• Christopher J. MacDonald, Susumu Tonegawa. Crucial role for CA2 inputs in the sequential organization of CA1 time cells supporting memory. Proceedings of the National Academy of Sciences (2021), 118 (3) e2020698118; DOI: 10.1073/pnas.2020698118


  • Bari, A., Xu, S., Pignatelli, M., Takeuchi, D., Feng, J., Li, Y., Tonegawa, S. Differential attentional control mechanisms by two distinct noradrenergic coeruleo-frontal cortical pathways. Proceedings of the National Academy of Sciences (2020); 202015635; DOI: 10.1073/pnas.2015635117
  • Lak, A., Hueske, E., Hirokawa, J., Masset, P., Ott, T., Urai, A.E., Donner, T.H., Carandini, M., Tonegawa, S., Uchida, N., Kepecs, A. Reinforcement biases subsequent perceptual decisions when confidence is low, a widespread behavioral phenomenon. eLife (2020);9:e49834. DOI:10.7554/eLife.49834
  • Sun, C., Yang, W., Martin, J., Tonegawa, S. Hippocampal neurons represent events as transferable units of experience. Nat Neurosci (2020). https://doi.org/10.1038/s41593-020-0614-x
  • Josselyn, S. and Tonegawa, S. Memory engrams: recalling the past and imagining the future. Science 367: eaaw4325 (2020).


  • Zhang, X., Kim, J., Tonegawa, S. Amygdala Reward Neurons Form and Store Fear Extinction Memory, Neuron (2019), https://doi.org/10.1016/j.neuron.2019.12.025
  • Pignatelli, M., Ryan, T., Roy. D., Lovett, C., Smith, L., Muralidhar, S., Tonegawa, S. Engram excitability state determines the efficacy of memory retrieval. Neuron 10 (2):274-284, (2019)


  • Tonegawa, S., Morrissey, M.D., Kitamura, T. The role of engram cells in the systems consolidation of memory. Nature Reviews Neuroscience. (2018).


  • Wagatsuma, A., Okuyama, T., Sun, C., Smith, L.M., Abe, K., Tonegawa, S. Locus coeruleus input to hippocampal CA3 drives single-trial learning of a novel context. PNAS. 115: 310-316 (2018). In the news »
  • Roy, D.S., Muralidhar, S., Smith, L.M., Tonegawa, S. Silent memory engrams as the basis for retrograde amnesia. PNAS. 114: 9972-9979 (2017). In the news »
  • Xu, S., Das, G., Hueske, E., Tonegawa, S. Dorsal Raphe Serotonergic Neurons Control Intertemporal Choice under Trade-Off. Current Biology. 27: 3111-3119 (2017).
  • Yamamoto, J., Tonegawa, S. Direct Medial Entorhinal Cortex Input to Hippocampal CA1 Is Crucial for Extended Quiet Awake Replay. Neuron. 96: 217-227 (2017).
  • Roy, D.S.*, Kitamura, T.*, Okuyama, T., Ogawa, S.K., Sun, C., Obata, Y., Yoshiki, A., Tonegawa, S. Distinct Neural Circuits for the Formation and Retrieval of Episodic Memories. Cell. 170: 1000-1012 (2017). In the news »
  • Kitamura, T.*, Ogawa, S.K.*, Roy, D.S.*, Okuyama, T., Morrissey, M.D., Smith, L., Redondo, R.L., Tonegawa, S. Engrams and circuits crucial for systems consolidation of a memory. Science. 356: 73-78 (2017). View PDF »  In the news »
  • Kim, J.*, Zhang, X.*, Muralidhar, S., LeBlanc, S.A., Tonegawa, S. Basolateral to Central Amygdala Neural Circuits for Appetitive Behaviors. Neuron. 93(6): 1464-1479 (2017).  View PDF »  In the news »


  • Kim, J., Pignatelli, M., Xu, S., Itohara, S., Tonegawa, S. Antagonistic negative and positive neurons of the basolateral amygdala. Nature Neuroscience. 19: 1636-1646 (2016).  View PDF » In the news »
  • Okuyama, T., Kitamura, T., Roy, D.S., Itohara, S., Tonegawa, S. Ventral CA1 neurons store social memory. Science. 353: 1536-1541 (2016). View PDF » In the news »
  • Roy, D.S., Arons, A., Mitchell, T.I., Pignatelli, M., Ryan, T.J., Tonegawa, S. Memory retrieval by activating engram cells in mouse models of early Alzheimer’s disease. Nature. 531: 1-5 (2016). View PDF » In the news »


  • Kitamura, T., Sun, C., Martin, J., Kitch, L.J., Schnitzer, M., and Tonegawa, S. Entorhinal Cortical Ocean Cells Encode Specific Contexts and Drive Context-Specific Fear Memory. Neuron. 87(6): 1317-1331. View PDF » In the news »
  • Tonegawa, S., Liu, X., Ramirez, S., Redondo, R. Memory Engram Cells Have Come of Age. Neuron. 87(5): 918-931. View PDF »
  • Kitamura, T., MacDonald, C.J., and Tonegawa, S. Entorhinal-hippocampal neuronal circuits bridge temporally discontiguous events. Learning & Memory. 22: 438-443 (2015). View PDF »
  • Sarinana, J. and Tonegawa, S. Differentiation of Forebrain and Hippocampal Dopamine 1-Class
    Receptors, D1R and D5R, in Spatial Learning and Memory. Hippocampus. DOI: 10.1002/hipo.22492. View PDF »
  • Tonegawa, S., Pignatelli, M., Roy, D.S., and Ryan, T.J. Memory engram storage and retrieval. Current Opinion in Neurobiology. 35: 101-109 (2015). View PDF »
  • Sun, C., Kitamura, T., Yamamoto, J., Martin, J., Pignatelli, M., Kitch, L.J., Schnitzer, M.J., Tonegawa, S. Distinct speed dependence of entorhinal island and ocean cells, including respective grid cells. PNAS 112(30): 9466-9471 (2015). View PDF »
  • Ramirez, S., Liu, X., MacDonald C.J., Moffa, A., Zhou, J., Redondo, R.L., Tonegawa, S. Activating positive memory engrams suppresses depression-like behaviour. Nature 522: 335-339 (2015). View PDF »  In the news »
  • Ryan, T.J., Roy, D.S., Pignatelli, M., Arons, A., Tonegawa, S. Engram cells retain memory under retrograde amnesia. Science 348: 941-1052 (2015). View PDF »  In the news »
  • Liu, X., Ramirez, S., Redondo R., Tonegawa S. Identification and Manipulation of Memory Engram Cells. Cold Spring Harbor Symposia on Quantitative Biology Volume LXXIX (2015). View PDF »


  • Redondo R., Kim J., Arons A., Ramirez S., Liu X., Tonegawa S. Bidirectional switch of the valence associated with a hippocampal contextual memory engram. Nature 513: 426-430 (2014). View PDF » In the news »
  • Sariñana, J., Kitamura, K., Künzler, P., Sultzman, L., Tonegawa S. Differential roles of the dopamine 1-class receptors, D1R and D5R, in hippocampal dependent memory. PNAS doi/10.1073/pnas.1407395111 (2014). View PDF »
  • Yamamoto, J., Suh, J., Takeuchi, D., Tonegawa, S. Successful Execution of Working Memory Linked to Synchronized High-Frequency Gamma Oscillations. Cell. 157: 1-13 (2014). View PDF »  In the news »
  • Kitamura, T., Pignatelli, M., Suh, J., Kohara, K., Yoshiki, A., Abe, K., Tonegawa, S. Island Cells Control Temporal Association Memory. Science. 343: 896-901 (2014). View PDF »  In the news »
  • Kohara K., Pignatelli, M., Rivest, A.J., Jung, H.Y., Kitamura, T., Suh, J., Frank, D., Kajikawa, K., Mise, N., Obata, Y., Wickersham, I., Tonegawa, S. Cell type–specific genetic and optogenetic tools reveal hippocampal CA2 circuits. Nat. Neuro. 17: 269-279 (2014). View PDF »  In the news »
  • Dragoi, G. and Tonegawa S. Selection of preconfigured cell assemblies for representation of novel spatial experiences. Phil. Trans. R. Soc. B. 369: 20120522 (2014). View PDF »
  • Ramirez, S., Tonegawa, S., Liu, X. Identification and optogenetic manipulation of memory engrams in the hippocampus. Front. Behav. Neurosci. 7: 226 (2014).. View PDF »
  • Liu, X., Ramirez, S., and Tonegawa, S. Inception of a false memory by optogenetic manipulation of a hippocampal memory engram. Phil. Trans. R. Soc. B. 369: 20130142 (2014). View PDF »


  • Dragoi, G. and Tonegawa S. Development of schemas revealed by prior experience and NMDA receptor knock-out. eLife 2:e01326(2013). View PDF »
  • Suh, J., Foster, D., Davoudi, H., Wilson, M., Tonegawa, S. Impaired Hippocampal Ripple-Associated Replay in a Mouse Model of Schizophrenia. Neuron. 80: 484–493, (2013). View PDF »  In the news »
  • Ramirez, S., Liu, X., Lin, P.A., Suh, J., Pignatelli, M., Redondo, R., Ryan, T.J., Tonegawa, S. Creating a False Memory in the Hippocampus. Science. 341: 387-391 (2013). View PDF 1 » View PDF 2 »  In the news »
  • Arsenault, D., Dal-Pan, A., Tremblay, C., Bennett, D. A., Guitton, M. J., De Koninck, Y., Tonegawa, S., & Calon, F. PAK inactivation impairs social recognition in 3xTg-AD mice without increasing brain deposition of tau and Aβ. J Neurosci. 33(26):10729-10740(2013). View PDF »
  • Dragoi, D. and Tonegawa, S. Distinct preplay of multiple novel spatial experiences in the rat. PNAS. 110 (22): 9100-9105 (2013). View PDF »
  • Dolan, B.M., Duron, S.G., Campbell, D.A., Vollrath, B., Rao, B.S.S., Ko, H.Y., Lin, G.G., Govindarajan, A., Choi, S.Y., Tonegawa, S. Rescue of fragile X syndrome phenotypes in Fmr1 KO mice by the small-molecule PAK inhibitor FRAX486. PNAS. 110 (14):5671-6(2013). View PDF »  In the news »


  • Liu X., Ramirez, S., Pang, P., Puryear, C., Govindarajan, A., Deisseroth, K., and Tonegawa S. Optogenetic stimulation of a hippocampal engram activates fear memory recall. Nature 484: 381–385 (2012). View PDF »  In the news »
  • Place R., Lykken C., Beer Z., Suh J., McHugh T.J., Tonegawa S., Eichenbaum H., Sauvage M.M. Nmda Signaling in Ca1 Mediates Selectively the Spatial Component of Episodic Memory. Learn. Mem. 19(4): 164-69 (2012). View PDF »
  • Nakashiba, T., Cushman, J., Pelkey, K., Renaudineau, S., Buhl, D., McHugh, T.J., Rodriguez Barrera, V., Chittajallu, R., Iwamoto, K.S., McBain, C.J., Fanselow, M.S. and Tonegawa, S. Young Dentate Granule Cells Mediate Pattern Separation whereas Old Granule Cells Facilitate Pattern Completion. Cell 149 (1): 188-201 (2012). View PDF 1 »  View PDF 2 »


