rather than clustered according to geographic region. This suggests that the Bossou-Nimba populations derived only recently from the common ancestral population of the West African chimpanzees, and did not pass through a bottleneck. For details, see ref. (5)

(6) Evolution of O alleles of the human ABO blood group gene

Francis Roubinet1, Stephanie Despiau1, Francesc Calafell2, Fen Jin3, Jaume Bertanpetit2, Naruya Saitou and Antoine Blancher1 (1Paul Sabatier University, Rangueil Hospital,2Universitat Pompeu Fabra,3The Institute of Genetics and Developmental Biology)

--To date, at least 40 different alleles O have been characterized on the basis of exon 6 and exon 7 sequences but not always for intron 6. STUDY DESIGN AND METHODS: Among 415 individuals, from four continents (Africa, Europe, South America, and Asia), studied for exon 6 and exon 7 sequences, we selected 46 individuals (of respective phenotypes O [39], AB [3], B [3], or A [1]) for sequencing 1800-bp amplicons spanning exon 6, intron 6, and exon 7. The amplicons were characterized either by direct sequencing or after cloning when required. We defined 14 new intron 6 O allele sequences, including four recombinant alleles. Based on sequence comparison, a phylogenetic network was constructed. It confirmed recombinant allele origins and that most O alleles are derived by point mutations from the two worldwide distributed alleles O01 and O02. CONCLUSION: Allele O phylogenetic analysis suggests that the most frequent silencing mutation (deletion of a G in exon 6) appeared once in human evolution in the ancient O02 allele lineage and that allele O01 resulted from an interallele exchange between O02 and A101. Assuming constancy of evolutionary rate, diversification of the representative alleles of the three human ABO lineages (A101, B101, and O02) was estimated at 4.5 to 6 million years ago. For details, see ref. (6)

(7) Comparative genetics of functional trinucleotide tandem repeats in humans and apes

Aida Andres1, Marta Soldevila1, Oscar Lao1, Victor Volpini1, Nayuya Saitou, Howard Jacobs1, Ikuo Hayasaka2, Francesc Calafell1 and Jaume Bertranpetit1 (1Universitat Pompeu Fabra, 2Kumamoto Primate Park, Sanwa Kagaku Kenkyusho)

--Several human neurodegenerative disorders are caused by the expansion of polymorphic trinucleotide repeat regions. Many of these loci are functional short tandem repeats (STRs) located in brain-expressed genes, and their study is thus relevant from both a medical and an evolutionary point of view. The aims of our study are to infer the comparative pattern of variation and evolution of this set of loci in order to show species-specific features in this group of STRs and on their potential for expansion (therefore, an insight into evolutionary medicine) and to unravel whether any human-specific feature may be identified in brain-expressed genes involved in human disease. We analyzed the variability of the normal range of seven expanding STR CAG/CTG loci (SCA1, SCA2, SCA3-MJD, SCA6, SCA8, SCA12, and DRPLA) and two nonexpanding polymorphic CAG loci (KCNN3 and NCOA3) in humans, chimpanzees, gorillas, and orangutans. The study showed a general conservation of the repetitive tract and of the polymorphism in the four species and high heterogeneity among loci distributions. Humans present slightly larger alleles than the rest of species but a more relevant diference appears in variability levels: Humans are the species with the largest variance, although only for the expanding loci, suggesting a relationship between variability levels and expansion potential. The sequence analysis shows high levels of sequence conservation among species, a lack of correspondence between interruption patterns and variability levels, and signs of conservative selective pressure for some of the STR loci. Only two loci (SCA1 and SCA8) show a human specific distribution, with larger alleles than the rest of species. This could account, at the same time, for a human-specific trait and a predisposition to disease through expansion. For details, see ref. (7)

(8) Polymorphisms in the Trace Amine Receptor 4 (TRAR4) Gene on Chromosome 6q23.2 Are Associated with Susceptibility to Schizophrenia

