Episodic encoding and recognition of pictures and words: role of the human medial temporal lobes

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1 Acta Psychologica 105 (2000) 159±179 Episodic encoding and recognition of pictures and words: role of the human medial temporal lobes Stefan Kohler a,*, Morris Moscovitch a,b,c, Gordon Winocur a,d, Anthony R. McIntosh a,c a Rotman Research Institute of Baycrest Centre for Geriatric Care, University of Toronto, Toronto, Canada b Department of Psychology, Baycrest Centre for Geriatric Care, North York, Canada c Department of Psychology, University of Toronto, Toronto, Canada d Department of Psychology, Trent University, Peterborough, Canada Abstract In the present PET study, we examined brain activity related to processing of pictures and printed words in episodic memory. Our goal was to determine how the perceptual format of objects (verbal versus pictorial) is re ected in the neural organization of episodic memory for common objects. We investigated this issue in relation to encoding and recognition with a particular focus on medial temporal-lobe (MTL) structures. At encoding, participants saw pictures of objects or their written names and were asked to make semantic judgments. At recognition, participants made yes±no recognition judgments in four di erent conditions. In two conditions, target items were pictures of objects; these objects had originally been encoded either in picture or in word format. In two other conditions, target items were words; they also denoted objects originally encoded either as pictures or as words. Our data show that right MTL structures are di erentially involved in picture processing during encoding and recognition. A posterior MTL region showed higher activation in response to the presentation of pictures than of words across all conditions. During encoding, this region may be involved in setting up a representation of the perceptual information that comprises the picture. At recognition, it may play a role in guiding retrieval processes based on the perceptual input, i.e. the retrieval cue. Another more anterior right MTL region was found to be di erentially involved in recognition of objects that had been encoded as pictures, irrespective of whether the retrieval cue provided was pictorial or verbal in nature; this region may be involved in accessing * Corresponding author. Present address: University of Western Ontario, Department of Psychology, London, Ontario, Canada N6A 5C2. Tel.: ext ; fax: address: stefank@julian.uwo.ca (S. KoÈhler) /00/$ - see front matter Ó 2000 Elsevier Science B.V. All rights reserved. PII: S (00)

2 160 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 stored pictorial representations. Our results suggest that left MTL structures contribute to picture processing only during encoding. Some regions in the left MTL showed an involvement in semantic encoding that was picture speci c; others showed a task-speci c involvement across pictures and words. Together, our results provide evidence that the involvement of some but not all MTL regions in episodic encoding and recognition is format speci c. Ó 2000 Elsevier Science B.V. All rights reserved. PsycINFO classi cation: 2343; 2540 Keywords: Hippocampus; Parahippocampal gyrus; PET; Functional neuroimaging; Visual; Memory It is a widely accepted notion in cognitive psychology that representations in episodic memory, i.e. memory for events that are distinct in time and space, are primarily based on the meaning of our experiences rather than on their sensory and perceptual characteristics. Episodic long-term memory is thought to be conceptually driven and very e cient at preserving the semantic gist of past experiences. However, there is clear evidence from research in cognitive psychology showing that some perceptual aspects of ongoing experiences are represented in episodic memory (Madigan, 1983; Mandler & Johnson, 1976; Nelson, Reed, & Walling, 1976). Common objects are unique in that they can be perceived and identi ed in the visual modality in two di erent perceptual formats: as words and as pictures. By comparing memory processing of pictures of objects with processing of their names, it is possible to examine whether the perceptual format of objects is re ected in the functional organization of episodic memory in the human brain. In the present study, we asked whether there are brain regions, in particular in the medial temporal lobe (MTL), whose involvement in episodic memory for common objects changes depending on whether the objects are encountered as two-dimensional pictures or as printed words. We compared picture and word processing at encoding and at retrieval in order to determine whether there are format-speci c e ects that may be associated with distinct neural correlates at each stage of processing. There is evidence from lesion studies in patients who underwent unilateral anterior temporal lobectomy that speaks to these issues. It is well established that leftsided temporal-lobe lesions that encroach on medial temporal structures result in episodic memory de cits for verbal material, especially when a delay intervenes the initial encoding and subsequent memory retrieval (Milner, 1958; Smith, 1989). In line with this general nding, several studies have demonstrated impairments in recall of the names of pictures and of the names of visually encountered three-dimensional objects after left-sided temporal lobectomies (Incisa della Rocchetta, 1986; Jaccarino, 1975; Smith & Milner, 1981). However, de cits in memory for pictures of objects have also been reported in patients who underwent right temporal lobectomies (Jaccarino, 1975; Nunn, Graydon, Polkey, & Morris, 1999; Pigott & Milner, 1993). Most notably, Jaccarino found de cits after right-sided lesions for delayed but not immediate free recall of the names of previously studied line drawings of objects.