  • Suh, J., Rivest, A.J., Nakashiba, T., Tominaga, T., Tonegawa, S. Entorhinal cortex layer III input to the hippocampus is crucial for temporal association memory. Science 334: 1415-2 (2011). View PDF 1 »  View PDF 2 »
  • Govindarajan, A., Israely, I., Huang, S.Y., and Tonegawa, S. The Dendritic Branch Is the Preferred Integrative Unit for Protein Synthesis-Dependent LTP. Neuron 69: 132-146 (2011). View PDF 1 »  View PDF 2 »
  • Dragoi, G. and Tonegawa, S. Preplay of future place cell sequences by hippocampal cellular assemblies. Nature 469: 397-403 (2011). View PDF 1 »  View PDF 2 »


  • Torborg, C.L., Nakashiba, T., Tonegawa, S., and McBain, C.J. Control of CA3 Output by Feedforward Inhibition Despite Developmental Changes in the Excitation-Inhibition Balance. J. Neurosci. 30(46): 15628–15637 (2010). View PDF »
  • Wiltgen, B.J., Royle, G.A., Gray, E.E., Abdipranoto, A., Thangthaeng, N., Jacobs, N., Saab, F., Tonegawa, S., Heinemann, S.F., O’Dell, T.J., Fanselow, M.S., and Vissel, B. A Role for Calcium-Permeable AMPA Receptors in Synaptic Plasticity and Learning. PLoS One 5(9): e12818 (2010). View PDF »
  • Liu, X. and Tonegawa, S. Optogenetics 3.0 (Preview). Cell 141: 154-165 (2010). View PDF »
  • Slutsky, I., Abumaria, N., Wu, L., Huang, C., Zhang, L., Li, B., Zhao, X., Govindarajan, A., Zhao, M., Zhuo, M., Tonegawa, S., Liu, G. Enhancement of Learning and Memory by Elevating Brain Magnesium. Neuron 65: 165-177 (2010). View PDF »
  • Kamsler, A., McHugh, T.J., Gerber, D., Huang, S.Y., and Tonegawa, S. Presynaptic m1 muscarinic receptors are necessary for mGluR LTD in the hippocampus. PNAS 107: 1618-1623 (2010). View PDF »


  • Jinde, S., Belforte, J.E., Yamamoto, J., Wilson, M.A., Tonegawa, S., and Nakazawa, K.Lack of kainic acid-induced gamma oscillations predicts subsequent CA1 excitotoxic cell death. Eur. J. Neurosci. 30: 1036–1055 (2009). View PDF »
  • McHugh, T.J., and Tonegawa, S. CA3 NMDA Receptors Are Required for the Rapid Formation of a Salient Contextual Representation. Hippocampus 19:1153–1158 (2009). View PDF »
  • Nakashiba, T., Buhl, D.L., McHugh, T.J., and Tonegawa, S. Hippocampal CA3 Output Is Crucial for Ripple-Associated Reactivation and Consolidation of Memory. Neuron 62: 781-787 (2009). View PDF 1 »  View PDF 2 »


  • Nakashiba, T., Young, J.Z., McHugh, T.J., Buhl, D.L., and Tonegawa, S. Transgenic inhibition of synaptic transmission reveals role of CA3 output in hippocampal learning. Science 319: 1260-1264 (2008). View PDF 1 »  View PDF 1 »
  • Tonegawa, S. and McHughoh, T.J. The ins and outs of hippocampal circuits. Neuron 57: 175-177 (2008). View PDF »
  • Leamey, C.A., Glendining, K.A., Kreiman, G., Kang, N.D., Wang, K.H., Fassler, R., Sawatari, A., Tonegawa, S., and Sur, M. Differential gene expression between sensory neocortical areas: potential roles for Ten_m3 and Bcl6 in patterning visual and somatosensory pathways. Cerebral Cortex 18(1): 53-66 (2008). View PDF »
  • McHugh, T.J. and Tonegawa, S. Molecular and Circuit Mechanisms for Hippocampal Learning. Fondation IPSEN, Research and Perspectives in Neurosciences 13-21 (2008). View PDF »
  • Gharami, K., Xie, Y., An, J.J., Tonegawa, S., and Xu, B. Brain-derived neurotrophic factor over-expression in the forebrain ameliorates Huntington’s disease phenotypes in mice. J. Neurochem. 105(2): 369-379 (2008). View PDF »


  • Caine, S.B., Thomsen, M., Gabriel, K.I., Berkowitz, J.S., Gold, L.H., Koob, G.F., Tonegawa, S., Zhang, J., and Xu, M. Lack of self-administration of cocaine in dopamine D1 receptor knock-out mice. J. Neurosci. 27(48):13140-50 (2007). View PDF »
  • Kent, K., Hess, K. Tonegawa, S., and Small, S.A. CA3 NMDA receptors are required for experience-dependent shifts in hippocampal activity. Hippocampus 1098-1063 (2007). View PDF »
  • Hayashi, M.L.,Shankaranarayana Rao, B.S., Seo, J.S., Choi, H.S., Dolan, B.M., Choi, S.Y., Chattarji, S., and Tonegawa, S. Inhibition of p21-activated kinase rescues symptoms of fragile X syndrome in mice. PNAS 104:11489-11494 (2007). View PDF »
  • McHugh, T.J., Jones, M.W., Quinn, J.J., Balthasar, N., Coppari, R., Elmquist, J.K., Lowell, B.B., Fanselow, M.S., Wilson, M.A., and Tonegawa, S. Dentate Gyrus NMDA Receptors Mediate Rapid Pattern Separation in the Hippocampal Network. Science 317:94-99 (2007). View PDF 1 » View PDF 2 »
  • Yamada, K., Gerger, D.J., Iwayama, Y., Ohnishi, T., Ohba, H., Toyta, T., Arga, J., Minabe, Y., Tonegawa, S., and Yoshikawa, T. Genetic analysis of the calcineurin pathway identifies the EGR family and specifically EGR3, as potential candidates in schizophrenia. PNAS 104:2815-2820 (2007). View PDF »
  • McHugh, T.J. and Tonegawa, S. Spatial exploration is required for the formation of contextual fear memory. Behav. Neurosci. 121(2):335-339 (2007). View PDF »


  • Rondi-Reig, L., Petit, G.H., Tobin, C., Tonegawa, S., Mariana, J., and Berthoz, A. Impaired sequential egocentric and allocentric memories in forebrain-specific-NMDA receptor knock0oug mice during a new task dissociating strategies of navigation. J. Neurosci. 26:4071-4081 (2006). View PDF »
  • Govindarajan, A., Shankarananarayana Rao, B.S., Nair, D., Trinh, M., Mawjee, N., Tonegawa, S., and Chattarji, S. Transgenic brain-derived neuroptrophic factor expression causes both anxiogenic and antidepressant effects. PNAS 103:13208-13213 (2006). View PDF »
  • Wang, K.H., Majewska, A., Schummers, J., Farley, B., Hu, C., Sur, M., and Tonegawa, S. In vivo two-photon imaging reveals a role of Arc in enhancing orientation specificity in visual cortex. Cell 126:389-402 (2006). View PDF »
  • Govindarajan, A., Kelleher, R.J., and Tonegawa, S. A clustered plasticity model of long-term memory engrams. Nature Rev. Neurosci. 7:575-583 (2006). View PDF »
  • Kishimoto, Y., Nakazawa, K., Tonegawa, S., Kirino, Y., and Kano, M. Hippocampal CA3 NMDA receptors are crucial for adaptive timing of trace eyeblink conditioned response. J. Neurosci. 26:1562-1570 (2006). View PDF »
  • Kremin, T., Gerber, D., Giocomo, L.M., Huang, S.Y., Tonegawa, S., and Hasselmo, M.E. Muscarinic suppression in stratum radiatum of CA1 shows dependence on presynaptic M1 receptors and is not dependent on effects at GABA(B) receptors. Neurobiol. Learn. Mem. 85:153-163 (2006). View PDF »


  • Chan, G.C., Tonegawa, S., and Storm, D.R. Hippocampal neurons express a calcineuring-actrivated adenylyl cyclase. J. Neurosci. 25:9913-9918 (2005). View PDF »
  • Anderson, M.P., Mochizuki, T., Xie, J., Fischler, W., Manager, J.P., Talley, E.M., Scammell, T.E., and Tonegawa, S. Thalamic Cav3.1 T-type Ca2+ channel plays a crucial role in stabilizing sleep. PNAS 102:1743-1748 (2005). View PDF »


  • Newton, J.R., Ellswirth, C., Miyakawa, T. Tonegawa, S., and Sur, M. Acceleration of visually cued conditioned fear through the auditory pathway. Nat. Neurosci. 7:968-973 (2004). View PDF »
  • Kelleher, R.J.III, Govindarajan, A., and Tonegawa, S. Translational regulatory mechanisms in persistent forms of synaptic plasticity. Neuron 44:59-73 (2004). View PDF »
  • Hayashi, M.L., Choi, S.-Y., Rao, B.S.S., Jung, H.-Y., Lee, H.-K., Zhang, D., Chattarji, S., Kirkwood, A., and Tonegawa, S. Altered cortical synaptic morphology and impaired memory consolidation in forebrain-specific dominant negative p21-activated kinase (PAK) transgenic mice. Neuron 42:773-787 (2004). View PDF »
  • Nakazawa, K., McHugh, T.J., Wilson, M.A., and Tonegawa, S. NMDA receptors, place cells and hippocampal spatial memory. Nature Rev. Neurosci. 5:361-372 (2004). View PDF »
  • Gerber, D.J., and Tonegawa, S. Psychotomimetic effects of drugs—a common pathway to schizophrenia? N. Engl. J. Med. 350:1047-1048 (2004). View PDF »
  • Kelleher, R.J., Govindarajan, A., Jung, H.-Y., Kang, H., and Tonegawa, S. Translational control by MAPK signaling in long-term synaptic plasticity and memory. Cell 116:467-479 (2004). View PDF »
  • Tonegawa, S. That great time in Basel. Cell 116:S99-101 (2004). View PDF »