Jubao Duan1, Alan Sanders1, Cuiping Hou1, Naruya Saitou, Takashi Kitano and Pablo Gejman1 (1Department of Psychiatry and Behavioral Sciences, Northwestern University)

--Several linkage studies across multiple population groups provide convergent support for a susceptibility locus for schizophrenia-and, more recently, for bipolar disorder-on chromosome 6q13-q26. We genotyped 192 European-ancestry and African American (AA) pedigrees with schizophrenia from samples that previously showed linkage evidence to 6q13-q26, focusing on the MOXD1-STX7-TRARs gene cluster at 6q23.2, which contains a number of prime candidate genes for schizophrenia. Thirty-one screening single-nucleotide polymorphisms (SNPs) were selected, providing a minimum coverage of at least 1 SNP/20 kb. The association observed with rs4305745 (P=.0014) within the TRAR4 (trace amine receptor 4) gene remained significant after correction for multiple testing. Evidence for association was proportionally stronger in the smaller AA sample. We performed database searches and sequenced genomic DNA in a 30-proband subsample to obtain a high-density map of 23 SNPs spanning 21.6 kb of this gene. Single-SNP analyses and also haplotype analyses revealed that rs4305745 and/or two other polymorphisms in perfect linkage disequilibrium (LD) with rs4305745 appear to be the most likely variants underlying the association of the TRAR4 region with schizophrenia. Comparative genomic analyses further revealed that rs4305745 and/or the associated polymorphisms in complete LD with rs4305745 could potentially affect gene expression. Moreover, RT-PCR studies of various human tissues, including brain, confirm that TRAR4 is preferentially expressed in those brain regions that have been implicated in the pathophysiology of schizophrenia. These data provide strong preliminary evidence that TRAR4 is a candidate gene for schizophrenia; replication is currently being attempted in additional clinical samples. For details, see ref. (8)

Kenta Sumiyama, Assistant Professor

(9) Evolutionary functional genomic analysis of cis-regulation in mammalian Dlx gene system

Kenta Sumiyama

--Dlx genes are important regulators for mammalian early development. Mammalian Dlx genes has been duplicated through evolution and resulted in 6 genes in 3 clusters that have overlapping but also distinct functions. In branchial arches three Dlx gene clusters are expressed in nested pattern that is essential to establish positional information in jaw patterning. This nested expression pattern is found only in higher vertebrates so that studying genomic regulation mechanism through cis-regulatory elements would be of great interest in mammalian jaw evolution. Recent expansion of genome sequence information enables us to compare intergenic noncoding genomic sequences among different species. Although intergenic regions have been sometimes called “junk DNA" due to lack of knowledge about their functional importance, a large number of sequence conservations have been found in intergenic regions through comparative genomic sequence analysis. Such conservations are likely to be functional elements that are repressing, initiating or maintaining gene expressions. I focus on genomic analysis of cis-regulatory elements in Dlx3-7 gene clusters by using comparative genomics and further functional analysis by using transgenic mouse system.

