sc.January1.1997.170359-36 REFERENCES MP.Schmitt MA (1980)The apl roke ir “expressive”aphasia syndrome(Mohr Other activated areas e of pure rd-deafness with autopsy.J Ner :Raichle may play a gnera l role in neur computa imaging of human orted to sho l35662-62 tems dem A major ion during the semanti on task the leou parahip 994.an n of lan This conne sugge men 。 ith eral region reno cause a verbal et al,1987 and Warring 1991).Retro encoding of the se Hum Brain Mapp 3:93-106. .p that ni G, manti is known to enhan storage of e pisodic memories compared dic ceptua 28 leve ted during task relati task MR function (von Cramon et al.1988:Zola oustic-phonctic processing deficits eurolog n task relative to the on task and.therefore vas not dered CM (192) of the a (We 1874 0,1965 ton EK (198 1995).One hypothesis that would re econcile these v arious findings 0 of 1971:C Craik FIM. ea h Be 1157- work for 11 al,1981:Paulesu et al,1993).The tone ision task Holt R Reld the task,which involved Price Cl.Wele ormal subjects.Studiest rus activation to the right hemisphere is con sistent with data ving right hemispher on lesions usually manifest global aphasia and significant comprehen￾sion disturbances in the acute period after injury, only later evolving into the typical “expressive” aphasia syndrome (Mohr, 1976; Mohr et al., 1978). Other activated areas Functional activation of the right cerebellum during word gener￾ation tasks is a consistent finding in PET studies that include the cerebellum (Petersen et al., 1989; Pardo and Fox, 1993; Raichle et al., 1994). The present data extend this observation to a semantic decision task that does not require word retrieval. The cerebellum may play a general role in facilitating complex neural computa￾tions (Keele and Ivry, 1990; Leiner et al., 1991), although the precise nature of this role remains somewhat speculative. Several patients were reported to show cognitive deficits in association with cerebellar damage (Bracke-Tolkmitt et al., 1989; Fiez et al., 1992), yet frank aphasic disturbances are rare. A major activation during the semantic decision task oc￾curred near the splenium of the corpus callosum. Much of this region probably coincides with retrosplenial cortex (Vogt, 1976), which has connections with hippocampus, parahip￾pocampus (Mufson and Pandya, 1984; Suzuki and Amaral, 1994), and anterolaterodorsal thalamus (Sripanidkulchai and Wyss, 1986). This connectivity pattern suggests an involvement in memory functions, and left hemisphere lesions in this gen￾eral region reportedly cause a verbal amnestic syndrome (Valenstein et al., 1987; Rudge and Warrington, 1991). Retro￾splenial activation may therefore be related to memory￾encoding processes that accompanied performance of the se￾mantic decision task. Although identification of memory systems was not an intended goal, processing at a semantic level is known to enhance storage of episodic memories compared with processing at a perceptual level (Craik and Lockhart, 1972). Thus, the episodic memory encoding system was likely activated during the semantic task relative to the tone task. This interpretation could also account for the activation observed in left parahippocampus, another structure closely tied to memory function (von Cramon et al., 1988; Zola￾Morgan et al., 1989). The supramarginal gyrus was activated bilaterally by the tone decision task relative to the semantic decision task and, therefore, was not considered to be a language area. This finding deserves comment, because the left supramarginal gyrus is usually consid￾ered part of the perisylvian language “core” (Wernicke, 1874; Marie, 1917; Penfield and Roberts, 1959; Geschwind, 1965; Ben￾son, 1985; Mayeux and Kandel, 1985). Lesions in or near this structure cause speech output disturbances characterized by pho￾nemic paraphasias in repetition (Geschwind, 1965; Damasio and Damasio, 1980) and other phonological deficits (Caplan et al., 1995). One hypothesis that would reconcile these various findings is that the supramarginal gyrus may be involved in short-term storage of auditory information (Warrington et al., 1971; Car￾amazza et al., 1981; Paulesu et al., 1993). The tone decision task, which involved monitoring arbitrary sequences of up to seven tones, probably placed more demand on short-term auditory memory resources than did the semantic task, which involved