3 S. Kohler et al. / Acta Psychologica 105 (2000) 159± Given that right-sided temporal-lobe lesions usually result in material-speci c memory de cits for information that is di cult to verbalize (Milner, 1968; Smith, 1989), this nding suggests that right temporal-lobe structures may play a distinct role in retention of visual characteristics of pictorially encoded objects. Moreover, it hints that this information is accessed even when no pictorial cues are present at the time of retrieval. Previous functional imaging ndings that address whether speci c brain structures show a di erential regional cerebral blood ow (rcbf) response to processing of pictures and words in healthy individuals come from a positron emission tomography (PET) study on neural correlates of episodic encoding of visually presented objects (Grady, McIntosh, Rajah, & Craik, 1998). Grady et al. presented pictures or names of objects under three di erent encoding instructions during scanning. Across the di erent instruction conditions, encoding of pictures as compared to words resulted in bilateral activation of medial temporal cortices that was more pronounced in the right hemisphere. The reverse comparison did not reveal activation in MTL regions. Two additional studies have con rmed the involvement of medial temporal-lobe structures in picture encoding (Kelley et al., 1998; Martin, Wiggs, & Weisberg, 1997). In these studies, only right-sided structures showed higher activation during picture than word encoding. It is noteworthy that the neuroimaging studies discussed this far (Grady et al., 1998; Kelley et al., 1998; Martin et al., 1997) have only focussed on MTL activation in relation to encoding of pictures and words. Given that no retrieval conditions were investigated, these studies do not address whether the format-speci c activation reported in MTL regions is speci cally tied to encoding processes or whether it can also be observed in relation to episodic retrieval. Evidence that speaks to whether memory processes involved in retrieval of pictures and words are also supported by distinct brain regions comes from a PET study by Buckner, Raichle, Miezin, and Petersen (1996). These investigators examined rcbf while subjects retrieved the names of pictorially encoded objects or of auditorily presented names of objects in a paired-associate recall task. The associates presented as cues during retrieval (and during encoding before scanning) were of the same type in both conditions, namely auditorily presented words. A direct comparison of both recall tasks revealed only subtle di erences in rcbf whose reliability could not be con rmed. These results led Buckner et al. to emphasize the commonalities in brain activity between retrieval of objects encoded as pictures and those encoded as words. It remains to be determined, however, whether the di erences between picture and word retrieval are similarly subtle for retrieval tasks other than cued recall. In the present PET investigation, we examined the neural correlates of episodic memory for pictures and words associated with encoding and those associated with retrieval within the same study. Our goal was to determine the overlap in formatspeci c activation at encoding and retrieval in order to see whether there are any format-related patterns of activation that are speci cally tied to encoding or retrieval processes. We used a recognition memory paradigm and a design in which the format of objects presented at encoding was crossed completely with the format of