  • Shutoh, F., Katoh, A., Ohki, M., Itohara, S., Tonegawa, S., and Nagao, S. Role of protein kinase C family in the cerebellum-dependent adaptive learning of horizontal optokinetic response eye movements in mice. Eur. J. Neurosci. 18:134-142 (2003). View PDF »
  • Gianfranceschi, L., Siciliano, R., Walls, J., Morales, B., Kirkwood, A., Huang, Z.J., Tonegawa, S., and Maffei, L. BDNF overexpression rescues the effects of dark rearing on the development of critical period in visual cortex. PNAS 100:12486-12491(2003). View PDF »
  • Sawtell, N.B., Frenkel, M.Y., Philpot, B.D., Nakazawa, K., Tonegawa, S., and Bear, M.F. NMRA Receptor-Dependent Ocular Dominance Plasticity on Adult Visual Cortex. Neuron 38:977-985 (2003). View PDF »
  • Gerber, D.J., Hall, D., Miyakawa, T., Demars, S., Gogos, J.A., Karayiorgou, M., and Tonegawa, S. Evidence for association of schizophrenia with genetic variation in the 8p21.3 gene, PPP3CC, encoding the calcineurin gamma subunit. PNAS 100:8993-8998 (2003). View PDF »
  • Miyakawa, T., Leiter, L.M., Gerber, D.J., Gainetdinov, R.R., Sotnikova, T.D., Zeng, H., Caron, M.G., and Tonegawa, S. Conditional calcineurin knockout mice exhibit multiple abnormal behaviors related to schizophrenia. PNAS 100:8987-8992 (2003). View PDF »
  • Fukaya, M., Kato, A., Lovett, C., Tonegawa, S., and Watanabe, M. Retention of NMDA receptor NR2 subunits in the lumen of endoplasmic reticulum in targeted NR1 knockout mice. PNAS 100:4855-4860 (2003). View PDF »
  • Tonegawa, S., Nakazawa, K., and Wilson, M.A. Genetic neuroscience of mammalian learning and memory. Phil. Trans. R. Soc. London 358:787-795 (2003). View PDF »
  • Hinds, H.L., Guossakov, I., Nakazawa, K., Tonegawa, S., and Bolshakov, V.Y. Essential function of α-CaMKII in neurotransmitter release at a glutamatergic central synapse. PNAS 100:4275-4280 (2003). View PDF »
  • Nakazawa, K., Sun, L.D., Quirk, M.C., Rondi-Reig, L., Wilson, M.A., and Tonegawa, S. Hippocampal CA3 NMDA receptors are crucial for memory acquisition of one-time experience. Neuron 38:305-315 (2003). View PDF »


  • Nakazawa, K., Quirk, M.C., Chitwood, R.A., Watanabe, M., Yeckel, M.F., Sun, L.D., Kato, A., Carr, C.A., Johnston, D., Wilson, M.A., and Tonegawa, S. Requirement for hippocampal CA3 NMDA receptors in associative memory recall. Science 297:211-218 (2002). View PDF »
  • Huang, Z.J., Yu, W., Lovett, C. and Tonegawa, S. Cre/loxP recombination-activated neuronal markers in mouse neocortex and hippocampus. Genesis 32:209-217 (2002). View PDF »


  • Gerber, D.J., Sotnikova, T.D., Gainetdinov, R.R., Huang, S.Y., Caron, M.G., and Tonegawa, S. Hyperactivity, elevated dopaminergic transmission, and response to amphetamine in M1 muscarinic acetylcholine receptor-deficient mice. PNAS 98:15312-15317 (2001). View PDF »
  • Zeng, H., Chattarji, S., Barbarosie, M., Rondi-Reig, L., Philpot, B.D., Miyakawa, T., Bear, M.F., and Tonegawa, S. Forebrain-specific calcineurin knockout selectively impairs bidirectional synaptic plasticity and working/episodic-like memory. Cell 107:617-629 (2001). View PDF »
  • Kang, H., Sun, L.D., Atkins, C.M., Soderling, T.R., Wilson, M.A., and Tonegawa, S. An important role of neural activity-dependent CaMKIV signaling in the consolidation of long-term memory. Cell 106:771-783 (2001). View PDF »
  • Rondi-Reig, L., Libbey, M., Eichenbaum, H., and Tonegawa, S. CA1-specific N-methyl-D-aspartate receptor knockout mice are deficient in solving a nonspatial transverse patterning task. PNAS 98:3543-3548 (2001). View PDF »


  • Iwasato, T., Datwani, A., Wolf, A.M., Nishiyama, H., Taguchi, Y., Tonegawa, S., Knopfel, T., Erzurumlu, R.S., and Itohara, S. Cortex-restricted disruption of NMAR1 impairs neuronal patterns in the barrel cortex. Nature 406:726-731 (2000). View PDF »
  • Wang, Y., Xu, R., Sasaoka, T., Tonegawa, S., Kung, M.P., and Sankoorika, E.B. Dopamine D2 long receptor-deficient mice display alterations in striatum-dependent functions. J. Neurosci. 20:8305-8314 (2000). View PDF »
  • Huerta, P.T., Sun, L.D., Wilson, M.A., Tonegawa, S. Formation of temporal memory requires NMDA receptors within CA1 pyramidal neurons. Neuron 25:473-480 (2000). View PDF »
  • Huerta, P.T. and Tonegawa, S. NMDA receptors in CA1 pyramidal cells are needed for spatial learning, but not for nonspatial learning. Taniguchi Symposia on Brain Sciences, Challenges for Neuroscience in the 21st Century. 22:161-169 (2000). View PDF »


  • Hanover, J.L., Huang, J.Z., Tonegawa, S., and Stryker, M.P. Brain-derived Neurotrophic Factor Overexpression Induces Precocious Critical Period in Mouse Visual Cortex. J. of Neurosci. 19 RC40:1-5 (1999). View PDF »
  • Berezovska, O., Frosch, M., McLean P., Knowles R., Koo, E., Kang, D., Shen J., Lu. F.M., Lux, S.E., and Tonegawa S. The Alzheimer-related gene presenilin 1 facilitates notch 1 in primary mammalian neurons. Brain Res. Mol. Brain 69:273-80 (1999). View PDF »
  • Oliveira-dos-Santos A.J., Ho A., Tada, Y., Lafaille, J.J., Tonegawa, S., Mak. T.M., and Penninger, J.M. CD28 costimulation is crucial for the development of spontaneous autoimmune encephalomyelitis. Journal of Immunology 162:4490-5 (1999). View PDF »
  • Bowers BJ., Owen, E.H., Collins, A.C., Abeliovich, A., Tonegawa, S., Wehner, J.M. Decreased ethanol sensitivity and tolerance development in gamma-protein kinase C null mutant mice is dependent on genetic background. Alcohol Clinical Experience Research 23:387-97 (1999). View PDF »
  • Huang, J.Z., Kirkwood, A., Pizzorusso, T., Porciatti, V., Morales, B., Bear, M. F., Maffei, L., and Tonegawa, S. BDNF Regulates the maturation of inhibition and the critical period of plasticity in mouse visual cortex. Cell 98:739-755 (1999). View PDF »
  • Levelt, C.N., DeJong, Y.P., Muzoguchi, E., O’Farrelly, C., Bhan, A.K., Tonegawa, S., Terhorst, C., and Simpson, S.J. High- and low-affinity single-peptide/MHC ligands have distinct effects on the development of mucosal CD8aa and CD8ab T lymphocytes. PNAS 96:5628-5633 (1999). View PDF »
  • Fujihashi, K., Dohi, T., Kweon, M.N., McGhee, J.R., Koga, T., Cooper, M.D., Tonegawa, S., and Kiyono, H. gd T cells regulate mucosally induced tolerance in a dose-dependent fashion. Int. Immunol. 11:1907-1916 (1999). View PDF »
  • Xu, M., Koeltzow, T.E., Cooper, D.C., Tonegawa, S., and White, F.J. Dopamine D3 receptor mutant and wild-type mice exhibit identical responses to putative D3 receptor-selective agonists and antagonists. Synapse 31:210-215 (1999). View PDF »


  • Levelt, C.N., Mizoguchi, E., Zacks, R., Bhan, A.K., and Tonegawa, S. Inhibition of intrathymic T cell development by expression of a transgenic antagonist peptide. PNAS 95:14349-14364 (1998). View PDF »
  • Van de Keere, F., and Tonegawa, S. CD4+ T cells prevent spontaneous experimental autoimmune encephalomyelitis in anti-myelin basic protein T cell receptor transgenic mice. J. Exp. Med. 188:1875-1882 (1998). View PDF »
  • Hinds, H.L., Tonegawa, S., and Malinow, R. CA1 long term potentiation is diminished but present in hippocampal slices from a-CaMKII mutant mice. Learning & Memory 5:344-354 (1998). View PDF »
  • Chen, D.F. and Tonegawa, S. Why do mature CNS neurons of mammals fail to reestablish connections following injury – Functions of BCL-2. Cell Death and Differentiation 5:816-822 (1998). View PDF »
  • Koeltzow, T.E., Cooper, D.C., Hu, X.-T., Xu, M., Tonegawa, S., Wolf, M.E., and White, F.J. Assessment of dopamine D3 receptor function using D3 receptor knockout mice. In: Dopamine Receptor Subtypes from Basic Science to Clinic (Jenner, P., and Demirdamar, R., Eds.) Amsterdam: ISO Press, 131-146 (1998). View PDF »
  • Koeltzow, T.E., Xu, M., Cooper, D.C., Hu, X.-T., Tonegawa, S., Wolf, M.E., and White, F.J. Alterations in dopamine release but not dopamine autoreceptor function in dopamine D3 receptor mutant mice. J. Neurosci. 18:2231-2238 (1998). View PDF »
  • Delaney, J.R., Sykulev, Y., Eisen, H., and Tonegawa, S. Differences in the level of expression of class 1 MHC proteins on thymic epithelial and dendritic cells influence the decision of immature thymocytes between positive and negative selections. PNAS 95:5235-5240 (1998). View PDF »
  • Burnett, A.L., Johns, D.G., Kriegsfeld, L.J., Klein, S.L., Calvin, D.C., Demas, G.E., Schramm, L.P., Tonegawa, S., Jenlson, R.J., Snyder, S.H., and Poss, K.D. Ejaculatory abnormalities in mice with targeted disruption of the gene for heme oxygenase-2. Nature Med. 4:84-87 (1998). View PDF »