familiar word stimuli. Relative lateralization of supramarginal gyrus activation to the right hemisphere is consistent with data showing right hemisphere dominance for short-term storage of nonlinguistic auditory information (Zatorre and Samson, 1991; Zatorre et al., 1994). REFERENCES Alexander MP, Schmitt MA (1980) The aphasia syndrome of stroke in the left anterior cerebral artery territory. Arch Neurol 37:97–100. Alexander MP, Hiltbrunner B, Fischer RS (1989) Distributed anatomy of transcortical sensory aphasia. Arch Neurol 46:885– 892. Barrett AM (1910) A case of pure word-deafness with autopsy. J Nerv Ment Dis 37:73–92. Benson DF (1985) Aphasia and related disorders: a clinical approach. In: Principles of behavioral neurology (Mesulam M-M, ed), pp 193–238. Philadelphia: Davis. Binder JR, Rao SM, Hammeke TA, Frost JA, Bandettini PA, Hyde JS (1994a) Effects of stimulus rate on signal response during functional magnetic resonance imaging of auditory cortex. Cognit Brain Res 2:31–38. Binder JR, Rao SM, Hammeke TA, Yetkin YZ, Jesmanowicz A, Bandet￾tini PA, Wong EC, Estkowski LD, Goldstein MD, Haughton VM, Hyde JS (1994b) Functional magnetic resonance imaging of human auditory cortex. Ann Neurol 35:662– 672. Binder JR, Rao SM, Hammeke TA, Frost JA, Bandettini PA, Jesmanow￾icz A, Hyde JS (1995) Lateralized human brain language systems dem￾onstrated by task subtraction functional magnetic resonance imaging. Arch Neurol 52:593– 601. Binder JR, Frost JA, Hammeke TA, Rao SM, Cox RW (1996a) Function of the left planum temporale in auditory and linguistic processing. Brain 119:1239 –1247. Binder JR, Swanson SJ, Hammeke TA, Morris GL, Mueller WM, Fischer M, Benbadis S, Frost JA, Rao SM, Haughton VM (1996b) Determi￾nation of language dominance using functional MRI: a comparison with the Wada test. Neurology 46:978 –984. Bogen JE, Bogen GM (1976) Wernicke’s region - where is it? Ann NY Acad Sci 290:834 – 843. Bookheimer SY, Zeffiro TA, Blaxton T, Gaillard T, Theodore W (1995) Regional cerebral blood flow during object naming and word reading. Hum Brain Mapp 3:93–106. Bottini G, Corcoran R, Sterzi R, Paulesu E, Schenone P, Scarpa P, Frackowiak RSJ (1994) The role of the right hemisphere in the inter￾pretation of figurative aspects of language. A positron emission tomog￾raphy activation study. Brain 117:1241–1253. Bracke-Tolkmitt R, Linden A, Canavan AGM, Rockstroh B, Scholz E, Wessel K, Diener H-C (1989) The cerebellum contributes to mental skills. Behav Neurosci 103:442– 446. Broca P (1861) Remarques sur le sie`ge de la faculte´ du langage articule´; suivies d’une observation d’aphemie. Bull Soc Anat Paris 6:330 –357. Caplan D, Gow D, Makris N (1995) Analysis of lesions by MRI in stroke patients with acoustic-phonetic processing deficits. Neurology 45:293–298. Caramazza A, Basili A, Koller J, Berndt R (1981) An investigation of repetition and language processing in a case of conduction aphasia. Brain Lang 14:235–271. Carr TH, McCauley C, Sperber RD, Parmalee CM (1982) Words, pic￾tures, and priming: on semantic activation, conscious identification, and the automaticity of information processing. J Exp Psychol Hum Percept Perform 8:757–777. Cohen JD, Forman SD, Braver TS, Casey BJ, Servan-Schreiber D, Noll DC (1994) Activation of the prefrontal cortex in a nonspatial working memory task with functional MRI. Hum Brain Mapp 1:293–304. Costello AL, Warrington EK (1989) Dynamic aphasia: the selective im￾pairment of verbal planning. Cortex 25:103–114. Cox RW (1996) AFNI: software for analysis and visualization of func￾tional magnetic resonance neuroimages. Comput Biomed Res 29:162–173. Craik FIM, Lockhart RS (1972) Levels of processing: a framework for memory research. J Verb Learn Verb Behav 11:671– 684. Crocker L, Algina J (1986) Introduction to classical and modern test theory. New York: Holt, Rinehart and Winston. Warburton E, Wise RJS, Price CJ, Weiller C, Hadar U, Ramsay S, Frackowiak RSJ (1996) Noun and verb retrieval by normal subjects. Studies with PET. Brain 119:159 –179. Damasio AR, Tranel D (1993) Nouns and verbs are retrieved with dif￾ferently distributed neural systems. Proc Natl Acad Sci USA 90:4957– 4960. Damasio H (1981) Cerebral localization of the aphasias. In: Acquired aphasia (Sarno MT, ed), pp 27–50. Orlando, FL: Academic. 360 J. Neurosci., January 1, 1997, 17(1):353–362 Binder et al. • FMRI of Language Areas