Host control of persistent Epstein–Barr virus infection

A Axel Schmidt T T. Madhusankha Alawathurage F Friederike S. David Y Yosuke Ogawa L Leonard Frach S Sylvia Richter M Merle Schaefer C Carina M. Mathey S Sabrina K. Henne G Genta Nagao H Hiromu Tanaka S Shuhei Azekawa K Ko Lee N Naoki Fukunaga J Junko Hamamoto H Hiroki Kabata K Katsunori Masaki H Hirofumi Kamata S Shinnosuke Ikemura S Shotaro Chubachi S Satoshi Okamori H Hideki Terai A Atsuho Morita T Takanori Asakura M Makoto Ishii K Koichi Fukunaga Y Yoshifumi Uwamino S Sho Uchida S Shunsuke Uno T Tomoyasu Nishimura N Naoki Hasegawa E Emmy Yanagita H Hiroshi Nishihara J Junichi Sasaki H Hiroshi Morisaki T Toshiro Sato Y Yuko Kitagawa Y Yuta Matsubara Y Yohei Mikami K Kosaku Nanki T Takanori Kanai R Ryuya Edahiro Y Yuya Shirai K Kyuto Sonehara D Daisuke Okuzaki D Daisuke Motooka M Masahiro Kanai T Tatsuhiko Naito K Kenichi Yamamoto Q Qingbo S. Wang Y Yasuhiro Kato T Takayoshi Morita S Shinichi Namba K Ken Suzuki Y Yoko Naito Y Yu-Chen Liu A Ayako Takuwa F Fuminori Sugihara J James B. Wing S Shuhei Sakakibara N Nobuyuki Hizawa T Takayuki Shiroyama S Satoru Miyawaki Y Yusuke Kawamura A Akiyoshi Nakayama H Hirotaka Matsuo Y Yuichi Maeda T Takuro Nii Y Yoshimi Noda T Takayuki Niitsu Y Yuichi Adachi T Takatoshi Enomoto S Saori Amiya R Reina Hara Y Yuta Yamaguchi T Teruaki Murakami T Tomoki Kuge K Kinnosuke Matsumoto Y Yuji Yamamoto M Makoto Yamamoto M Midori Yoneda T Toshihiro Kishikawa S Shuhei Yamada S Shuhei Kawabata N Noriyuki Kijima M Masatoshi Takagaki N Noah Sasa Y Yuya Ueno M Motoyuki Suzuki N Norihiko Takemoto H Hirotaka Eguchi T Takahito Fukusumi T Takao Imai M Munehisa Fukushima H Haruhiko Kishima H Hidenori Inohara K Kazunori Tomono K Kazuto Kato M Meiko Takahashi F Fumihiko Matsuda H Haruhiko Hirata Y Yoshito Takeda A Atsushi Kumanogoh T Takanori Hasegawa K Kunihiko Takahashi T Tatsuhiko Anzai S Satoshi Ito Y Yuji Uchimura A Akifumi Endo Y Yasunari Miyazaki T Takayuki Honda T Tomoya Tateishi S Shuji Tohda N Naoya Ichimura K Kazunari Sonobe C Chihiro Tani Sassa J Jun Nakajima M Masumi Ai R Ryuji Koike A Akinori Kimura S Satoru Miyano T Tomomi Takano K Kazuhiko Katayama K Koji Okudela R Ryunosuke Saiki Y Yasuhito Nannya S Seishi Ogawa T Takayoshi Hyugaji E Eigo Shimizu K Kotoe Katayama S Seiya Imoto Y Yosuke Omae K Katsushi Tokunaga T Takafumi Ueno Y Yoshinori Fukui H Hiroyuki Hayashi Y Yukihiro Yoshimura N Natsuo Tachikawa K Kazuhisa Takahashi N Norihiro Harada Y Yuki Tanabe T Toshio Naito M Makoto Hiki Y Yasushi Matsushita H Haruhi Takagi R Ryousuke Aoki A Ai Nakamura S Sonoko Harada H Hitoshi Sasano T Takashi Ishiguro T Taisuke Isono S Shun Shibata Y Yuma Matsui C Chiaki Hosoda K Kenji Takano T Takashi Nishida Y Yoichi Kobayashi Y Yotaro Takaku N Noboru Takayanagi S Soichiro Ueda N Natsumi Yazaki A Ai Tada M Masayoshi Miyawaki M Masaomi Yamamoto E Eriko