  • Iwasato, T., Erzurumlu, R.S., Huerta, P.T., Chen, D.F., Sasaoka, T., Ulupinar, E., and Tonegawa, S. NMDA receptor-dependent refinement of somatotopic maps. Neuron 19:1201-1210 (1997). View PDF »
  • Silva, A.J., Simpson, E.M., Takahasi, J.S., Lipp. H.-P., Nakanishi, S., Wehner, J.M., Giese, K.P., Tully, T., Abel, T., Chapman, P.F., Fox, K., Grant, S., Itohara, S., Lathe, R., Mayford, M., McNamara, J.O., Morris, R.J., Picciotto, M., Roder, J., Shin, H.-S., Slesinger, P.A., Storm, D.R., Stryker, M.P., Tonegawa, S., Wang, Y., and Wolfer, D.P. Mutant mice and neuroscience: Recommendation concerning genetic background. Neuron 19:755-759 (1997). View PDF »
  • Zakhary, R., Poss, K.D., Jaffrey, S.R., Ferris,C.D., Tonegawa, S., and Snyder, S.H. Targeted gene deletion of heme oxygenase 2 reveals neural role for carbon monoxide. PNAS 94:14848-14853 (1997). View PDF »
  • Poss, K.D. and Tonegawa, S. Reduced stress defense in heme oxygenase 1-deficient cells. PNAS 94:10925-10930 (1997). View PDF »
  • Poss, K.D. and Tonegawa, S. Heme-oxygenase-1 is required for mammalian iron reutilization. PNAS 94:10919-10924 (1997). View PDF »
  • Xu, M., Koeltzow, T., Santiago, G.T., Moratalla, R., Cooper, D., Hu, X.-T., White, N.M., Graybiel, A.M., White, F.J., and Tonegawa, S. Dopamine D3 receptor mutant mice exhibit increased behavioral sensitivity to concurrent stimulation of D1 and D2 receptors. Neuron 19:837-848 (1997). View PDF »
  • Malmberg, A.B., Chen, C., Tonegawa, S., and Basbaum, A.I. Preserved acute pain and reduced neuropathic pain in mice lacking PKCg. Science 278:279-283 (1997). View PDF »
  • Dave, V.P., Cao, Z., Browne, C., Alarcon, B., Fernandez-Miguel, G., Lafaille, J., de la Hera, A., Tonegawa, S., and Kappes, D.J. CD3 delta deficiency arrests development of the alpha beta but not the gamma delta T cell lineage. EMBO J. 16:1360-1370 (1997). View PDF »
  • Marusic-Galesic, S. and Tonegawa, S. Tolerance induction and autoimmune encephalomyelitis amelioration after administration of myelin basic protein-derived peptide. J. Exp. Med. 186:507-515 (1997). View PDF »
  • Lafaille, J., Vliebergh, F., and Tonegawa, S. Myelin basic protein-specificT helper 2 (Th2) cells cause experimental autoimmune encephalomyelitis in immunodeficient hosts rather than protect them from disease. J. Exp.Med. 186:307-312 (1997). View PDF »
  • Pereira, P., Lafaille, J.J., Gerber, D., and Tonegawa, S. The T-Cell receptor repertoire of intestinal intraepithelial g/d T lymphocytes is influenced by genes linked to the major histocompatibility complex and to the T-cel receptor loci. PNAS 94:5761-5766 (1997). View PDF »
  • Shen, J., Roderick, T.B., Chen, D.F., Xia, W., Selkoe, D.J., and Tonegawa, S. Skeletal and CNS defects in Presenilin-1 deficient mice. Cell 89:629-639 (1997). View PDF »
  • Chen, C. and Tonegawa, S. Molecular genetic analysis of synaptic plasticity, activity-dependent neural development, learning, and memory in the mammalian brain. Ann. Rev. Neurosci. 20:157-184 (1997). View PDF »
  • Wilson, M.A. and Tonegawa, S. Synaptic plasticity, place cells and spatial memory: study with second generation knockouts. TINS 20:102-106 (1997). View PDF »
  • Chen, D.F., Schneider, G.E., Martinou, J.-C., and Tonegawa, S. BCL-2 promotes regeneration of severed axons in mammalian CNS. Nature 385:434-439 (1997). View PDF »
  • Kano, M., Hashimoto, K., Kurihara, H., Watanabe, M., Inoue, Y., Aiba, A., and Tonegawa, S. Persistent multiple climbing fiber innervation of cerebellar Purkinje cells in mice lacking mGluR1. Neuron 18:71-79 (1997). View PDF »


  • Tonegawa, S., Tsien, J.Z., McHugh, T.J., Huerta, P., Blum, K.I., and Wilson, M.A. Hippocampal CA1-region-restricted knockout of NMDAR1 gene disrupts synaptic plasticity, place fields, and spatial learning. Cold Spring Harbor Symposia on Quantitative Biology 61:225-238 (1996). View PDF »
  • Xu, M., Hu, X.T., Cooper, D.C., White, F.J., and Tonegawa, S. A genetic approach to study mechanisms of cocain action. Ann. NY Acad. Sci. 801:51-63 (1996). View PDF »
  • Gerber, D.J., Periera, P., Huang, S.Y., Pelletier, C., and Tonegawa, S. Expression of aV and b3 integrin chains on murine lymphocytes. PNAS 93:14698-14703 (1996). View PDF »
  • Goda, Y., Stevens, C.F., and Tonegawa, S. Phorbol ester effects at hippocampal synapses act independently of the g isoform of PKC. Learning and Memory 3:182-187 (1996). View PDF »
  • Moratalla, R., Xu, M., Tonegawa, S., and Graybiel, A. Cellular responses to psychomotor stimulant and neuroleptic drugs are abnormal in mice lacking the D1 dopamine receptor. PNAS 93:14928-14933 (1996). View PDF »
  • McHugh, T.J., Blum, K.I., Tsien, J.Z., Tonegawa, S., and Wilson, M.A. Impaired hippocampal representation of space in CA1-specific NMDAR1 knockout mice. Cell 87, 1339-1349. (1996). View PDF »
  • Tsien, J.Z., Huerta, P.T., and Tonegawa, S. The essential role of hippocampal CA1 NMDA receptor-dependent synaptic plasticity in spatial memory. Cell 87:1327-1338 (1996). View PDF »
  • Tsien, J.Z., Chen, D.F., Gerber, D., Tom, C., Mercer, E.H., Anderson, D.J., Mayford, M., Kandel, E.R., and Tonegawa, S.T. Subregion-and cell type-restricted gene knockout in mouse brain. Cell 87:1317-1326 (1996). View PDF »
  • Chen, C., Kim, J.J., Thompson, R.F., Tonegawa, S. Hippocampal lesions impair contextual fear conditioning in two strains of mice. Behav. Neurosci. 110:1177 (1996). View PDF »
  • Jacobs, H., Iacomini, J., van de Ven, M., Tonegawa, S., and Berns, A. Domains of the TCRb-chain required for ea rly thymocyte development. J. Exp. Med. 184:1833 (1996). View PDF »
  • Mizoguchi, A., Mizoguchi, E. Tonegawa, S., and Bhan, A.K. Alteration of polyclonal to oligoclonal immune response to cecal aerobic bacterial antigens in TCR-a mutant mice with inflammatory bowel disease. International Immunology, 8:1387-1394 (1996). View PDF »
  • Ji, Z., Ebralidze, A., Tonegawa, S., and Vogel, M.W. Spinocerebellar mossy fiber terminal topography in the NMDAR2C/PKCg double mutant cerebellum. Develop. Brain Res. 97:138 (1996). View PDF »
  • Ebralidze, A.K., Rossi, D.J., Tonegawa, S., and Slater, N.T. Modification of NMDA receptor-channels and synaptic transmission by targeted disruption of the NR2C gene. J. Neurosci. 16:5014 (1996). View PDF »
  • Tonegawa, S. Analysis of synaptic plasticity and memory in the mammalian brain by the gene knockout technology. Proceedings of the 1995 Yakult Symposium. (1996). View PDF »
  • Mizoguchi, A., Mizoguhci, E., Chiba, C., Spiekermann, G.M., Tonegawa, S., Nagler-Anderson, C., and Bhan, A.K. Cytokine imbalance and aitoantibody production in TCR-a mutant mice with inflammatory bowel disease. J. Exp. Medicine 183:847-856 (1996). View PDF »
  • Tonegawa, S. A genetic approach to synaptic plasticity and learning. In: Konnerth, A., Tsien, R.Y., Mikoshiba, K, and Altman, J. (Eds.) Coincidence Detection in the Nervous System. Strasbourg, HFSP, (1996), pp.159-164. View PDF »
  • Fujihashi, K., McGhee, J.R., Kweon, M.-N., Cooper, M.D., Tonegawa, S., Takahashi, I., Hiroi, T., Mestecky J., and Kiyono, H. gd T cell dificient mice have impaired mucosal IgA responses. J. Exp. Medicine, 183:1-8 (1996). View PDF »
  • Wu, M., Chen., D.F., Sasaoka, T., and Tonegawa, S. Neural tube defects and abnormal brain development in F52-deficient mice. PNAS 93:2110-2115 (1996). View PDF »
  • Pereira, P., Gerber, D., Regnault, A., Hermitte, V., Coutinho, A., and Tonegawa, S. Rearrangement and expression of Vg1, Vg2, and Vg3 T cell receptor g genes in C57BL/6 mice. International Immunity 8:83-90 (1996). View PDF »