4 162 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 the stimulus cues presented at recognition. This allowed us to compare formatspeci c e ects at both stages of memory processing under conditions of equivalent visual stimulation. In addition, our design allowed us to separate retrieval e ects that depend on the format of the recognition cue from retrieval e ects that depend on the format in which the objects were originally encoded. Given the prominent role that the MTLs' have played in the literature on format speci city in episodic memory, special emphasis will be placed on the examination and discussion of task-related changes in rcbf in this region. For the present analyses, the MTLs' were de ned broadly as including the hippocampal formation, the amygdala, as well as surrounding cortex in the uncus and parahippocampal gyrus in their entire rostro-caudal extent (Amaral, 1999). 1. Methods 1.1. Overview Participants underwent PET scanning in 6 di erent experimental conditions with one scan per condition. The rst two conditions required semantic encoding of objects that were presented either as words (EW) or as pictures (EP). Subsequently, participants performed four di erent yes/no-recognition memory tasks for the objects encountered during encoding. These tasks required: (i) recognition of objects presented as pictures at recognition and at encoding (RPEP), (ii) recognition of objects presented as pictures at recognition and as words at encoding (RPEW), (iii) recognition of objects presented as words at recognition and at encoding (RWEW), or (iv) recognition of objects presented as words at recognition and as pictures at encoding (RWEP) Participants Twelve young right-handed individuals participated in the PET study (7 male, 5 female; mean age ˆ 25:3, S:D: ˆ 3:2). They were reimbursed for their participation. Participants were screened to ensure the absence of any medical, neurological, or psychiatric disorder. The study was approved by the Human Subjects Use Committee of Baycrest Centre. All participants gave written informed consent Material The stimuli were representational line drawings of common objects and their verbal labels. A sample of 198 target items and 16 practice items was selected from a set of common objects, for which line drawings, corresponding picture norms, and word-frequency norms were available (Snodgrass & Vanderwart, 1980). Only objects that satis ed the following two criteria were selected: (i) the name agreement for the line drawing was larger than 70%, and (ii) the word length of the object name was between 3 and 12 letters. The 198 target objects were assigned to 18 lists

5 in such a way that the average name agreement, picture width, picture luminance, word length, and word frequency were equivalent for all lists. The assignment of lists to the di erent experimental conditions was counterbalanced across participants. Every participant encountered 16 lists of 11 items during the experiment; the remaining 2 lists were used for counterbalancing purposes only. For each participant, 12 lists of items provided the targets that were presented at encoding (66 items in EP; 66 items in EW); four additional lists provided the lures during recognition. Half of the items presented during EP served as targets during RPEP; the other half served as targets during RWEP. Similarly, half of the items presented during EW served as targets during RPEW; the other half served as targets during RWEW Behavioral procedure S. Kohler et al. / Acta Psychologica 105 (2000) 159± In each of the six experimental conditions, participants were presented with objects (words or pictures) that were centered in the middle of a computer screen at a viewing distance of approximately 1m. The font size for words was chosen such that the average subtended visual angle for each list of items (i.e. number of pixels covered horizontally) was equivalent for words and pictures. Each item appeared for 2.5 s and was followed by a blank screen during an inter-stimulus interval of 0.5 s. For all tasks, participants reported their responses by pressing one of the two buttons of a computer mouse with their right hand. The number of items presented within the actual scan-acquisition period was identical (20 items) for all tasks. Prior to the actual scanning session, participants were familiarized with all tasks of the study with practice trials. Experimental testing under scanning always started with the two encoding tasks and was followed by the four recognition-memory tasks. This setup made it possible to test memory in the recognition memory scans for information that was previously encoded under scanning. The presentation order of the two encoding tasks was counterbalanced across participants, as was the order of the four recognition memory tasks. In the two encoding tasks, participants made semantic judgments on the 66 objects presented in random order; they were instructed to decide for each item whether it was a living or a non-living object. In the four recognition-memory tasks, participants made yes/no recognition judgments on the 44 objects presented. Thirtythree of these objects had previously been shown and 11 objects were novel lures. The presentation order of objects was pseudo-random with the constraint that the middle set of 28 items, including the 20 items to be presented during the scan-acquisition period, consisted of previously encountered (`old') items only. Participants were instructed to decide for each object whether they had seen it in the encoding phase. They were informed that they might have encountered the objects previously as words or as pictures and that they should make their decision irrespective of whether the object format was the same as during encoding. They were also informed that the majority of items in each condition were targets.