Yoshida R Reina Hayashi T Tomoki Nagasaka S Sawako Arai Y Yutaro Kaneko K Kana Sasaki E Etsuko Tagaya M Masatoshi Kawana K Ken Arimura Y Yasushi Nakano Y Yukiko Nakajima R Ryusuke Anan R Ryosuke Arai Y Yuko Kurihara Y Yuko Harada K Kazumi Nishio T Tetsuya Ueda M Masanori Azuma R Ryuichi Saito T Toshikatsu Sado Y Yoshimune Miyazaki R Ryuichi Sato Y Yuki Haruta T Tadao Nagasaki Y Yoshinori Yasui Y Yoshinori Hasegawa A Akihiro Noda Y Yusei Fukushima R Reina Kitagawa Y Yoshikazu Mutoh T Tomoki Kimura T Tomonori Sato R Reoto Takei S Satoshi Hagimoto Y Yoichiro Noguchi Y Yasuhiko Yamano H Hajime Sasano S Sho Ota Y Yasushi Nakamori K Kazuhisa Yoshiya F Fukuki Saito T Tomoyuki Yoshihara D Daiki Wada H Hiromu Iwamura S Syuji Kanayama S Shuhei Maruyama T Takashi Yoshiyama K Ken Ohta H Hiroyuki Kokuto H Hideo Ogata Y Yoshiaki Tanaka K Kenichi Arakawa M Masafumi Shimoda T Takeshi Osawa H Hiroki Tateno I Isano Hase S Shuichi Yoshida S Shoji Suzuki M Miki Kawada H Hirohisa Horinouchi F Fumitake Saito K Keiko Mitamura M Masao Hagihara J Junichi Ochi T Tomoyuki Uchida R Rie Baba D Daisuke Arai T Takayuki Ogura H Hidenori Takahashi S Shigehiro Hagiwara S Shunichiro Konishi I Ichiro Nakachi K Koji Murakami M Mitsuhiro Yamada H Hisatoshi Sugiura H Hirohito Sano S Shuichiro Matsumoto N Nozomu Kimura Y Yoshinao Ono H Hiroaki Baba Y Yusuke Suzuki S Sohei Nakayama K Keita Masuzawa H Hidefumi Koh T Tadashi Manabe Y Yohei Funatsu F Fumimaro Ito T Takahiro Fukui K Keisuke Shinozuka S Sumiko Kohashi M Masatoshi Miyazaki T Tomohisa Shoko T Takashi Inoue T Takahiro Asami T Toshiyuki Hirano K Keigo Kobayashi H Hatsuyo Takaoka K Kazuyoshi Watanabe N Naoki Miyazawa Y Yasuhiro Kimura R Reiko Sado H Hideyasu Sugimoto A Akane Kamiya N Naota Kuwahara A Akiko Fujiwara T Tomohiro Matsunaga Y Yoko Sato T Takenori Okada Y Yoshihiro Hirai H Hidetoshi Kawashima A Atsuya Narita K Kazuki Niwa Y Yoshiyuki Sekikawa K Koichi Nishi M Masaru Nishitsuji M Mayuko Tani J Junya Suzuki H Hiroki Nakatsumi T Takashi Ogura H Hideya Kitamura E Eri Hagiwara K Kota Murohashi H Hiroko Okabayashi T Takao Mochimaru S Shigenari Nukaga R Ryosuke Satomi Y Yoshitaka Oyamada N Nobuaki Mori T Tomoya Baba Y Yasutaka Fukui M Mitsuru Odate S Shuko Mashimo Y Yasushi Makino K Kazuma Yagi M Mizuha Hashiguchi J Junko Kagyo T Tetsuya Shiomi S Satoshi Fuke H Hiroshi Saito T Tomoya Tsuchida S Shigeki Fujitani M Mumon Takita D Daiki Morikawa T Toru Yoshida T Takehiro Izumo M Minoru Inomata N Naoyuki Kuse N Nobuyasu Awano M Mari Tone A Akihiro Ito Y Yoshihiko Nakamura K Kota Hoshino J Junichi Maruyama H Hiroyasu Ishikura T Tohru Takata T Toshio Odani M Masaru Amishima T Takeshi Hattori Y Yasuo Shichinohe T Takashi Kagaya T Toshiyuki Kita K Kazuhide Ohta S Satoru Sakagami K Kiyoshi Koshida K Kentaro Hayashi T Tetsuo Shimizu Y Yutaka Kozu H Hisato Hiranuma Y Yasuhiro Gon N Namiki Izumi K Kaoru