  • Tonegawa, S. Mammalian learning and memory studied by gene targeting. Ann. N.Y. Acad. Sci. 758:213-217 (1995). View PDF »
  • Tonegawa, S. Somatic generation of antibody diversity. In: Immunology: The Making of a Modern Science. Academic Press (1995). View PDF »
  • Oxenius, A., Bachmann, M.F., Ashton-Rickardt, P.G., Tonegawa, S., Zinkernagel, R.M., and Hengartner, H. Presentation of endogenous viral proteins in association with major histocompatibility complex class II: on the role of intracellular compartmentalization, invariant chain and the TAP transporter system. Eur. J. Immunol. 25:3402-3411 (1995). View PDF »
  • Prosser, H.M., and Tonegawa, S. T cell receptor V(D)K recombinations: mechanisms and developmental regulation. In: Bell, J.I., Owen, M.J., and Simpson, E. (Eds.) T Cell Receptors, Oxford: Oxford University Press, 326-351 (1995). View PDF »
  • Tonegawa, S., Li, Y., Erzurumlu, R.S., Jhaveri, S., Chen, C., Goday, Y., Paylor, R., Silva, A.J., Kim., J.J., Wehner, J.M., Stevens, C.F., and Abeliovich, A. The gene knockout technology for the analysis of learning and memory, and neural development. Prog. Brain Res. 105:3-14 (1995). View PDF »
  • Poss, K., Thomas, M.J., Ebralidze, A.K., O’Dell, T.T., and Tonegawa, S. Hippocampal long-term potentiation is normal in hemo-oxygenase-2 mutant mice. Neuron 15:867-873 (1995). View PDF »
  • Ljunggren, H.-G., Van Kaer, L., Sabatine, M.S., Auchincloss, Jr., H., Tonegawa, S., and Ploegh, H.L. MHC class I expression and CD8+ T cell development in TAP1/B2-microglobulin double mutant mice. Intl. Immunology 7:975-984 (1995). View PDF »
  • Hombach, J., Pircher, H., Tonegawa, S., and Zinkernagel, R.M. Strictly transporter of antigen presentation (TAP)-dependent presentation of an immunodominant cytotoxic T lymphocyte epitope in the signal sequence of a virus protein. J. Exp.Med 182:1615-1619 (1995). View PDF »
  • Ladel, C.H., Hess, J., Daugelat, S., Mombaerts, P., Tonegawa, S., and Kaufman, S.H.E. Contribution of a/b and g/d T lymphocytes to immunity against Mycobacterium bovis Bacillus Calmette Guerin: studies with T cell receptor-deficient mutant mice. Eur. J. Immunol. 25:838-846 (1995). View PDF »
  • Chen, C., Kano, M., Abeliovich, A., Chen, L., Bao, S., Kim, Jeansok J., Hashimoto, K., Thompson, R., and Tonegawa, S. Impaired motor coordination correlates with persistent multiple climbing fiber innervation in PKCg mutant mice. Cell 83:1233-1242 (1995). View PDF »
  • Kano, M., Hashimoto, K., Chen, C., Abeliovich, A., Aiba, A., Kurihara, H., Watanabe, M., Inoue, Y., and Tonegawa, S. Impaired synapse elimination during cerebellar development in PKCg mutant mice. Cell 83:1223-1231 (1995). View PDF »
  • Komano, H., Fujiura, Y., Kawaguchi, M., Matsumoto, S., Hashimoto, Y., Obana, S., Mombaerts, P., Tonegawa, S., Yamamoto, H., Itohara, S., Nanno, M., and Ishikawa, H. Homeostatic regulation of intestinal epithelia by intraepithelial gd T cells. PNAS 92:6147-6151 (1995). View PDF »
  • Butler, L.S., Silva, A.J., Abeliovich, A., Watanabe, Yoshinori, Tonegawa, S., and McNamara, J.O. Limbic epilepsy in a-Calcium-Calmodulin Kinase II mutant mice. PNAS 92:6852-6855 (1995). View PDF »
  • Hsia, A.Y., Salin, P.A. , Castillo, P.E., Aiba, A., Abeliovich, A., Tonegawa, S., and Nicoll, R.A. Evidence against a role for metabotropic glutamate receptors in mossy fiber LTP: the use of mutant mice and pharmacological antagonists. Neuropharmacology 34:1567-1572 (1995). View PDF »
  • Langhorne, J., Mombaerts, P., and Tonegawa, S. ab T cells and gd T cells in the immune response to the erythrocytic stages of malaria in mice. Immunity 7:1005-1011 (1995). View PDF »
  • Bachmann, M., Oxenius, A., Pircher, H., Hengartner, H., Ashton-Rickardt, P., Tonegawa, S., and Zinkernagel, R. TAP1-Independent loading of Class I molecules by exogenous viral proteins. Eur. J. Immunol. 25:1739-1743 (1995). View PDF »
  • Pereira, P., Gerber, D., Huang, S.Y., and Tonegawa, S. Ontogenic development and tissue distribution of Vg1- expressing gd T lymphocytes in normal mice. Journal of Experimental Medicine 182:1921-1930 (1995). View PDF »
  • Ljunggren, H-G., Van Kaer, L., Ashton-Rickardt, P.G., Tonegawa, S., and Ploegh, H.L. Differential reactivity of residual CD8+ T lymphocytes in TAP1 and b2-microglobulin mutant mice. European Journal of Immunology 25:174-178 (1995). View PDF »
  • Levelt, C.N., Huang, M., Mombaerts, P., Mak T.W., Tonegawa, S., Eichmann, K., and Terhorst, C. Regulation of Thymocyte Development through CD3. Functional Dissociation between p56 lck and CD3z in early and late thymic selection.Immunity, 3:215-222 (1995). View PDF »
  • Eichelberger, M., McMickle, A., Blackman, M., Mombaerts, P., Tonegawa, S., and Doherty, P.C. Functional analysis of the TCR a-b+ cells that accumulate in the pneumonic lung of influenza virus infected TCR a (-/-) mice. Journal of Immunology, 154:1569-1576 (1995). View PDF »
  • Van Santen, H.M., Machold, R., Woolsey, A., Ploegh, H.L., Ashton-Rickardt, P., Van Kaer, L., and Tonegawa, S. Increase in Positive Selection of CD8+ T Cells in TAP1-Mutant Mice by Human b2-Microglobulin Transgene. J. Exp. Med 181:787-792 (1995). View PDF »
  • Harris, R. A., McQuilkin, S.J., Paylor, R., Abelovich, A., Tonegawa, S., and Wehner, J.M. Mutant mice lacking the gamma isoform of protein kinase C show decreased behavioral actions of ethanol and altered function of gamma-aminobuytrate type A receptors. PNAS 92:3658-3662 (1995). View PDF »
  • Mombaerts, P., Terhorst, C., Jacks, T., Tonegawa, S., and Sancho, J. Characterization of immature thymocyte lines derived from T-cell receptor or recombination activating gene 1 and p53 double mutant mice. PNAS 92:7420-7424 (1995). View PDF »


  • Godfrey, D.I., Kennedy, J., Mombaerts, P., Tonegawa, S., and Zlotnik, A. Onset of TCRb gene rearrangementand role of TCRb expression during thymocyte expression during CD3- CD4- CD8- thymocyte differentiation. J. Immunol. 152:4783-4792 (1994). View PDF »
  • Tonegawa, S. Gene targeting: a new approach for the analysis of mammalian memory and learning. Prog. Clin. Biol. Res. (PZ5) 390:5-18 (1994). View PDF »
  • Xu, M., Hu, X-T, Cooper, D.C., Moratalla, R., Graybiel, A.M., White, F.J., and Tonegawa, S. Elimination of cocaine-induced hyperactivity and dopamine-mediated neurophysioloigcal effects in dopamine D1 Receptor Mutant Mice. Cell 79:945-955 (1994). View PDF »
  • Xu, M., Moratalla, R., Gold, L.H., Hiroi, N., Koob, G.F., Graybiel, A.M., and Tonegawa, S. Dopamine D1 Receptor mutant mice are deficient in striatal expression of dynorphin and in dopamine-mediated behavioral responses. Cell 79:729-742 (1994). View PDF »
  • Aiba, A., Chen, C., Herrup, K., Rosenmund, C., Stevens, C., and Tonegawa, S. Reduced hippocampal long-term potentiation and context-specific deficit in associative learning in mGluR1 mutant mice. Cell 79:365-375 (1994). View PDF »
  • Aiba, A., Kano, M., Chen, C., Stanton, M., Fox, G., Herrup, K., Zwingman, T. and Tonegawa, S. Deficient cerebellar long-term depression and impaired motor learning in mGluR1 mutant mice. Cell 79:377-388 (1994). View PDF »
  • Stevens, C.F., Tonegawa, S., and Wang, Y. The role of calcium-calmodulin kinase II in three forms of synaptic plasticity. Current Biology 4:687-693 (1994) View PDF »
  • Lafaille, J.J., Nagashima, K., Katsuki, M., and Tonegawa, S. High incidence of spontaneous autoimmune encephalomyelitis in immunodeficient anti-myelin basic protein T cell receptor transgenic mice. Cell 78:399-408 (1994). View PDF »
  • Mombaerts, P., Anderson, S., Perlmutter, R., Mak, T.W., and Tonegawa, S. An activated lck transgene promotes thymocyte development in RAG-1 mutant mice. Immunity 1:261-267 (1994). View PDF »
  • Ashton-Rickardt, P.G., and Tonegawa, S. A differential avidity model for T cell selection. Immunology Today 15:362-366 (1994). View PDF »
  • Chen, C., Rainnie, D., Greene, R and Tonegawa, S. Abnormal fear response and aggressive behavior in mutant mice deficient for a-Calcium-Calmodulin Kinase II. Science 265:291-294 (1994). View PDF »
  • Van Kaer, L., Ashton-Rickardt, P.G., Eichelberger, M., Gaczynska, M., Nagashima, K., Rock, K., Goldberg, A.L., Doherty, P.C., and Tonegawa, S. Altered peptidase and viral-specific T cell response in LMP2 mutant mice. Immunity 1:533-541 (1994). View PDF »
  • Ashton-Rickardt, P.G., Bandeira, A., Delaney J.R., Van Kaer, L., Pircher, H-P., Zinkernagel, R.M., and Tonegawa, S. Evidence for a differential avidity model of T Cell selection in the thymus. Cell 76:651-663 (1994). View PDF »
  • Mombaerts, P., Mizoguchi, E., Ljunggren, H-G., Iacomini, J., Ishikawa, H., Wang, L., Grusby, M., Glimcher, L., Winn, H.J., Bhan, A.K., and Tonegawa, S. Peripheral lymphoid development and function in TCR mutant mice. International Immunology 6:1061-1070 (1994). View PDF »
  • Aldrich, C.J., Ljunggren H.-G., Van Kaer, L., Ashton-Rickardt, P.G., Tonegawa, S., and Forman J. Positive selection of self-and alloreactive CD8+ T cells in Tap-1 mutant mice. PNAS 91:6525-6528 (1994). View PDF »
  • Li, Y., Erzurumlu, R., Chen, C., Jhaveri, S., and Tonegawa, S. Whisker-Related neuronal patterns fail to develop in the trigeminal brainstem nuclei of NMDAR1 knockout mice. Cell 76:427-437 (1994). View PDF »
  • Tsuji, M., Mombaerts, P., Lefrancois, L., Nussenzweig, R.S., Zavala, F., and Tonegawa, S. gd T cells contribute to immunity against the liver stages of malaria in ab T-cell-deficient mice. PNAS 91:345-349 (1994). View PDF »
  • Schumacher, T.N.M., Kantesaria, D.V., Heemels, M-T., Ashton-Rickardt, P.G., Shepherd, J.C., Fruh, K., Yang, Y., Peterson, P.A., Janeway, C.A Jr., Tonegawa, S., and Ploegh, H.L. Peptide length and sequence specificity of the mouse TAP1/TAP2 translocator. J. Exp. Med. 179:533-540 (1994). View PDF »
  • Ljunggren, H.-G., Van Kaer, L., Ploegh, H.L. and Tonegawa, S. Altered natural killer cell repertoire in TAP1 mutant mice. PNAS 91: 6520-6524 (1994). View PDF »


  • Levelt, C.N., Mombaerts, P., Iglesias, A., Tonegawa, S., and Eichmann, K. Restoration of early thymocyte differentiation in T-cell receptor b-chain-deficient mutant mice by transmembrane signaling through CD3e. PNAS 90:11401-11405 (1993). View PDF »
  • Abeliovich, A., Paylor, R., Chen, C., Kim, J.J., Wehner, J.M., and Tonegawa, S. PKCg mutant mice exhibit mild deficits in spatial and contextual learning. Cell 75:1263-1271 (1993). View PDF »
  • Abeliovich, A., Chen, C., Goda, Y., Silva, A.J., Stevens, C.F., and Tonegawa, S. Modified hippocampal long term potentiation in PKCg mutant mice. Cell 75:1253-1262 (1993). View PDF »
  • Moriwaki, S., Korn, B., Ichikawa, Y., Van Kaer, L., and Tonegawa, S. Amino acid substitutions in the floor of the putative antigen-binding site of H-2T22 affect recognition by a gd T-cell receptor. PNAS 90:11396-11400 (1993). View PDF »
  • Spaner, D., Migita, K., Ochi, A., Shannon, J., Miller, R.G., Pereira, P., Tonegawa, S., and Phillips, R.A. gdT cells differentiate into a functional but nonproliferative state during a normal immune response. PNAS 90:8415- 8419 (1993). View PDF »
  • Ishikawa, H., Li, Y., Abeliovich, A., Yamamoto, S., Kaufmann, S.H.E., and Tonegawa, S. Cytotoxic and Interferon g-producing activities of gd T cells in the mouse intestinal epithelium are strain dependent. PNAS 90:8204- 8208 (1993). View PDF »
  • Ashton-Rickardt, P.G., Van Kaer, L., Schumacher, T.N.M., Ploegh, H.L., and Tonegawa, S. Repertoire-Determining Role of Peptide in the Positive Selection of CD8+ T cells. Immunol. Rev 135:157-182 (1993). View PDF »
  • Mombaerts, P., Arnoldi, J., Friedemann, R., Tonegawa, S., and Kaufmann, S.H.E. Differential Roles of ab and gd T cells in Immunity Against an Intracellular Bacterial Pathogen. Nature 365:53-56 (1993). View PDF »
  • Mombaerts, P., Mizoguchi, E., Grusby, M.J., Glimcher, L.H., Bhan, A.K., and Tonegawa, S. Spontaneous Development of Inflammatory Bowel Disease in T Cell Receptor Mutant Mice. Cell 75:275-282 (1993). View PDF »
  • Shepherd, J.C., Schumacher, T.N.M., Ashton-Rickardt, P.G., Imaeda, S., Ploegh, H.L., Janeway, C.A. Jr., and Tonegawa, S. TAP1-Dependent Peptide Translocation in vitro is ATP Dependent and Peptide Selective. Cell 74:577-584 (1993). View PDF »
  • Ashton-Rickardt, P.G., Van Kaer, L., Schumacher, T.N.M., Ploegh, H.L., and Tonegawa, S. Peptide Contributes to the Specificity of Positive Selection of CD8+ T cells in the Thymus. Cell 73:1041-1049 (1993). View PDF »
  • Itohara, S., Mombaerts, P., Lafaille, J., Iacomini, J., Nelson, A., Clarke, A., Hooper, M., Farr, A., and Tonegawa, S. T Cell Receptor d Gene Mutant Mice: Independent Generation of ab T cells and Programmed Rearrangements of gd TCR Genes. Cell 72:337-348 (1993). View PDF »
  • Haas, W., Pereira, P., and Tonegawa, S. gd Cells. Ann. Rev. of Immun. 11: 637-685 (1993). View PDF »


