Research ArticleDevelopmentNeuroscience Open Access | 10.1172/JCI176631

Minkyung Kang,1,2 Jihye Choi,3 Jeongho Han,4 Toshiyuki Araki,5 Soo-Whee Kim,6,7 Hyun-Hee Ryu,1 Min-Gyun Kim,1,2 Seoyeon Kim,6,7 Hanbyul Jang,1,2 Sun Yong Kim,1,2 Kyoung-Doo Hwang,1,2 Soobin Kim,1,2 Myeongjong Yoo,1,2 Jaegeon Lee,1,2 Kitae Kim,1,2 Pojeong Park,8 Ja Eun Choi,8 Dae Hee Han,8 Yujin Kim,6,7 Jeongyeon Kim,9 Sunghoe Chang,1,2 Bong-Kiun Kaang,8 Jung Min Ko,10,11 Keun-Ah Cheon,12,13 Joon-Yong An,6 Sang Jeong Kim,1,2,14 Hyungju Park,4 Benjamin G. Neel,5 Chul Hoon Kim,3 and Yong-Seok Lee1,2,14

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kang, M. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Choi, J. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Han, J. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Araki, T. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, S. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Ryu, H. in: JCI | PubMed | Google Scholar |

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, M. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, S. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Jang, H. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, S. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Hwang, K. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, S. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Yoo, M. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Lee, J. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, K. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Park, P. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Choi, J. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Han, D. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Republic of Korea.

3Department of Pharmacology, Graduate School of Medical Science, Brain Korea 21 Project, Yonsei University College of Medicine, Seoul, Republic of Korea.

4Research Group of Neurovascular Unit, Korea Brain Research Institute, Daegu, Republic of Korea.

5Laura and Isaac Perlmutter Cancer Center, New York University Langone Medical Center, New York, New York, USA.

6Department of Integrated Biomedical and Life Science, and

7BK21FOUR R&E Center for Learning Health Systems, Korea University, Seoul, Republic of Korea.

8School of Biological Sciences, Seoul National University, Seoul, Republic of Korea.

9Emotion, Cognition and Behavior Research Group, Korea Brain Research Institute, Daegu, Republic of Korea.

10Department of Pediatrics, Division of Clinical Genetics, Seoul National University College of Medicine, Seoul, Republic of Korea.

11Rare Disease Center, Seoul National University Hospital, Seoul, Republic of Korea.

12Department of Child and Adolescent Psychiatry, Severance Hospital, and

13Institute of Behavioral Science in Medicine, Yonsei University College of Medicine, Yonsei University Health System, Seoul, Republic of Korea.

14Neuroscience Research Institute, Medical Research Center, Seoul National University, Seoul, Republic of Korea.

Address correspondence to: Chul Hoon Kim, 50-1 Yonsei-ro, Department of Pharmacology, Yonsei University College of Medicine, Seoul 03722, Republic of Korea. Phone: 82.2.2228.1738; Email: KIMHOON@yuhs.ac. Or to: Yong-Seok Lee, 103 Daehakro, Jongro-gu, Department of Physiology, Seoul National University College of Medicine, Seoul 03080, Republic of Korea. Phone: 82.2.740.8225; Email: yongseok7@snu.ac.kr.

Authorship note: MK and JC are co–first authors and contributed equally to this work.

Find articles by Kim, Y. in: JCI | PubMed | Google Scholar

1Department of Physiology, and

2Department