Nagata K Ken Ueda R Reiko Taki S Satoko Hanada K Kodai Kawamura K Kazuya Ichikado K Kenta Nishiyama H Hiroyuki Muranaka K Kazunori Nakamura N Naozumi Hashimoto K Keiko Wakahara S Sakamoto Koji N Norihito Omote A Akira Ando N Nobuhiro Kodama Y Yasunari Kaneyama S Shunsuke Maeda T Takashige Kuraki T Takemasa Matsumoto K Koutaro Yokote T Taka-Aki Nakada R Ryuzo Abe T Taku Oshima T Tadanaga Shimada M Masahiro Harada T Takeshi Takahashi H Hiroshi Ono T Toshihiro Sakurai T Takayuki Shibusawa Y Yoshifumi Kimizuka A Akihiko Kawana T Tomoya Sano C Chie Watanabe R Ryohei Suematsu H Hisako Sageshima A Ayumi Yoshifuji K Kazuto Ito S Saeko Takahashi K Kota Ishioka M Morio Nakamura M Makoto Masuda A Aya Wakabayashi H Hiroki Watanabe S Suguru Ueda M Masanori Nishikawa Y Yusuke Chihara M Mayumi Takeuchi K Keisuke Onoi J Jun Shinozuka A Atsushi Sueyoshi Y Yoji Nagasaki M Masaki Okamoto S Sayoko Ishihara M Masatoshi Shimo Y Yoshihisa Tokunaga Y Yu Kusaka T Takehiko Ohba S Susumu Isogai S Satoru Fukuyama Y Yoshihiro Eriguchi A Akiko Yonekawa K Keiko Kan-o K Koichiro Matsumoto K Kensuke Kanaoka S Shoichi Ihara K Kiyoshi Komuta Y Yoshiaki Inoue S Shigeru Chiba K Kunihiro Yamagata Y Yuji Hiramatsu H Hirayasu Kai K Koichiro Asano T Tsuyoshi Oguma Y Yoko Ito S Satoru Hashimoto M Masaki Yamasaki Y Yu Kasamatsu Y Yuko Komase N Naoya Hida T Takahiro Tsuburai B Baku Oyama M Minoru Takada H Hidenori Kanda Y Yuichiro Kitagawa T Tetsuya Fukuta T Takahito Miyake S Shozo Yoshida S Shinji Ogura S Shinji Abe Y Yuta Kono Y Yuki Togashi H Hiroyuki Takoi R Ryota Kikuchi S Shinichi Ogawa T Tomouki Ogata S Shoichiro Ishihara A Arihiko Kanehiro S Shinji Ozaki Y Yasuko Fuchimoto S Sae Wada N Nobukazu Fujimoto K Kei Nishiyama M Mariko Terashima S Satoru Beppu K Kosuke Yoshida O Osamu Narumoto H Hideaki Nagai N Nobuharu Ooshima M Mitsuru Motegi A Akira Umeda K Kazuya Miyagawa H Hisato Shimada M Mayu Endo Y Yoshiyuki Ohira M Masafumi Watanabe S Sumito Inoue A Akira Igarashi M Masamichi Sato H Hironori Sagara A Akihiko Tanaka S Shin Ohta T Tomoyuki Kimura Y Yoko Shibata Y Yoshinori Tanino T Takefumi Nikaido H Hiroyuki Minemura Y Yuki Sato Y Yuichiro Yamada T Takuya Hashino M Masato Shinoki H Hajime Iwagoe H Hiroshi Takahashi K Kazuhiko Fujii H Hiroto Kishi M Masayuki Kanai T Tomonori Imamura T Tatsuya Yamashita M Masakiyo Yatomi T Toshitaka Maeno S Shinichi Hayashi M Mai Takahashi M Mizuki Kuramochi I Isamu Kamimaki Y Yoshiteru Tominaga T Tomoo Ishii M Mitsuyoshi Utsugi A Akihiro Ono T Toru Tanaka T Takeru Kashiwada K Kazue Fujita Y Yoshinobu Saito M Masahiro Seike H Hiroko Watanabe H Hiroto Matsuse N Norio Kodaka C Chihiro Nakano T Takeshi Oshio T Takatomo Hirouchi S Shohei Makino M Moritoki Egi A Andreas J. Forstner A Alexander T. Dilthey A Anne-Katrin Pröbstel K Kaan Boztug M Markus M. Nöthen H Ho Namkoong Y Yukinori Okada E Eva C. Beins K Kerstin U. Ludwig