PUBLICATIONS

Papers
1. The International Chimpanzee Chromosome 22 Consortium [Watanabe, H., Fujiyama, A., Hattori, M., Taylor, T.D., Toyoda, A., Kuroki, Y., Noguchi, H., Ben, Kahla, A., Lehrach, H., Sudbrak, R., Kube, M., Taenzer, S., Galgoczy, P., Platzer, M., Scharfe, M., Nordsiek, G., Blocker, H., Hellmann, I., Khaitovich, P., Paabo, S., Reinhardt, R., Zheng, H.-J., Zhang, X.-L., Zhu, G.-F., Wang, B.-F., Fu, G., Ren9, S.-X., Zhao, G.-P., Chen, Z., Lee, Y.-S., Cheong, J.-E., Choi, S.-H., Wu, K.-M., Liu, T.-T., Hsiao, K.-J., Tsai, S.-F., Kim, C.-G., Oota, S., Kitano, T., Kohara, Y., Saitou, N., Park, H.-S., Wang, S.-Y., Yaspo, M.-L. and Sakaki, Y.] (2004). DNA sequence and comparative analysis of chimpanzee chromosome 22. Nature, 429, 382-388.
2. Kitano, T., Liu, Y.-H., Ueda, S. and Saitou, N. (2004). Human specific amino acid changes found in 103 protein coding genes. Molecular Biology and Evolution. 21, 936-944.
3. Takahashi, A., Liu, Y.-H. and Saitou, N. (2004). Genetic variation versus recombination rate in a structured population of mice. Molecular Biology and Evolution, 21, 404-409.
4. Tomiki, T. and Saitou, N. (2004). Phylogenetic analysis of proteins associated in four major energy metabolism systems: photosynthesis, oxidative phosphorylation, nitrogen metabolism and sulfur metabolism. Journal of Molecular Evolution, 59, 158-176.
5. Shimada, M., Hayakawa, S., Hamle, T., Fujita, S., Hirata, S., Sugiyama, Y. and Saitou, N. (2004). Mitochondrial DNA Genealogy of Chimpanzees in Nimba Mountains and Bossou, West Africa. American Journal of Primatology, 64, 261-275.
6. Roubinet, F., Despiau, S.-P., Calafell, F., Jin, F., Bertanpetit, J., Saitou, N. and Blancher, A. (2004). Evolution of O alleles of the human ABO blood group gene. Transfusion, 44, 707-715.
7. Andres, A.M., Soldevila, M., Lao, O., Volpini, V., Saitou, N., Jacobs, H.T., Hayasaka, I., Calafell, F. and Bertranpetit, J. (2004). Comparative genetics of functional trinucleotide tandem repeats in humans and apes. Journal of Molecular Evolution, 59, 329-339.