  • Van Kaer, L., Ashton-Rickardt, P.G., Ploegh, H.L., and Tonegawa, S. TAP1 Mutant Mice are Deficient in Antigen Presentation, Surface Class I Molecules, and CD4-8+ T Cells. Cell 71:1205-1214 (1992). View PDF »
  • Silva, A., Wang, Y., Paylor, R., Wehner, J., Stevens, C., and Tonegawa, S. a Calcium/Calmodulin Kinase II Mutant Mice: Deficient Long Term Potentiation and Impaired Spatial Learning. Cold Spring Harbor Symposium on Quantitative Biology ‘The Cell Surface,’ vol. 57 (1992). View PDF »
  • Silva, A., Paylor, R., Wehner, J.M., and Tonegawa, S. Impaired Spatial Learning in a Calcium-Calmodulin Kinase II Mutant Mice. Science 257:206-211 (1992). View PDF »
  •  Silva, A., Stevens, C., Tonegawa, S., and Wang, Y. Deficient Hippocampal Long-Term Potentiation in a Calcium-Calmodulin Kinase II Mutant Mice. Science 257:201-206 (1992). View PDF »
  • Abeliovich, A., Gerber, D., Tanaka, O., Katsuki, M., Graybiel, A., and Tonegawa, S. On Somatic Recombination in the Central Nervous System of Transgenic Mice. Science 257:404-408 (1992). View PDF »
  • Mombaerts, P., Clarke, A.R., Rudnicki, M.A., Iacomini, J., Itohara, S., Lafaille, J.J., Wang, L., Ishikawa, Y., Jaenisch, R., Hooper, M.L., and Tonegawa, S. Mutations in T-cell antigen receptor genes a and b block thymocyte development at different stages. Nature 360:225-231 (1992). View PDF »
  • Mombaerts, P., Iacomini, J., Johnson, R.S., Herrup, K., Tonegawa, S., and Papaioannou, V.E. RAG-1 Deficient Mice Have no Mature B and T Lymphocytes. Cell 68:869-877 (1992). View PDF »
  • Haas, W., and Tonegawa, S. Development and Selection of gd T cells. Current Opinions in Immunology 4:147-155 (1992). View PDF »
  • Pereira, P., Zijlstra, M., McMaster, J., Loring, J.M., Jaenisch, R., and Tonegawa, S. Blockade of Transgenic gd T Cell Development in b2-microglobulin Deficient Mice. EMBO J. 11:25-31 (1992). View PDF »


  • Kappes, D.J., and Tonegawa, S. Surface Expression of Alternative forms of the TCR/CD3 Complex. PNAS 88:10619-10623 (1991). View PDF »
  • Wu, M., Van Kaer, L., Itohara, S., and Tonegawa, S. Highly Restricted Expression of the Thymus Leukemia Antigens on Intestinal Epithelial Cells. J. Exp. Med. 174:213-218 (1991). View PDF »
  • Mombaerts, P., Clarke, A.R., Hooper, M.L., and Tonegawa, S. Creation of a Large Genomic Deletion at the T-Cell Antigen Receptor b-subunit locus in Mouse Embryonic Stem Cells by Gene Targeting. PNAS 88:3084-3087 (1991). View PDF »
  • Kappes, D.J., Browne, C.P., and Tonegawa, S. Identification of a T Cell-Specific Enhancer at the Locus Encoding T-cell Antigen Receptor g Chain. PNAS 88:2204- 2208 (1991). View PDF »
  • Van Kaer, L., Wu, M., Ichikawa, Y., Ito, K., Bonneville, M., Ostrand-Rosenberg, S., Murphy, D.B., and Tonegawa, S. Recognition of MHC TL Gene Products by gd T Cells. Immunological Reviews 120:89-115 (1991). View PDF »
  • Holland, G.D., Ito, K., Kaehler, D.A., Tonegawa, S., and Risser, R. Thymic Targets for Abelson Murine Leukemia Virus Are Early g/d Lymphocytes. PNAS 88:3700-3704 (1991). View PDF »
  • Bandeira, A., Itohara, S., Bonneville, M., Burlen-Defranoux, O., Mota-Santos, T., Coutinho, A., and Tonegawa, S. Extra Thymic Origin of Intestinal Intraepithelial T lymphocytes Bearing T-Cell Antigen Receptor gd. PNAS 88:43-47 (1991). View PDF »


  • Ito, K., Van Kaer, L., Bonneville, M., Hsu, S., Murphy, D., and Tonegawa, S. Recognition of the Product of a Novel MHC TL Region Gene (27b) by a Mouse gd T Cell Receptor. Cell 62:549-561 (1990). View PDF »
  • Itohara, S., and Tonegawa, S. Selection of gd T Cells with Canonical T-Cell Antigen Receptors in Fetal Thymus. PNAS 87:7935-7938 (1990). View PDF »
  • Tonegawa, S., Berns, A., Bonneville, M., Farr, A.G., Ishida, I., Ito, K., Itohara, S., Janeway, C.A. Jr., Kanagawa, O., Katsuki, M., Kubo, R., Lafaille, J.J., Mombaerts, P., Murphy, D.B., Natanishi, N., Takagaki, Y., VanKaer, L., and Veebeek, S. Diversity, Development, Ligands, and Probable Functions of gd T Cells. Progress in Immunology VII Springer-Verlag, Eds. F. Melchers et al. pp. 243-257 (1990). View PDF »
  • Bandeira, A., Mota-Santos, T., Itohara, S., Degermann, S., Heusser, C., Tonegawa, S., and Coutinho, A. Localization of g/d T Cells to the Intestinal Epithelium is Independent of Normal Microbial Colonization. J. Exp. Med. 172:239-244 (1990). View PDF »
  • Bonneville, M., Ishida, I., Itohara, S., Verbeek, S., Berns, A., Kanagawa, O., Haas, W., and Tonegawa, S. Self Tolerance to Transgenic gd T cells by Intrathymic Inactivation. Nature 344:163-165 (1990). View PDF »
  • Lafaille, J.J., Haas, W., Coutinho, A., and Tonegawa, S. Positive Selection of gd T Cells. Immunology Today 11:75-78 (1990). View PDF »
  • Farr, A., Hosier, S., Nelson, A., Itohara, S., and Tonegawa, S. Distribution of thymocytes expressing gd receptors in the murine thymus during development. J. Immunol. 144:492-498 (1990). View PDF »
  • Bonneville, M., Itohara, S., Krecko, E.G., Mombaerts, P., Ishida, I., Katsuki, M., Berns, A., Farr, A.G., Janeway, C.A. Jr., and Tonegawa, S. Transgenic Mice Demonstrate that Epithelial Homing of g / d T Cells is Determined by Cell Lineages Independent of T Cell Receptor Specificity. J. Exp. Med. 171:1015-1026 (1990). View PDF »
  • Itohara, S., Farr, A.G., Lafaille, J., Bonneville, M., Takagaki, Y., Haas, W., and Tonegawa, S. Homing of a gd Thymocyte Subset with Homogeneous T-Cell Receptors to Mucosal Epithelia. Nature 343:754-757 (1990). View PDF »
  • Ishida, I., Verbeek, S., Bonneville, M., Itohara, S., Berns, A., and Tonegawa, S. T Cell Receptor gd Transgenic Mice Suggest a Role of a g Gene Silencer in the Generation of ab T cells. PNAS 87:3067-3071 (1990). View PDF »


  • Bonneville, M., Ishida, I., Mombaerts, P., Katsuki, M., Verbeek, S., Berns, A., and Tonegawa, S. Blockage of ab T-cell Development by TCR gd transgenes. Nature 342:931-934 (1989). View PDF »
  • Minoprio, P., Itohara, S., Heusser, C., Tonegawa, S., and Coutinho, A. Immunobiology of Murine T. Cruzi Infection: The Predominance of Parasite-nonspecific Responses and the Activation of TCRI T Cells. Immunological Reviews 112:183-207 (1989).
  • Tonegawa, S., Berns, A., Bonneville, M., Farr, A., Ishida, I., Ito, K., Itohara, S., Janeway, C.A. Jr., Kanagawa, O., Katsuki, M., Kubo, R., Lafaille, J.J., Mombaerts, P., Murphy, D., Nakanishi, N., Takagaki, Y., Van Kaer, L., and Verbeek, S. Diversity, development, ligands, and probable functions of gd T cells. Cold Spring Harbor Quant. Biol. 14:31-44 (1989). View PDF »
  • Lafaille, J.J., DeCloux, A., Bonneville, M., Takagaki, Y., and Tonegawa, S. Junctional Sequences of T Cell Receptor gd genes: Implications for gd T Cell Lineages and for a Novel Intermediate of V-(D)-J Joining. Cell 59:859-870 (1989). View PDF »
  • Bonneville, M., Ito, K., Krecko, E.G., Itohara, S., Kappes, D., Ishida, I., Kanagawa, O., Janeway, C.A. Jr., Murphy, D.B., and Tonegawa, S. Recognition of a Self Major Histocompatibility Complex TL Region Product by gd T-Cell Receptors. PNAS 86: 5928-5932 (1989). View PDF »
  • Itohara, S., Nakanishi, N., Kanagawa, O., Kubo, R., and Tonegawa, S. Monoclonal Antibodies Specific to Native Murine T-Cell Receptor gd: Analysis of gd T cells in Thymic Ontogeny and in Peripheral Lymphoid Organs. PNAS 86:5094-5098 (1989). View PDF »
  • Takagaki, Y., DeCloux, A., Bonneville, M., and Tonegawa, S. Diversity of gd T-cell Receptors on Murine Intestinal Intra-Epithelial Lymphocytes. Nature 339:712-714 (1989). View PDF »
  • Takagaki, Y., Nakanishi, N., Ishida, I., Kanagawa, O., and Tonegawa, S. T Cell Receptor g and d Genes Preferentially Utilized by Adult Thymocytes for the Surface Expression. J. Immunol 142:2112-2121 (1989). View PDF »
  • Ito, K., Bonneville, M., Takagaki, Y., Nakanishi, N., Kanagawa, O., Krecko, E.G., and Tonegawa, S. Different gd T-Cell Receptors are Expressed on Thymocytes at Different Stages of Development. PNAS 86:631-635 (1989). View PDF »
  • Bonneville, M., Janeway, C.A. Jr., Ito, K., Haser, W., Ishida, I., Nakanishi, N., and Tonegawa, S. Intestinal Intraepithelial Lymphocytes are a Distinct Set of gd T Cells. Nature 336:479-481 (1989). View PDF »