Abstract

Abstract Epstein–Barr virus (EBV) infects approximately 90–95% of the global population 1,2 and persists in B cells as a lifelong infection 3 . Previous EBV infection is associated with autoimmune and neoplastic disease 4 . Still, the biological basis of host control during EBV persistence remains unclear. Here we report the identification of non-genetic and genetic factors that are associated with EBV control during persistent infection. Using blood-based genome sequence data from 486,315 UK Biobank and 336,123 All of Us participants, we identified short-read pairs mapping to the EBV genome in 16.2% and 21.8% of individuals, respectively. EBV read detection (EBVread + ) reflects increased viral load in blood cells, as shown by orthogonal measurements, and was associated with HIV infection, immunosuppressive drug intake and current smoking. Genome-wide analyses of EBVread + identified strong associations at the major histocompatibility complex (MHC), including 54 independent human leukocyte antigen (HLA) alleles of MHC classes I and II, and at 27 genomic regions outside MHC. Epistasis with distinct HLA alleles of MHC class I was observed at the ERAP2 locus. Analysis of individuals with EBV-associated diseases 4 revealed a higher polygenic burden of EBVread + for HLA alleles at MHC class I in multiple sclerosis (driven by HLA-A*02:01) and at MHC class II in rheumatoid arthritis. Phenome-wide analyses identified a polygenic overlap of EBVread + with inflammatory bowel disease, hypothyroidism and type 1 diabetes. Our study establishes by-products of human genome sequencing as a surrogate marker of EBV viral load. This will facilitate investigation and treatment for EBV and other persistent viral infections.

Article Details

Journal Nature
Volume / Issue Vol. 653, Issue 8114
Published May 14, 2026
Pages 444-454
ISSN 0028-0836
Publisher Nature Portfolio

Journal Info

Nature

Nature Portfolio

ISSN: 0028-0836 Health Sciences

Authors (503)