6 164 S. Kohler et al. / Acta Psychologica 105 (2000) 159± PET scanning Participants were scanned in a GEMS-PC B scanner with a custom- tted thermoplastic face-mask, which permitted unobstructed viewing of the computer screen on which the stimulus displays were presented. For the purpose of the present experiment, every participant underwent six scans. Each scan was acquired after a bolus-injection of 40 mci (1.48 GBq) of 15 OŠ±H 2 O had been injected into the left forearm vein through an indwelling catheter. The cognitive task was started approximately 30 s prior to the 60 s data acquisition period for each scan. All scans were performed 11 min apart. The PET images were attenuation-corrected using a transmission scan acquired prior to the rst PET scan. The scans were reconstructed using a Hanning lter with a cuto frequency of 0.5 Hz. Normalized integrated regional counts were used as an index of rcbf Image analysis Pre-processing of images For each participant, the reconstructed PET images were realigned to the rst image to correct for head movements using AIR software (Woods, Cherry, & Mazziotta, 1992). The realigned images from each subject were then transformed into standard stereotactic space (Talairach & Tournoux, 1988) using the Statistical Parametric Mapping software (SPM 95; Friston et al., 1995). They were subsequently smoothed using an isotropic Gaussian kernel with a full width at halfmaximum of 10 mm in all three dimensions. To correct for changes in global blood ow, the voxel values in the transformed images were expressed as a ratio of the average counts for all brain voxels within a scan Examination of experimental e ects Experimental e ects on rcbf were evaluated using a two-step procedure (Cabeza et al., 1997). In the rst step, we determined the most prominent di erences in rcbf between our tasks by examining activity in the entire brain with the method of partial least squares (PLS; McIntosh, Bookstein, Haxby, & Grady, 1996). The emphasis of PLS is on the covariances between brain images on one side and contrasts between tasks on the other. Based on a singular-value decomposition of these covariances, PLS serves to determine a set of variables (latent variables, LVs) that optimally relate patterns of change in rcbf across the brain to the strongest task contrasts. These LVs are extracted in order of the proportion of covariance they account for. The resulting patterns associated with each LV are mutually independent singular images, which re ect those brain regions that are most a ected by the corresponding experimental manipulation. Our speci c interest was to determine whether the singular images extracted with PLS had local maxima or minima in the MTLs'. In the second step of our analyses, we performed univariate statistical tests on the maxima and minima of the singular images extracted. These tests were employed to determine whether the rcbf changes observed in peak regions, in particular those in

7 S. Kohler et al. / Acta Psychologica 105 (2000) 159± the MTLs', could also be considered signi cant when examined locally (i.e. without taking rcbf changes in other brain regions into account) with the task contrasts from the PLS analysis. Complex e ects in MTL regions involving multiple conditions were followed up by additional univariate tests to allow for more speci c comparisons (e.g. picture versus word encoding) Assessment of statistical signi cance The statistical signi cance of the obtained PLS results was assessed using permutation tests and bootstrap estimation of standard errors. The permutation test served to determine whether the e ect represented in each singular image as a whole is su ciently strong in statistical terms (McIntosh et al., 1996). Results for this test were considered to be signi cant at P < 0:05. The bootstrap estimates of standard errors were computed to determine stable maxima and minima of the singular images (McIntosh, Lobaugh, Cabeza, Bookstein, & Houle, 1998). The estimation procedure involved randomly resampling subjects with replacement and computing the standard error for the saliences (i.e. image weights) of all voxels based on 100 bootstrap samples taken. Peak voxels with a ratio of salience/standard error P 2.0 were considered stable. The univariate tests were performed on all stable peaks identi ed with PLS that were part of a contiguous blob of at least 30 voxels. This blob size was chosen based on the degree of autocorrelation that can be expected when data are smoothed with a 10 mm Gaussian kernel. For each peak, the experimental e ect re ected in the corresponding PLS contrast was tested using a repeated-measures regression approach to ANOVA (McIntosh et al., 1998). The tables list those peaks of the singular images that were con rmed locally with the univariate tests at an uncorrected signi cance level of P < 0:001, except for speci c regions of interest that are reported at P < 0:01. Follow-up tests for the further characterization of signi cant complex e ects were performed with a signi cance criterion of P < 0: Results 2.1. Behavioral results of the PET study Response accuracy was P 0.70 in each of the recognition memory tasks under scanning, indicating that the brain activity being measured was associated with successful memory encoding and recognition in all tasks (Table 1). A two-way ANOVA revealed that the main e ect of recognition format was signi cant F 1; 11 ˆ5:89; P < 0:05 while the main e ect of encoding format was not (P > 0:10). The interaction between the two factors also reached statistical signi cance F 1; 11 ˆ11:43; P < 0:01. A planned pairwise comparison revealed the typical picture superiority e ect, which is most robustly observed in recognition memory when object format at recognition matches that at encoding (Madigan, 1983): Performance in RPEP was higher than performance in RWEW t 11 ˆ3:05; P < 0:05.

