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1 modality + + quality Skramlik + / ph ph ph,./..* Shallenberger Acree, +, taster supertaster non-taster * ** * 20* 2/ ** -*/ 2/

2 J. Japan Association on Odor Environment Vol. -1 No. 0,**0 399 phenylthiourea :PTU 0ono 0-npropylthiouracil :PROP PTU +2 - PTU PROP.,, + +.* 1* - + / /* 1* +/ -* / +/ + : 0,**/ Deg ENaC Degridin G 0 T + R, T,R T +R, T +R - T +R + T +R - T,R s G,, X, - + -, + 1 2

3 , NaCl N, HCl H, sucrose S, quinine Q, sucrose, NaCl, HCl, quinine O Sato et al. +31/ J Gen Physiol. 00 : * Rockefeller Univ P Fig. 1 nucleus tractus solitarius : NTS VPMpc : nucleus ventralis posteromedialis, pars parvicellularis primary somatosensory cortex frontal operculum insula area G G * PrCO : precentral operculum OFO : orbitofrontal operculum OFC : orbitofrontal cortex +* OFA : orbitofrontal area.. + /, blackcurrant juice : Scott ++.A Pritchard +, Lenz +- - ai, ;as, ; ipc, ;p, ;ss, :.2,**.., Rolls Scott Miyaoka Pritchard +1 Ogawa +2, +3,*., + Rolls +., +/

4 J. Japan Association on Odor Environment Vol. -1 No. 0,** area G BJ : blackcurrent juice, G : glucose, W:, +331 G sensory specific satiety Ogawa +2 G - *, +, * - G.B S Hirata Ogawa,+ G -

5 ,, - GO NOGO *.,, G +, - - +, G,,., -,- LED LED GO, / GO,. / / GO NOGO, cue A, ;B, peristimulus time histogram PSTH Ifuku et al,**- Neuroscience Research.1 : / Elsevier,. Bornstein,/.- * / Penfield,0

6 J. Japan Association on Odor Environment Vol. -1 No. 0,**0 403 / + / + + Kobayakawa,1 MEG : Magnetoencephalography +M 1* 0A.- 0B - mm,.* 0A 0B G,2 0 A B 0. Kobayakawa et al Neuroscience Letters,+, : +// +/2 Elsevier Fig. +, Fig...

404-1 0 +2 PET : positron emission tomography fmri : functional Magnetic Resonance Imaging,3 -+.1* G -,, MEG G.- fmri G.- -- 1 G.- P + +,0.1 ms N +,0,.

7 PET : positron emission tomography fmri : functional Magnetic Resonance Imaging,3 -+.1* G -,, MEG G.- fmri G G.- P + +,0.1 ms N +,0,.1 ms P,.-+.0 ms * 0 G G -.,1, -,, -- -/.- Cerf-Ducastel -0.- / +, 1 fmri A, ;B, ;a, ;b, ;c, Ogawa et al.,**/ Chemical Senses -* : /2- /3,. Oxford U P Fig. 1

8 J. Japan Association on Odor Environment Vol. -1 No. 0,** MEG fmri Onoda.* MEG G - fmri.+ /, -*.,.-.-,.../,.0.1 : + areag G : Benjamin +302 Sanides G, - : Brodmann +3*3 - +, - - +, : - +,. GO NOGO : GO NOGO, - *.+ M GO *.- M *./ ml GO LED LED GO NOGO /.- : precentral sulcus intraparietal sulcus. - 0 P +, P, N +, N,. P, N + von Skramlik, E. : Handbach der Physiologie der niederen Sinne, Georg Thieme, Leipzig, (+3,0)., Shallenberger, R.S. and Acree, T.E. : Molecular theory of sweet taste. Nature,,+0 :.2*.2,, (+301). - : p /2 2, Bartoshuk, L.M., Du#y, V.B. and Miller, I.J. Jr. : PTC/ PROP tasting : anatomy, psychophysics, and sex e#ects, Physiol. Behav., /0, ++0/ ++1+, (+33.). / : p Hoon, M.A., Adler, E., Lindemeier, J., Battey, J.F., Ryba, N.J. and Zuker, C.S. : Putative mammalian taste receptors : a class of taste-specific gpcrs with distinct topographic selectivity, Cell, 30, /.+ //+, (+333). 1 Sato, M., Ogawa, H. and Yamashita, S. : Response properties of macaque monkey chorda tympani fibers, J. Gen. Physiol., 00, *, (+31/).