8. Duan, J., Martinez, M., Sanders, A.R., Hou, C., Saitou, N., Kitano, T., Mowry, B.J., Crowe, R.R., Silverman, J.M., Levinson, D.F. and Gejman, P.V. (2004). Polymorphisms in the Trace Amine Receptor 4 (TRAR4) Gene on Chromosome 6q23.2 Are Associated with Susceptibility to Schizophrenia. American Journal of Human Genetics, 75, 624-638.
9. Abe, K., Noguchi, H., Tagawa, K., Yuzuriha, M., Toyoda, A., Kojima, T., Ezawa, K., Saitou, N., Hattori, M., Sakaki, Y., Moriwaki, K. and Shiroishi, T. (2004). Contribution of Asian mouse subspecies Mus musculus molossinus to genomic constitution of strain C57BL/6J, as defined by BAC end sequence-SNP analysis. Genome Research, 14, 2239-2247.
10. Imanishi, T. and many other authors including Saitou, N. and Sugano, S. (2004). Integrative annotation of 21,037 human genes validated by full-length cDNA clones. PLoS Biology, 2, 1-20.
11. Nakajima, T., Wooding, S., Sakagami, T., Emi, M., Tokunaga, K., Tamiya, G., Ishigami, T., Umemura, S., Munkhbat, B., Jin, F., Guan-Jun, J., Hayasaka, I., Ishida, T., Saitou, N., Pavelka, K., Lalouel, J.M., Jorde, L.B. and Inoue, I. (2004). Natural selection and population history in the human angiotensinogen gene (AGT): 736 complete AGT sequences in chromosomes from around the world. American Journal of Human Genetics, 74, 898-916.
12. 斎藤成也(2004)近隣結合法をはじめとする距離行列法.連載「ゲノム進化学の展開」第4回.数理科学,42巻12号.60-67頁.
13. 斎藤成也(2004)人類進化学の将来像.科学,74巻10号,1246-1249頁.
14. 斎藤成也(2004)系統樹とは(その2).連載「ゲノム進化学の展開」第3回.数理科学,42巻10号,72-78頁.
15. 斎藤成也(2004)『ゲノムと進化―ゲノムから立ち昇る生命―』.ワードマップシリーズ,新曜社(9月10日発行).
16. 斎藤成也(2004)言語能力の遺伝的基礎.大航海,52号,114-121頁.
17. 斎藤成也(2004)系統樹とは(その1).連載「ゲノム進化学の展開」第2回.数理科学,42巻9号,68-74頁.
18. 瀬名秀明・斎藤成也(2004)類人猿のゲノムで探る人間らしさの起源(対談).瀬名秀明編『科学の最前線で研究者は何を見ているのか』,52-70頁.(日経サイエンス,2003年6月号,94-99頁所収より)
19. 斎藤成也・隅山健太(2004)Hox遺伝子クラスターの進化―脊椎動物の誕生からヒトまで―.実験医学,Vol.22, No.12(8月号),1677-1683頁.
20. 斎藤成也(2004)ネアンデルタールシンポジウムに参加して.GSJコミュニケーションズ,79巻2号,7-9頁.
21. 北野誉・斎藤成也(2004)類人猿とヒトのゲノム進化研究.生体の科学,[特集]分子進化学の現在,55巻3号,252-256頁.
22. 斎藤成也(2004)はじめに.連載「ゲノム進化学の展開」第1回.数理科学,42巻7号,64-70頁.
23. 隅山健太・斎藤成也(2004)HOX遺伝子.榊佳之・笹月健彦・油谷浩幸編『ヒトゲノム―生命システムの理解と医学への展開―』,91-93頁.中山書店.
24. Lubert Stryer著,入村達郎ら共訳(2004)『ストライヤー生化学(第5版)』.2章・7章の翻訳.東京化学同人.