A

Axel Schmidt

T

T. Madhusankha Alawathurage

F

Friederike S. David

Y

Yosuke Ogawa

L

Leonard Frach

S

Sylvia Richter

M

Merle Schaefer

C

Carina M. Mathey

S

Sabrina K. Henne

G

Genta Nagao

H

Hiromu Tanaka

S

Shuhei Azekawa

K

Ko Lee

N

Naoki Fukunaga

J

Junko Hamamoto

H

Hiroki Kabata

K

Katsunori Masaki

H

Hirofumi Kamata

S

Shinnosuke Ikemura

S

Shotaro Chubachi

S

Satoshi Okamori

H

Hideki Terai

A

Atsuho Morita

T

Takanori Asakura

M

Makoto Ishii

K

Koichi Fukunaga

Y

Yoshifumi Uwamino

S

Sho Uchida

S

Shunsuke Uno

T

Tomoyasu Nishimura

N

Naoki Hasegawa

E

Emmy Yanagita

H

Hiroshi Nishihara

J

Junichi Sasaki

H

Hiroshi Morisaki

T

Toshiro Sato

Y

Yuko Kitagawa

Y

Yuta Matsubara

Y

Yohei Mikami

K

Kosaku Nanki

T

Takanori Kanai

R

Ryuya Edahiro

Y

Yuya Shirai

K

Kyuto Sonehara

D

Daisuke Okuzaki

D

Daisuke Motooka

M

Masahiro Kanai

T

Tatsuhiko Naito

K

Kenichi Yamamoto

Q

Qingbo S. Wang

Y

Yasuhiro Kato

T

Takayoshi Morita

S

Shinichi Namba

K

Ken Suzuki

Y

Yoko Naito

Y

Yu-Chen Liu

A

Ayako Takuwa

F

Fuminori Sugihara

J

James B. Wing

S

Shuhei Sakakibara

N

Nobuyuki Hizawa

T

Takayuki Shiroyama

S

Satoru Miyawaki

Y

Yusuke Kawamura

A

Akiyoshi Nakayama

H

Hirotaka Matsuo

Y

Yuichi Maeda

T

Takuro Nii

Y

Yoshimi Noda

T

Takayuki Niitsu

Y

Yuichi Adachi

T

Takatoshi Enomoto

S

Saori Amiya

R

Reina Hara

Y

Yuta Yamaguchi

T

Teruaki Murakami

T

Tomoki Kuge

K

Kinnosuke Matsumoto

Y

Yuji Yamamoto

M

Makoto Yamamoto

M

Midori Yoneda

T

Toshihiro Kishikawa

S

Shuhei Yamada

S

Shuhei Kawabata

N

Noriyuki Kijima

M

Masatoshi Takagaki

N

Noah Sasa

Y

Yuya Ueno

M

Motoyuki Suzuki

N

Norihiko Takemoto

H

Hirotaka Eguchi

T

Takahito Fukusumi

T

Takao Imai

M

Munehisa Fukushima

H

Haruhiko Kishima

H

Hidenori Inohara

K

Kazunori Tomono

K

Kazuto Kato

M

Meiko Takahashi

F

Fumihiko Matsuda

H

Haruhiko Hirata

Y

Yoshito Takeda

A

Atsushi Kumanogoh

T

Takanori Hasegawa

K

Kunihiko Takahashi

T

Tatsuhiko Anzai

S

Satoshi Ito

Y

Yuji Uchimura

A

Akifumi Endo

Y

Yasunari Miyazaki

T

Takayuki Honda

T

Tomoya Tateishi

S

Shuji Tohda

N

Naoya Ichimura

K

Kazunari Sonobe

C

Chihiro Tani Sassa

J

Jun Nakajima

M

Masumi Ai

R

Ryuji Koike

A

Akinori Kimura

S

Satoru Miyano

T

Tomomi Takano

K

Kazuhiko Katayama

K

Koji Okudela

R

Ryunosuke Saiki

Y

Yasuhito Nannya

S

Seishi Ogawa

T

Takayoshi Hyugaji

E

Eigo Shimizu

K

Kotoe Katayama

S

Seiya Imoto

Y

Yosuke Omae

K

Katsushi Tokunaga

T

Takafumi Ueno

Y

Yoshinori Fukui

H

Hiroyuki Hayashi

Y

Yukihiro Yoshimura

N

Natsuo Tachikawa

K

Kazuhisa Takahashi

N

Norihiro Harada

Y

Yuki Tanabe

T

Toshio Naito

M

Makoto Hiki

Y

Yasushi Matsushita

H

Haruhi Takagi

R

Ryousuke Aoki

A

Ai Nakamura

S

Sonoko Harada

H

Hitoshi Sasano

T

Takashi Ishiguro

T

Taisuke Isono

S

Shun Shibata

Y

Yuma Matsui

C

Chiaki Hosoda

K

Kenji Takano

T

Takashi Nishida

Y

Yoichi Kobayashi

Y

Yotaro Takaku

N

Noboru Takayanagi

S

Soichiro Ueda

N

Natsumi Yazaki

A

Ai Tada

M

Masayoshi Miyawaki

M

Masaomi Yamamoto

E

Eriko Yoshida

R

Reina Hayashi

T

Tomoki Nagasaka

S

Sawako Arai

Y

Yutaro Kaneko

K

Kana Sasaki

E

Etsuko Tagaya

M

Masatoshi Kawana

K

Ken Arimura

Y

Yasushi Nakano

Y

Yukiko Nakajima

R

Ryusuke Anan

R

Ryosuke Arai

Y

Yuko Kurihara

Y

Yuko Harada

K

Kazumi Nishio

T

Tetsuya Ueda

M

Masanori Azuma

R

Ryuichi Saito

T

Toshikatsu Sado

Y

Yoshimune Miyazaki

R

Ryuichi Sato

Y

Yuki Haruta

T