9 Hellekant, G. and Ninomiya, Y. : Umami taste in the chimpanzee (Pan troglodytes) and rhesus monkey (Macaca mulatta), in Olfaction and Taste XI, p -0/ -02, Spinger-Verlag, Tokyo, (+33.). 3 Sanides, F. : The architecture of the cortical taste nerve areas in squirrel monkey (Saimiri sciureus) and their relationships to insular, sensorimotor and prefrontal regions, Brain Res. 2, 31 +,., (+303). +* :,+ +* p -30.*2,*** ++ Scott, T.R., Yaxley, S., Sienkiewicz, Z.J. and Rolls, E.T. : Gustatory responses in the nucleus tractus solitarius of the alert cynomolgus monkey, J. Neurophysiol., //, +2,,**, (+320). +, Pritchard, T.C., Hamilton, R.B. and Norgren, R. : Neural coding of gustatory information in the thalamus of Macaca mulatta. J. Neurophysiol., 0+, + +., (+323). +- Lenz, F.A., Gracely, R.H., Zir, T.A., Leopold, D.A., Rowland, L.H. and Dougherty, P.M. : Human thalamic nucleus mediating taste and multiple other sensations related to ingestive behavior, J. Neurophysiol., 11, -.*0 -.*3, (+331). +. Rolls, E.T. : Information processing in the taste system of primates, J. Exp. Biol., +.0, , (+323). +/ Rolls, E.T. : Taste and related systems in primates including humans, Chem. Senses, -* (Suppl +) : i10-i11, (,**/). +0 Scott, T.R. and Plata-Salaman, C.R. : Taste in monkey cortex, Physiol. Behav., 01,.23 /++, (+333). +1 Miyaoka, Y. and Pritchard, T.C. : Responses of primate cortical neurons to unitary and binary taste stimuli. J. Neurophysiol., 1/, , (+330). +2 Ogawa, H. : Gustatory cortex in primates. Anatomy and physiology, Neurosci. Res.,,*, + +-, (+33.). +3 Ogawa, H., Ifuku, H., Nakamura, T. and Hirata, S. : Possible changes in information from the primary to higher-order gustatory cortices studied by recording neural activities during a taste discrimination GO/ NOGO task in monkeys, Chem. Senses, -* (Suppl +), i12 13, (,**/).,* Ito, S. and Ogawa, H. : Neural activities in the frontoopercular cortex of macaque monkeys during tasting and mastication, Jpn. J. Physiol.,.., +.+ +/0, (+33.).,+ Hirata, S., Nakamura, T., Ifuku, H. and Ogawa, H. : Gustatory coding in the precentral extension of area - in Japanese macaque monkeys ; comparison with area G, Exp. Brain Res., +0-,.-/..0, (,**/).,, Ito, S., Ohgushi, M., Ifuku, H. and Ogawa, H. : Neuronal activity in the monkey fronto-opercular and adjacent insular/prefrontal cortices during a taste discrimination GO/NOGO task : response to cues, Neurosci. Res.,.,,,/1,00, (,**+).,- Ifuku, H., Ohgushi, M., Ito. S. and Ogawa, H. : Neurons associated with behavioral context errors in the primary and higher order gustatory cortex in the monkey, Neurosci. Lett., -+3, +,+ +,-, (,**,).,. Ifuku, H., Hirata, S., Nakamura, T. and Ogawa, H. : Neuronal activities in the monkey primary and higher order gustatory cortices during a taste discrimination delayed GO/NOGO task and after reversal, Neurosci. Res.,.1, /, (,**-).,/ Bornstein, W.S. : Cortical representation of taste in man and monkey. II. The localization of the cortical taste area in man and a method of measuring impairment of taste in man, Yale J. Biol. Med., +-, +-- +/0, (+3.*).,0 Penfield, W. and Jasper, H. : Epilepsy and the functional anatomy of the brain. Little Brown, Roston, MA, (+3/.).,1 Kobayakawa, T., Endo, H., Ayabe-Kanamura, S., Kumagai, T., Yamaguchi, Y., Kikuchi, Y., Tateda, T., Saito, S. and Ogawa, H. : The primary gustatory area in human cerebral cortex studied by magnetoencephalography, Neurosci. Lett.,,+,, +// +/2, (+330).,2 Saito, S., Endo, H., Kobayakawa, T., Ayabe-Kanamura, S., Kikuchi, Y., Takeda, T. and Ogawa, H. ; Temporal process from receptors to higher brain in taste detection studied by gustatory-evoked magnetic fields and reaction times, in Olfaction and Taste XII, Ann. N.Y. Acad. Sci., 2//,.*-.31, (+332).,3 Kinomura, S., Kawashima. R., Yamada, K., Ono, S., Itoh, M., Yoshida, S., Yamauchi, T., Matsui, H., Miyazawa, H., Itoh, H., Goto, R., Fujiwara, T., Satoh, K. and Fukuda, H. : Functional anatomy of taste perception in the human brain studied with positron emission tomography, Brain Res., 0/3,,0-,00, (+33.). -* Small, D.M., Jones-Gotman, M., Zatorre, R.J., Petrides, M. and Evans, A.C. : A role for the right anterior temporal lobe in taste quality recognition, J. Neurosci., +1, /+-3 /+.,, (+331). -+ Faurion, A., Cerf, B., LeBihan, D. and Pillias, A.M. : fmri study of taste cortical areas in humans, in Olfaction and Taste XII, Ann. N.Y. Acad. Sci., 2//, /-/ /./, (+332). -, Kobayakawa, T., Ogawa, H., Kaneda, H., Ayabe- Kanamura, S., Endo, H. and Saito, S. : Spatio-temporal analysis of cortical activities evoked by gustatory stimulation in humans, Chem. Senses,,.,,*+,*3, (+333). -- Ogawa, H., Wakita, M., Hasegawa, K., Kobayakawa, T., Sakai, N., Hirai, T., Yamashita, Y. and Saito, S. : Functional MRI detection of activation in the primary gustatory cortices in human, Chem. Senses, -*, /2- /3,, (,**/). -. Mizoguchi, C., Kobayakawa, T., Saito, S. and Ogawa, H. : Gustatory evoked cortical activity in humans studied by simultaneous EEG and MEG recording, Chem. Senses,,1, 0,3 0-., (,**,).