8 166 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 Table 1 Behavioral performance in the PET study: proportion of correct recognition decisions as a function of object format at encoding and recognition a Encoding format Recognition format Picture (RP) Word (RW) Picture (EP) (0.07) (0.07) Word (EW) (0.13) (0.09) a Note. Standard deviations are shown in parentheses Results of the image analysis The PLS analysis extracted 3 LVs that were clearly signi cant by permutation test (P ˆ 0:00; 0:00; 0:01 for LV 1, 2, 3, respectively) and one LV that revealed a signi cant trend (LV 4, P ˆ 0.08). LVs 2, 3, and 4 were found to re ect e ects of object format on rcbf and will be presented rst Format-related e ects comparable at encoding and recognition The singular image of the second LV from the PLS analysis identi ed those brain regions that demonstrated a di erential rcbf response to pictures and words that was comparable at encoding and recognition (see Fig. 1(a)). At recognition, these regions responded to the format of the stimulus present, regardless of whether the tobe-retrieved objects had been encoded as pictures or words initially. Maxima and Fig. 1. Task-related e ects on rcbf that were extracted with the PLS analysis for the entire brain. Mean standardized brain scores are plotted against the di erent task conditions to illustrate the e ects re ected in LV 2 (plot a), LV 3 (plot b), LV 4 (plot c) and LV 1 (plot d).

9 S. Kohler et al. / Acta Psychologica 105 (2000) 159± Table 2 Maxima and minima of singular image 2 re ecting format-related e ects comparable at encoding and recognition a Region Hem BA x y z Ratio F F prob Maxima (picture processing > word processing) Middle/inferior frontal L 11 )16 58 ) gyrus (frontopolar) Orbital gyrus/gyrus L 47/11 )14 26 ) < rectus Posterior parahippocampal R )28 ) < gyrus Brainstem R 4 )32 ) < Inferior temporal/fusiform/lingual R )52 ) < gyrus R )86 ) < R )90 ) < L 18 )8 )92 ) < L 17/18 )16 )92 ) Lateral cerebellum R 20 )88 ) < L )24 )84 ) < Minima (word processing > picture processing) Posterior putamen L )26 ) Inferior parietal lobule L 40 )44 ) Cuneus/lingual gyrus R 17/18 8 ) L 19 )10 ) < a Note: BA: Brodmann Area; Hem: Hemisphere; Ratio: stability coe cient in bootstrap analysis; degrees of freedom for F ˆ 1, 55; coordinates of representative maxima and minima listed in millimeter according to Talairach and Tournoux (1988). minima of this image are listed in Table 2. Maxima, which re ect regions with higher rcbf during picture than word processing, were found predominantly in bilateral visual extrastriate cortex (lingual and fusiform gyrus) and posterior temporal cortex. Their spatial extent was larger in the right hemisphere and included a peak in the right posterior parahippocampal gyrus. This peak is shown in Fig. 2(a) together with its response pro le. Minima of singular image 2, which re ect regions with higher rcbf during word than picture processing, were found primarily in the left hemisphere. They included portions of inferior parietal lobule and medial cuneus but no MTL regions Format-related e ects speci c to encoding The singular image associated with LV 3 identi ed those brain regions that demonstrated an interaction between object format and task; they showed a di erential rcbf response to pictures and words at encoding that was not paralleled by a comparable response at recognition (see Fig. 1(b)). Maxima and minima of this image are listed in Table 3. Maxima re ect regions with a higher rcbf to picture encoding as compared to word encoding whereas minima re ect regions with the opposite type of response. Among the regions with a picture-speci c increase in