  • Tonegawa, S. Somatic Generation of Immune Diversity. (Nobel Lecture in Physiology or Medicine 1987.) Bioscience Reports 8:3-26 (1988). View PDF »
  • Tonegawa, S. Antibody and T-Cell Receptors. JAMA 259:1845-1847 (1988). View PDF »
  • Von Boehmer, H., Bonneville, M., Ishida, I., Ryser, S., Lincoln, G., Smith, R.T., Kishi, H., Scott, B., Kisielow, P., and Tonegawa, S. Early Expression of a T-Cell Receptor b-chain Transgene Suppresses Rearrangement of the Vg4 Gene Segment. PNAS 85:9729-9732 (1988). View PDF »


  • Saito, H., Koyama, T., Georgopoulos, K., Clevers, H., Haser, W.G., LeBien, T., Tonegawa, S., and Terhorst, C. Close Linkage of the Mouse and Human CD3 g-and d-chain Genes Suggests that their Transcription is Controlled by Common Regulatory Elements. PNAS 84:9131-9134 (1987). View PDF »
  • Heilig, J.S. and Tonegawa, S. T-Cell g Gene is Allelically but not Isotypically Excluded and is not Required in Known Functional T-Cell Subsets. PNAS 84:8070-8074 (1987). View PDF »
  • Haser, W.G., Saito, H., and Tonegawa, S. Cloning and Sequencing of Murine T3g cDNA from a Subtractive cDNA Library. J. Exp. Med. 166:1186-1191 (1987). View PDF »
  • Maeda, K., Nakanishi, N., Rogers, B.L., Haser, W.G., Shitara, K., Yoshida, H., Takagaki, Y., Augustin, A.A., and Tonegawa, S. Expression of the T-cell Receptor g -Chain Products on the Surface of Peripheral T-Cells and T-Cell Blasts Generated by Allogeneic mixed Lymphocyte Reaction. PNAS 84:6536-6540 (1987). View PDF »
  • Koyama, T., Hall, L.R., Haser, W.G., Tonegawa, S., and Saito, H. Structure of a Cytotoxic T-lymphocyte-specific Gene Shows a Strong Homology to Fibrinogen b and g Chains. PNAS 84:1609-1613 (1987). View PDF »
  • Nakanishi, N., Maeda, K., Ito, K., Heller, M., and Tonegawa, S. Tg Protein is Expressed on Murine Fetal Thymocytes as a Disulfide-Linked Heterodimer. Nature 325:720-723 (1987). View PDF »


  • Tonegawa, S., and Saito, H. Evolutionary Aspects of Immunoglobulin-Related Genes. Chemica Scripta 26:343-349 (1986). View PDF »
  • Lieberman, J., Verret, C.R., Kranz, D.M., Hubbard, S.C., Saito, H., Raulet, D., Tonegawa, S., and Eisen, H.N. A Phosphorylated, Disulfide-linked Membrane Protein in Murine Cytotoxic T Lymphocytes. PNAS 83:7870-7874 (1986). View PDF »
  • Heilig, J.S. and Tonegawa, S. Diversity of Murine Gamma Genes and Expression in Fetal and Adult T Lymphocytes. Nature 322:836-840 (1986). View PDF »
  • Reilly, E.B., Kranz, D.M., Tonegawa, S., and Eisen, H.N. The functional g gene formed from known g gene segments is not necessary for antigen-specific responses of murine cytotoxic T lymphocytes. Nature 321:878-880 (1986). View PDF »
  • Novotny, J., Tonegawa, S., Saito, H., Kranz, D.M., and Eisen, H. Secondary, Tertiary and Quaternary Structure of T Cell-Specific Immunoglobulin-Like Polypeptide Chains. PNAS 83:742-746 (1986). View PDF »


  • Tonegawa, S. The Molecules of the Immune System. Scientific American 253:122-131 (1985). View PDF »
  • Ucker, D., Kurosawa, Y., and Tonegawa, S. An Unusual Clonotypic Determinant on a Cytotoxic T Lymphocyte Line is Encoded by an Immunoglobulin Heavy Chain Variable Region Gene. The Journal of Immunology 135:4204-4214 (1985). View PDF »
  • Collins, M.K.L., Tanigawa, G., Kissonerghis, A.M., Ritter, M., Price, K.M., Tonegawa, S., and Owen, M.J. Regulation of T-cell Receptor Gene Expression in Human T-Cell Development. PNAS 82:24503-4507 (1985). View PDF »
  • Hayday , A.C., Diamond, D.J., Tanigawa, G., Heilig, J.S., Folsom, V., Saito, H., and Tonegawa, S. Unusual Organization and Diversity of T-Cell Receptor a Chain Genes. Nature 316:828-832 (1985). View PDF »
  • Heilig, J.S., Glimcher, L.H., Kranz, D.M., Clayton, L.K., Greenstein, J.L., Saito, H., Maxam, A.M., Burakoff, S.J., Eisen, H.N., and Tonegawa, S.Expression of the T Cell-Specific g Gene is Unnecessary in T Cells Recognizing Class II MHC Determinants. Nature 317:68-70 (1985). View PDF »
  • Yamamura, K., Kikutani, H., Folsom, V., Clayton, L.K., Kimoto, M., Akira, S., Kashiwamura, S., Tonegawa, S., and Kishimoto, T. Functional Expression of a Microinjected Ead gene in C57BL/6 Transgenic Mice. Nature 316:67-69 (1985). View PDF »
  • Collins, M.K.L., Goodfellow, P.N., Spurr, N.K., Solomon, E., Tanigawa, G., Tonegawa, S., and Owen, M.J. The Human T-cell Receptor a-chain Gene Maps to Chromosome 14. Nature 314:273-274 (1985). View PDF »
  • Raulet, D.H., Garman, R.D., Saito, H., and Tonegawa, S. Developmental Regulation of T- Cell Receptor Gene Expression. Nature 314:103-107 (1985). View PDF »
  • Kranz, D.M., Saito, H., Disteche, C.M., Swisshelm, K., Pravtcheva, D., Ruddle, F.H., Eisen, H.N., and Tonegawa, S. Chromosomal Locations of the Murine T-Cell Receptor Alpha-Chain Gene and the T-Cell Gamma Gene. Science 227:941-945 (1985). View PDF »
  • Kranz, D.M., Saito, H., Heller, M., Takagaki, Y., Haas, W., Eisen, H.N., and Tonegawa, S. Limited Diversity of the Rearranged T cell g Gene. Nature 313:752-755 (1985). View PDF »
  • Church, G.M., Ephrussi, A., Gilbert, W., and Tonegawa, S. Cell-Type-Specific Contacts to Immunoglobulin Enhancers in Nuclei. Nature 313:798-801 (1985). View PDF »
  • Ephrussi, A., Church, G.M., Tonegawa, S., and Gilbert, W. B Lineage Specific Interactions of an Immunoglobulin Enhancer with Cellular Factors. In Vivo. Science 227:134-140 (1985). View PDF »
  • Hayday, A.C., Saito, H., Gillies, S.D., Kranz, D.M., Tanigawa, G., Eisen, H.N., and Tonegawa, S. Structure, Organization, and Somatic Rearrangement of T Cell Gamma Genes. Cell 40:259-269 (1985). View PDF »
  • Folsom, V., Gay, D., and Tonegawa, S. The b1 Domain of the Mouse Eb Chain is Important for Restricted Antigen Presentation to Helper T Cell Hybridomas. PNAS 82:1678-1682 (1985). View PDF »


  • Kranz, D.M., Tonegawa, S., and Eisen, H.N. Attachment of an Anti-Receptor Antibody to Non-target Cells Renders Them Susceptible to Lysis by a Clone of Cytotoxic T Lymphocytes. PNAS 81:7922-7926 (1984). View PDF »
  • Saito, H., Kranz, D.M., Takagaki, Y., Hayday, A.C., Eisen, H.N., and Tonegawa, S. A Third Rearranged and Expressed Gene in a Clone of Cytotoxic T Lymphocytes. Nature 312:36-40 (1984). View PDF »
  • Saito, H., Kranz, D.M., Takagaki, Y., Hayday, A.C., Eisen, H.N., and Tonegawa, S. Complete Primary Structure of a Heterodimeric T-cell Receptor Deduced from cDNA Sequences. Nature 309:757-762 (1984). View PDF »
  • Wiman, K.G., Clarkson, B., Hayday, A.C., Saito, H., Tonegawa, S., and Hayward, W.S. Activation of a Translocated c-myc Gene: Role of Structural Alterations in the Upstream Region. PNAS 81:6798-6802 (1984). View PDF »
  • Alt, F.W., Yancopoulos, G.D., Blackwell, T.K., Wood, C., Thomas, E., Boss, M., Coffman, R., Rosenberg, N., Tonegawa, S., and Baltimore, D. Ordered Rearrangement of Immunoglobulin Heavy Chain Variable Region Segments. EMBO 3:1209-1219 (1984). View PDF »
  • Gillies, S.D., Folsom, V., and Tonegawa, S. Cell Type-specific Enhancer Element Associated with a Mouse MHC gene, Eb. Nature 310: 549-554 (1984). View PDF »
  • Reilly, E.B., Blomberg, B., Imanishi-Kari, T., Tonegawa, S., and Eisen, H.N. Restricted Association of V and J-C Gene Segments for Mouse l Light Chains. PNAS 81:2484-2488 (1984). View PDF »
  • Folsom, V., Gold, D.P., White, J., Marrack, P., Kappler, J., and Tonegawa, S. Functional and Inducible Expression of a Transfected Murine Class II Major Histocompatibility Complex Gene. PNAS 80:2045-2049 (1984). View PDF »
  • Heinrich, G., Traunecker, A., and Tonegawa, S. Somatic Mutation Creates Diversity in the Major Group of Mouse Immunoglobulin k Light Chains. J. Exp. Med. 159:417-435 (1984). View PDF »
  • Hayday, A, C., Gillies, S. D., Saito, H., Wood, C., Wiman, K., Hayward, W. S., and Tonegawa, S. Activation of Translocated Human c-myc Gene by an Enhancer in the Immunoglobulin Heavy-Chain Locus. Nature 307:334-340 (1984). View PDF »