10 J. Japan Association on Odor Environment Vol. -1 No. 0,** / Komiyama, A., Kobayakawa, T., Goto, N., Saito, S. and Ikeda, M. : Trigeminal nerve response in human for - M NaCl solutions, studied by MEG, in Abstracts for Annual Meeting of the AChmS XXVI, Sarasota, FL, p 0/, (,**/). -0 Cerf-Ducastel, B., Van de Moortele, P.F., MacLeod, P., LeBihan, D. and Faurion, A. : Interaction of gustatory and lingual somatosensory perception, at the cortical level in the human : a functional magnetic resonance imaging study, Chem. Senses,,0, , (,**+). -1 Motta, G. : The cortical taste centers, Bull. Sci. Med. (Bologna), +-+,.2*.3-, (+3/3). -2 Onoda, K. and Ikeda, M. : Gustatory disturbance due to cerebrovascular disorder, Laryngoscope, +*3, +,- +,2, (+333). -3 Pritchard, T.C., Macculuso, D.A. and Eslinger, P.J. : Taste perception in patients with insular cortex lesions, Behav. Neurosci., ++-, , (+333)..* Onoda, K., Kobayakawa, T., Ikeda, M., Saito, S. and Kida, A. : Laterality of human primary gustatory cortex studied by MEG, Chem. Senses, -*, 0/1 000, (,**/)..+ : laterality, 02 1/,**0., O Doherty, J., Rolls, E.T., Francis, S., Bowtell, R. and McGlone, F. : Representation of pleasant and aversive taste in the human brain, J. Neurophysiol., 2/, +-+/ +-,+, (,**+)..- Small, D.M., Gregory, M.D., Mak, Y.E., Gitelman, D., Mesulam, M.M. and Parrish, T. : Dissociation of neuronal representation of intensity and a#ective valuation in human gustation, Neuron, -3, 1*+ 1++, (,**-)... Zald, D.H., Hagen, M.C. and Pardo, J.V. : Neural correlates of tasting concentrated quinine and sugar solutions, J. Neurophysiol., 21, +*02 +*1/, (,**,)../ O Doherty, J., Deichmann, R., Critchley, H.D. and Dolan, R. J. : Neural responses during anticipation of a primary taste reward, Neuron, --, 2+/ 2,0, (,**,)..0 Kringelbach, M.L., de Araujo, I.E. and Rolls, E.T. : Taste-related activity in the human dorsolateral prefrontal cortex, Neuroimage,,+, , (,**.)..1 Small, D.M., Zatorre, R.J., Dagher, A., Evans, A.C. and Jones-Gotman, M. Changes in brain activity related to eating chocolate : from pleasure to aversion. Brain, +,., +1,* +1--, (,**+). Brain mechanism of taste sensation Hisashi OGAWA*, Tatsu KOBAYAKAWA** * Dept. of Neurol., Kumamoto Kinoh Hospital (Professsor Emeritus in Kumamoto University) Yamamuro, Kumamoto City 20* 2/+2 ** Institute for Human Science and Biomedical Engineering, National Institute of Advanced Industrial Science and Technology Higashi, Tsukuba City, Ibaragi Prefecture, -*/ 2/00 Abstract Since the nose and mouth are situated near to each other on the face, olfaction and gustation are generally thought to have close positions in the nervous system to interact with each other. However, neural information about both sensations are not integrated until they reach association cortex. The present review first described the general nature of gustation in comparison with olfaction, and then stated how gustatory information is analyzed or integrated along the central pathway to the gustatory cortex. Further, it described the position of the primary gustatory cortex in humans recently revealed by non-invasive brain imaging and the function of gustation-related cortex, including the primary gustatory cortex. Key words : taste receptive mechanism, central gustatory pathway, cortical taste neurons, noninvasive brain imaging, primates

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