Tadao Nagasaki

Y

Yoshinori Yasui

Y

Yoshinori Hasegawa

A

Akihiro Noda

Y

Yusei Fukushima

R

Reina Kitagawa

Y

Yoshikazu Mutoh

T

Tomoki Kimura

T

Tomonori Sato

R

Reoto Takei

S

Satoshi Hagimoto

Y

Yoichiro Noguchi

Y

Yasuhiko Yamano

H

Hajime Sasano

S

Sho Ota

Y

Yasushi Nakamori

K

Kazuhisa Yoshiya

F

Fukuki Saito

T

Tomoyuki Yoshihara

D

Daiki Wada

H

Hiromu Iwamura

S

Syuji Kanayama

S

Shuhei Maruyama

T

Takashi Yoshiyama

K

Ken Ohta

H

Hiroyuki Kokuto

H

Hideo Ogata

Y

Yoshiaki Tanaka

K

Kenichi Arakawa

M

Masafumi Shimoda

T

Takeshi Osawa

H

Hiroki Tateno

I

Isano Hase

S

Shuichi Yoshida

S

Shoji Suzuki

M

Miki Kawada

H

Hirohisa Horinouchi

F

Fumitake Saito

K

Keiko Mitamura

M

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J

Junichi Ochi

T

Tomoyuki Uchida

R

Rie Baba

D

Daisuke Arai

T

Takayuki Ogura

H

Hidenori Takahashi

S

Shigehiro Hagiwara

S

Shunichiro Konishi

I

Ichiro Nakachi

K

Koji Murakami

M

Mitsuhiro Yamada

H

Hisatoshi Sugiura

H

Hirohito Sano

S

Shuichiro Matsumoto

N

Nozomu Kimura

Y

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H

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Y

Yusuke Suzuki

S

Sohei Nakayama

K

Keita Masuzawa

H

Hidefumi Koh

T

Tadashi Manabe

Y

Yohei Funatsu

F

Fumimaro Ito

T

Takahiro Fukui

K

Keisuke Shinozuka

S

Sumiko Kohashi

M

Masatoshi Miyazaki

T

Tomohisa Shoko

T

Takashi Inoue

T

Takahiro Asami

T

Toshiyuki Hirano

K

Keigo Kobayashi

H

Hatsuyo Takaoka

K

Kazuyoshi Watanabe

N

Naoki Miyazawa

Y

Yasuhiro Kimura

R

Reiko Sado

H

Hideyasu Sugimoto

A

Akane Kamiya

N

Naota Kuwahara

A

Akiko Fujiwara

T

Tomohiro Matsunaga

Y

Yoko Sato

T

Takenori Okada

Y

Yoshihiro Hirai

H

Hidetoshi Kawashima

A

Atsuya Narita

K

Kazuki Niwa

Y

Yoshiyuki Sekikawa

K

Koichi Nishi

M

Masaru Nishitsuji

M

Mayuko Tani

J

Junya Suzuki

H

Hiroki Nakatsumi

T

Takashi Ogura

H

Hideya Kitamura

E

Eri Hagiwara

K

Kota Murohashi

H

Hiroko Okabayashi

T

Takao Mochimaru

S

Shigenari Nukaga

R

Ryosuke Satomi

Y

Yoshitaka Oyamada

N

Nobuaki Mori

T

Tomoya Baba

Y

Yasutaka Fukui

M

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S

Shuko Mashimo

Y

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K

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M

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J

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T

Tetsuya Shiomi

S

Satoshi Fuke

H

Hiroshi Saito

T

Tomoya Tsuchida

S

Shigeki Fujitani

M

Mumon Takita

D

Daiki Morikawa

T

Toru Yoshida

T

Takehiro Izumo

M

Minoru Inomata

N

Naoyuki Kuse

N

Nobuyasu Awano

M

Mari Tone

A

Akihiro Ito

Y

Yoshihiko Nakamura

K

Kota Hoshino

J

Junichi Maruyama

H

Hiroyasu Ishikura

T

Tohru Takata

T

Toshio Odani

M

Masaru Amishima

T

Takeshi Hattori

Y

Yasuo Shichinohe

T

Takashi Kagaya

T

Toshiyuki Kita

K

Kazuhide Ohta

S

Satoru Sakagami

K

Kiyoshi Koshida

K

Kentaro Hayashi

T

Tetsuo Shimizu

Y

Yutaka Kozu

H

Hisato Hiranuma

Y

Yasuhiro Gon

N

Namiki Izumi

K

Kaoru Nagata

K

Ken Ueda

R

Reiko Taki

S

Satoko Hanada

K

Kodai Kawamura

K

Kazuya Ichikado

K

Kenta Nishiyama

H

Hiroyuki Muranaka

K

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N

Naozumi Hashimoto

K

Keiko Wakahara

S

Sakamoto Koji

N

Norihito Omote

A

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N

Nobuhiro Kodama

Y

Yasunari Kaneyama

S

Shunsuke Maeda

T

Takashige Kuraki

T

Takemasa Matsumoto

K

Koutaro Yokote

T

Taka-Aki Nakada