  • Saito, H., Hayday, A.C., Wiman, K., Hayward, W.S., and Tonegawa, S. Activation of the c-myc Gene by Translocation: A model for Translational Control. PNAS 80:7476-7480 (1983). View PDF »
  • Saito, H., Maki, R.A., Clayton, L.K., and Tonegawa, S. Complete Primary Structures of the Eb Chain and Gene of the Mouse Major Histocompatibility Complex. PNAS 80:5520-5524 (1983). View PDF »
  • Gillies, S.D., Morrison, S.L., Oi, V.T., and Tonegawa, S. A tissue Specific Transcription Enhancer Element is Located in the Major Intron of a Rearranged Immunoglobulin Heavy Chain Gene. Cell 33:717-728 (1983). View PDF »
  • Wood, C., and Tonegawa, S. Diversity and Joining Segments of Mouse Immunoglobulin Heavy Chain Genes are Closely Linked and in the Same Orientation: Implications for the Joining Mechanism. PNAS 80:3030-3034 (1983). View PDF »
  • Tonegawa, S. Somatic Generation of Antibody Diversity. Nature 302:575-581 (1983). View PDF »
  • Gillies, S. D., and Tonegawa, S. Expression of Cloned Immunoglobulin Genes Introduced into Mouse L Cells. Nucleic Acids Res. 11:7981-7997 (1983). View PDF »


  • Blomberg, B., and Tonegawa, S.Organization and Expression of Mouse l Light Chain Immunoglobulin Genes. In: Gene Regulation UCLA Symposium on Molecular and Cellular Biology vol. 26 (1982). View PDF »
  • Tonegawa, S., Kurosawa, Y., Blomberg, B., Heinrich, G., Haas, W., and von Boehmer, H. Genetic Origins of B and T Cell Antibodies. Nucleic Acids Res.: Future Development. (K. Mizobuchi, I. Watanabe, and J.D. Watson. eds.) pp. 197-211, Academic Press Japan, Inc. Tokyo. View PDF »
  • Kurosawa, Y., and Tonegawa, S. Organization, Structure, and Assembly of Immunoglobulin Heavy Chain Diversity DNA Segments. J. Exp. Med. 155:201-218 (1982). View PDF »
  • Blomberg, B., and Tonegawa, S. DNA sequences of the joining regions of mouse l light chain immunoglobulin genes. PNAS 9:530-533 (1982). View PDF »


  • Blomberg, B., Traunecker, A., Eisen, H., and Tonegawa, S. Organization of four mouse l light chain immunoglobulin genes. PNAS 78:3765-3769 (1981). View PDF »
  • Maki, R., Roeder, W., Traunecker, A., Sidman, C., Wabl, M., Raschke, W., and Tonegawa, S. The Role of DNA Rearrangement and Alternative RNA Processing in the Expression of Immunoglobulin Delta Genes. Cell 24:353-365 (1981). View PDF »
  • Kurosawa, Y., von Boehmer, H., Haas, W., Sakano, H., Traunecker, A., and Tonegawa, S. Identification of D segments of immunoglobulin heavy-chain genes and their rearrangement in T Lymphocytes. Nature 290:565-570 (1981). View PDF »
  • Sakano, H., Kurosawa, Y., Weigert, M., and Tonegawa, S. Identification and nucleotide sequence of a diversity DNA segment (D) of immunoglobulin heavy-chain genes. Nature 290:562-565 (1981). View PDF »
  • Roeder, W., Maki, R., Traunecker, A., and Tonegawa, S. Linkage of the four g subclass heavy chain genes. PNAS 78:474-478 (1981). View PDF »
  • Tonegawa, S. Somatic Recombination and Mosaic Structure of Immunoglobulin Genes. The Harvey Lecture Series 75:61-83 (1981). View PDF »


  • Tonegawa, S., Sakano, H., Maki, R., Traunecker, A., Heinrich, G., Roeder, W., and Kurosawa, Y. Somatic Reorganization of Immunoglobulin Genes During Lymphocyte Differentiation. Cold Spring Harbor Symp. Quant. Biol. 45:839-858 (1980). View PDF »
  • Maki, R., Kearney, J., Paige, C., and Tonegawa, S. Immunoglobulin Gene Rearrangement in Immature B Cells. Science 209:1366-1369 (1980). View PDF »
  • Sakano, H., Maki, R., Kurosawa, Y., Roeder, W., and Tonegawa, S. Two types of somatic recombination are necessary for the generation of complete immunoglobulin heavy-chain genes. Nature 286:676-683 (1980). View PDF »
  • Maki, R., Traunecker, A., Sakano, H., Roeder, W., and Tonegawa, S. Exon shuffling generates an immunoglobulin heavy chain gene. PNAS 77:2138-2142 (1980). View PDF »
  • Perry, R.P., Kelley, D.E., Coleclough, C., Seidman, J.G., Leader, P., Tonegawa, S., Mattheyssens, G., and Weigert, M. Transcription of mouse k genes: Implications for allelic exclusion. PNAS 77:1937-1941 (1980). View PDF »


  • Sakano, H., Huppi, K., Heinrich, G., and Tonegawa, S. Sequences at the somatic recombination sites of immunoglobulin light-chain genes. Nature 280:288-294 (1979). View PDF »
  • Sakano, H., Rogers, J.H., Huppi, K., Brack, C., Traunecker, A., Maki, R., Wall, R., and Tonegawa, S. Domains and the hinge region of an immunoglobulin heavy chain are encoded in separate DNA segments. Nature 277:627-633 (1979). View PDF »


  • Tonegawa, S., Brack, C., Hirama, M., and Lenhard-Schuller, R. Somatic Recombination and Structure of an Immunoglobulin Gene. In: Cells of Immunoglobulin Synthesis Benevenuto Pernis & Henry J. Vogel (eds.), New York, Academic Press, (1978) pp.15-32. View PDF »
  • Bernard, O., Hozumi, N., and Tonegawa, S. Sequences of Mouse Immunoglobulin Light Chain Genes before and after Somatic Changes. Cell 15:1133-1144 (1978). View PDF »
  • Lenhard-Schuller, R., Hohn, B., Brack, C., Hirama, M., and Tonegawa, S. DNA clones containing mouse immunoglobulin k chain genes isolated by in vitro packaging into phage l coats. PNAS 75:4709-4713 (1978). View PDF »
  • Brack, C., Hirama, M., Lenhard-Schuller, R., and Tonegawa, S. A Complete Immunoglobulin Gene is Created by Somatic Recombination. Cell 15:1-14 (1978). View PDF »
  • Matthyssens, G., and Tonegawa, S. V and C parts of Immunoglobulin k-Chain Genes Are Separate in Myeloma. Nature 273:763-765 (1978). View PDF »
  • Hozumi, N., Brack, C., Pirrotta, V., Lenhard-Schuller, R., and Tonegawa, S. Characterization of a mouse DNA clone containing an Immunoglobulin variable region gene. Nucleic Acids Res. 5:1779-1799 (1978). View PDF »
  • Tonegawa, S., Maxam, A.M., Tizard, R., Bernard, O., and Gilbert, W. Sequence of a mouse germ-line gene for a variable region of an immunoglobulin light chain. PNAS 75:1485-1489 (1978). View PDF »
  • Tonegawa, S., Brack, C., Hozumi, N., and Pirrotta, V. Organization of Immunoglobulin Genes. Cold Spring Harbor Symp. Biol. 42:921-931 (1978). View PDF »


  • Tonegawa, S., Hozumi, N., Brack, C., and Schuller, R. Arrangement and Rearrangement of Immunoglobulin Genes. In: Regulatory Genetics of the Immune System, L. Herzenberg and C.F. Fox, (eds.), New York, Academic Press, (1977), vol. VI, pp. 43-55. View PDF »
  • Brack, C., and Tonegawa, S. Variable and constant parts of the immunoglobulin light chain gene of a mouse myeloma cell are 1250 nontranslated bases apart. PNAS 74:5652-5656 (1977). View PDF »
  • Tonegawa, S., Brack, C., Hozumi, N., and Schuller, R. Cloning of an immunoglobulin variable region gene from mouse embryo. PNAS 74:3518-3522 (1977). View PDF »
  • Tonegawa, S., Brack, C., Hozumi, N., Matthyssens, G., and Schuller, R. Dynamics of Immunoglobulin Genes. Immunological Reviews 36:73-94 (1977). View PDF »
  • Tonegawa, S., Hozumi, N., Matthyssens, G., and Schuller, R. Somatic Changes in the Content and Context of Immunoglobulin Genes. Cold Spring Harbor Symp. Quant. Biol. 41:877-889 (1977). View PDF »


  • Tonegawa, S., and Hozumi, N. Differentiation of Immunoglobulin Genes. The Immune System. 27 Colloquium Mosbach 1976, F. Melchers and K. Rajewsky (eds.). Berlin, Heidelberg, Springer-Verlag, (1976), pp. 86-97. View PDF »
  • Matthyssens, G., Hozumi, N., and Tonegawa, S. Somatic Generation of Antibody Diversity. Ann. Immunol. (Inst. Pasteur) 127C:439-448 (1976). View PDF »
  • Hozumi, N., and Tonegawa, S. Evidence for somatic rearrangement of immunoglobulin genes coding for variable and constant regions. PNAS 73:3628-3632 (1976). View PDF »
  • Tonegawa, S., and Steinberg, C. Too many chains- too few genes. In: The Generation of Antibody Diversity: A New Look. A. Cunningham, (ed.). New York, Academic Press (1976), pp. 175-182. View PDF »
  • Tonegawa, S. Reiteration frequency of immunoglobulin light chain genes: Further evidence for somatic generation of antibody diversity. PNAS 73:203-207 (1976). View PDF »


  • Tonegawa, S., Steinberg, C., Dube, S., and Bernardini, A. Evidence for Somatic Generation of Antibody Diversity. PNAS 71:4027-4031 (1974). View PDF »
  • Bernardini, A., and Tonegawa, S. Hybridization Studies with an Antibody Heavy Chain mRNA. FEBS Letters 41:73-77 (1974). View PDF »
  • Tonegawa, S., Bernardini, A., Weimann, B.J., and Steinberg, C. Reiteration Frequency of Antibody Genes. Studies with k-chain mRNA. FEBS Letters 40:92-96 (1974). View PDF »


  • Tonegawa, S., and Baldi, I. Electrophoretically Homogeneous Myeloma Light Chain mRNA and its Transcription in vitro. Biochim. Biophys. Res. Comm. 51:81-87 (1973). View PDF »


  • Tonegawa, S., Walter, G., and Dulbecco, R. Transcription of SV40 Genome in Transformed and Lytically Infected Cells. Le Petit Coloquia on Biology and Medicine-The Biology of Oncogenic Viruses 2:65-76 (1970). View PDF »
  • Tonegawa, S., Walter, G., Bernardini, A., and Dulbecco, R. Transcription of the SV40 Genome in Transformed Cells and during Lytic Infection. Cold Spring Harbor Symp. Quant. Biol. 35:823-831 (1970). View PDF »
  • Tonegawa, S., and Hayashi, M. Intermediates of Assembly of øX-174. J. Mol. Biol. 48:219-242 (1970). View PDF »


  • Tonegawa, S., and Hayashi, M. Genetic Transcription Directed by the b2 Region of l Bacteriophage. PNAS 61:1320-1327 (1968). View PDF »


  • Tonegawa, S., and Hayashi, M. The direction of in vitro genetic transcription on circular DNA. Biochim. Biophys. Acta. 123:634-637 (1966). View PDF »