ORAL PRESENTATIONS

1. Saitou N.: Comparative genomics of human and apes. Heron Island, Invited talk Australia, February
2. Sumiyama K.: Gene duplication and functional diversification in mammalian Dlx genes, International Mammalian Symposium, Invited talk Hayama Japan, February
3. Saitou N.: Silver project: simultaneous analysis of human and ape genome sequences. HOPE symposium, Invited talk Kyoto, Japan, March
4. Sumiyama K.: Evolution of microRNA and other non-coding RNAs within Hox clusters. Yale University. Informal seminar USA March
5. Saitou N.: Comparative genome analysis of human/ape and mouse/rat. KRIBB, Invited talk Deajong, Korea, April
6. Sumiyama K.: Functional genomic analysis of cis-regulation in mammalian Dlx gene system. Symposium on Comparative Genomics, KRIBB, Invited talk Deajong, Korea, April
7. Saitou N.: Human and hominoid specific changes and conservation during primate evolution. Genomes & Evolution 2004, Invited talk Penn-State University, State College. USA, June
8. Saitou N.: Human and ape genome comparison as a model for evolutionary genomics of closely related species. Symosium on human evolution. SESJ annual meeting Invited talk University of Tokyo, Tokyo, Japan, August
9. Saitou N.: What can we extract from comparative genomic analysis? Plenary Lecture ICONIP, Kolcutta India, November
10. Saitou N.: Human and ape comparative genomics. Indian Statistical Institute, Informal talk Kolcutta India, November
11. Saitou N.: Before and after encountering neutral theory. Mishima Workshop, Invited talk Hakone, Japan, November
12. 斎藤成也:ネアンデルタールとは何者:遺伝子から視る.第18回大学と科学公開シンポジウム「アイデンティティに悩むネアンデルタール」.国際交流館,東京,1月.
13. 斎藤成也:人間性の基礎を与えるヒトゲノム中の遺伝子―霊長類比較ゲノム研究から.「ゲノム」4領域成果公開シンポジウム「生命システムの理解をめざして」.コクヨホール,東京,1月.
14. 斎藤成也:DNAから見たヒトの進化史.サイエンス講演会.山梨県立科学館,甲府,1月.
15. 斎藤成也:霊長類の比較ゲノム解析.ゲノム科学研究集会.名古屋観光ホテル,名古屋,2月.
16. 隅山健太:「ほ乳類顎発生関連遺伝子Dlx3-7クラスターの発現調節機構解析」東北大学加齢医学研究所研究員会セミナー,仙台,3月.
17. 斎藤成也:遺伝子とゲノムから見た生物の進化.日本機会学会,第23回バイオサロン.信濃町煉瓦館,東京,4月.
18. 斎藤成也:情報生物学の展開.ワークショップ「情報生物学の展開」.総合研究大学院大学,葉山,5月.
19. 斎藤成也:近隣結合法の開発とその後の発展.第12回木原記念財団学術賞受賞講演.横浜市立大学木原生物学研究所,横浜,5月.
20. 斎藤成也:人間へとたどるゲノムの進化.スーパーサイエンスハイスクール講義.山梨県立甲府南高等学校,甲府,6月.
21. 斎藤成也:日本における類人猿資源の遺伝学への利用〜特に比較ゲノム解析について〜.日本霊長類学会(自由集会)フロイデ犬山,犬山,7月.
22. 斎藤成也:新しいHPを使ったDDBJing.DDBJing講習会.国立遺伝学研究所,三島,7月.
23. 斎藤成也:ABO式血液型遺伝子を中心とする糖転移酵素遺伝子の進化.FCCA第9回糖質若手フォーラム,特別講演.東京大学薬学部,東京,7月.
24. 斎藤成也:古代DNAを用いた中国歴史時代の人間の移動の推定.中国語方言科学研究費補助金班会議,金沢大学,金沢,7月.
25. 斎藤成也:環境としてのゲノム.三島市民環境大学.日本大学国際関係学部,三島,8月.
26. 斎藤成也:遺伝子の系統樹と系統ネットワーク.第3回「数学者のための分子生物学入門」.けいはんなプラザ,京都,9月.
27. 斎藤成也:近縁な塩基配列の系統ネットワークを構築するSSJ法とその可視化ソフトPhyloNet.日本遺伝学会第76回大会,大阪大学,吹田市,9月.
28. 隅山健太「マウス鰓弓内で遠近軸情報を規定するDlx3遺伝子の発現調節機構の解析」特定領域研究発生システムのダイナミクス班会議,湘南国際村センター,葉山,9月.
29. 斎藤成也:人間への進化の過程を遺伝子でたどる.国立遺伝学研究所公開講演会.東京,10月.
30. 斎藤成也:哺乳類を例とした比較ゲノム解析の諸手法.第21回資源生物科学シンポジウム.倉敷市,12月.
31. 斎藤成也:ゲノム進化-生命を統合的に理解するための出発点.葉山セミナー.総合研究大学院大学,葉山,12月.
32. 隅山健太「発生調節遺伝子群Dlx3-7の発現調節cis-因子の同定とその進化」第27回日本分子生物学会年会,神戸市,12月.

EDUCATION

1. 斎藤成也:福井大学医学部法医学非常勤講義「中立進化とゲノムの進化」,3月
2. 斎藤成也:総合研究大学院大学先導科学研究科生命体科学専攻集中講義「系統樹解析の理論と問題点について」・「ヒトと類人猿の比較ゲノム解析」,5月
3. 斎藤成也:東京大学理学部講義「分子進化学」5月,6月
4. 斎藤成也:熊本大学理学部集中講義「遺伝子とゲノムから見た生物の進化」,11月
5. 斎藤成也:埼玉大学理学部集中講義「遺伝子の進化と生物の進化」,12月

SOCIAL CONTRIBUTIONS AND OTHERS

1. 斎藤成也:Molecular Biology and Evolution, Associate Editor
2. 斎藤成也:GENE-Evolutionary Genetics, Editor
3. 斎藤成也:シリーズ進化学全7巻(岩波書店) 編集委員
4. 斎藤成也:日本遺伝学会 会計幹事
5. 斎藤成也:日本DNA多型学会 評議員
6. 斎藤成也:日本人類学会 評議員
7. 斎藤成也:財団法人遺伝学普及会 監事
8. 斎藤成也:公益信託進化学振興木村資生基金 運営委員|
9. 斎藤成也:木原記念横浜生命科学振興財団より 第12回木原記念財団学術賞 受賞