R

Ryuzo Abe

T

Taku Oshima

T

Tadanaga Shimada

M

Masahiro Harada

T

Takeshi Takahashi

H

Hiroshi Ono

T

Toshihiro Sakurai

T

Takayuki Shibusawa

Y

Yoshifumi Kimizuka

A

Akihiko Kawana

T

Tomoya Sano

C

Chie Watanabe

R

Ryohei Suematsu

H

Hisako Sageshima

A

Ayumi Yoshifuji

K

Kazuto Ito

S

Saeko Takahashi

K

Kota Ishioka

M

Morio Nakamura

M

Makoto Masuda

A

Aya Wakabayashi

H

Hiroki Watanabe

S

Suguru Ueda

M

Masanori Nishikawa

Y

Yusuke Chihara

M

Mayumi Takeuchi

K

Keisuke Onoi

J

Jun Shinozuka

A

Atsushi Sueyoshi

Y

Yoji Nagasaki

M

Masaki Okamoto

S

Sayoko Ishihara

M

Masatoshi Shimo

Y

Yoshihisa Tokunaga

Y

Yu Kusaka

T

Takehiko Ohba

S

Susumu Isogai

S

Satoru Fukuyama

Y

Yoshihiro Eriguchi

A

Akiko Yonekawa

K

Keiko Kan-o

K

Koichiro Matsumoto

K

Kensuke Kanaoka

S

Shoichi Ihara

K

Kiyoshi Komuta

Y

Yoshiaki Inoue

S

Shigeru Chiba

K

Kunihiro Yamagata

Y

Yuji Hiramatsu

H

Hirayasu Kai

K

Koichiro Asano

T

Tsuyoshi Oguma

Y

Yoko Ito

S

Satoru Hashimoto

M

Masaki Yamasaki

Y

Yu Kasamatsu

Y

Yuko Komase

N

Naoya Hida

T

Takahiro Tsuburai

B

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M

Minoru Takada

H

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Y

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T

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T

Takahito Miyake

S

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S

Shinji Ogura

S

Shinji Abe

Y

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Y

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H

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R

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S

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T

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S

Shoichiro Ishihara

A

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S

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Y

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S

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N

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K

Kei Nishiyama

M

Mariko Terashima

S

Satoru Beppu

K

Kosuke Yoshida

O

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H

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N

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M

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A

Akira Umeda

K

Kazuya Miyagawa

H

Hisato Shimada

M

Mayu Endo

Y

Yoshiyuki Ohira

M

Masafumi Watanabe

S

Sumito Inoue

A

Akira Igarashi

M

Masamichi Sato

H

Hironori Sagara

A

Akihiko Tanaka

S

Shin Ohta

T

Tomoyuki Kimura

Y

Yoko Shibata

Y

Yoshinori Tanino

T

Takefumi Nikaido

H

Hiroyuki Minemura

Y

Yuki Sato

Y

Yuichiro Yamada

T

Takuya Hashino

M

Masato Shinoki

H

Hajime Iwagoe

H

Hiroshi Takahashi

K

Kazuhiko Fujii

H

Hiroto Kishi

M

Masayuki Kanai

T

Tomonori Imamura

T

Tatsuya Yamashita

M

Masakiyo Yatomi

T

Toshitaka Maeno

S

Shinichi Hayashi

M

Mai Takahashi

M

Mizuki Kuramochi

I

Isamu Kamimaki

Y

Yoshiteru Tominaga

T

Tomoo Ishii

M

Mitsuyoshi Utsugi

A

Akihiro Ono

T

Toru Tanaka

T

Takeru Kashiwada

K

Kazue Fujita

Y

Yoshinobu Saito

M

Masahiro Seike

H

Hiroko Watanabe

H

Hiroto Matsuse

N

Norio Kodaka

C

Chihiro Nakano

T

Takeshi Oshio

T

Takatomo Hirouchi

S

Shohei Makino

M

Moritoki Egi

A

Andreas J. Forstner

A

Alexander T. Dilthey

A

Anne-Katrin Pröbstel

K

Kaan Boztug

M

Markus M. Nöthen

H

Ho Namkoong

Y

Yukinori Okada

E

Eva C. Beins

K

Kerstin U. Ludwig