10 168 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 Fig. 2. MTL regions showing format-speci c e ects on rcbf. Sections from the singular images of the PLS analysis are superimposed on horizontal slices from an MRI template that conforms to the atlas space of Talairach and Tournoux (1988). Slices are taken at the Z-level speci ed in the coordinates XYZ for each peak. Only voxels with a bootstrap ratio P 2.0 are displayed. Regions with an rcbf increase during picture processing are coded in white (maxima); those with an increase during word processing are coded in gray (minima). Response pro les for these regions re ect the mean adjusted rcbf response and corresponding SEM for each condition. (a) A peak from LV 2 in right posterior parahippocampal gyrus exhibited an rcbf increase during picture as compared to word processing at encoding and recognition. At recognition, it responded to the format of the recognition cue. (b) A peak from LV 3 in left anterior parahippocampal gyrus showed an rcbf increase during picture as compared to word encoding with no comparable e ect at recognition. Note that there is a more posterior region (small arrow) that showed a weaker e ect in the same direction. (c) A peak from LV 4 in right anterior parahippocampal gyrus showed a picture-speci c response during recognition that was related to the format in which target information had been encoded.

11 S. Kohler et al. / Acta Psychologica 105 (2000) 159± Table 3 Maxima and minima of singular image 3 re ecting format-related e ects speci c to encoding a Region Hem BA x y z Ratio F F prob Maxima (picture encoding > word encoding) Anterior parahippocampal L 38 )28 10 ) gyrus/uncus Caudate nucleus L )8 ) < Middle temporal gyrus/ R )10 ) < inferior temporal sulcus Brainstem R 10 )20 ) < Posterior parahippocampal L 36 )28 )24 ) gyrus Fusiform gyrus/inferior R )58 ) temporal sulcus L 37 )40 )50 ) Middle occipital gyrus L 19 )50 )76 ) Minima (word encoding > picture encoding) Anterior cingulate gyrus R 24/ Caudate nucleus R < Posterior cingulate gyrus L 31 )16 ) Midline cerebellum R 14 )56 ) Cuneus L 18 )2 ) Lingual gyrus L 19 )2 ) a Note: BA: Brodmann Area; Hem: Hemisphere; Ratio: stability coe cient in bootstrap analysis; degrees of freedom for F ˆ 1; 55; coordinates of representative maxima and minima listed in mm according to Talairach and Tournoux (1988). rcbf at encoding were anterior and posterior portions of left parahippocampal gyrus. These MTL regions are shown in Fig. 2(b) together with a corresponding response pro le Format-related e ects speci c to recognition LV 4 revealed only a signi cant trend when assessed for signi cance in terms of the entire singular image. Brain regions associated with maxima and minima of this image showed a di erential rcbf response during recognition that depended on the format in which objects were presented at encoding; this di erential response was observed irrespective of the format of the stimulus present during recognition (see Fig. 1(c)). Several regions made stable contributions to the singular image identi ed with LV 4. When examined with univariate tests, none of them showed rcbf differences that met the signi cance criterion of P < 0:001 but several regions showed signi cant di erences at the less stringent level of P < 0:01 (see Table 4). Among these regions was a maximum in right anterior parahippocampal gyrus that showed higher rcbf during recognition of objects encoded as pictures (RPEP, RWEP) than during recognition of objects encoded as words (RPEW, RWEW). Fig. 2(c) displays this region together with a further anterior right parahippocampal region (XYZ ˆ ) that showed the same e ect at P < 0:03.

12 170 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 Table 4 Maxima and minima of singular image 4 re ecting format-speci c e ects speci c to recognition a Region Hem BA x y z Ratio F F prob Maxima (recognition of pictorially encoded objects > recognition of objects encoded as words) Anterior parahippocampal gyrus/amygdala R )10 ) Minima (recognition of objects encoded as words > recognition of pictorially encoded objects) Inferior frontal gyrus R 47/ R Brainstem R 4 )4 ) Inferior parietal lobule/ L 40 )34 ) intraparietal sulcus Inferior/middle occipital L 19 )42 ) gyrus Parieto-occipital sulcus L 7 )28 ) a Note: BA: Brodmann Area; Hem: Hemisphere; Ratio: stability coe cient in bootstrap analysis; degrees of freedom for F ˆ 1; 33; coordinates of representative maxima and minima are listed in mm according to Talairach & Tournoux (1988) Task-related e ects independent of object format It is noteworthy that the PLS analysis also extracted a task-related e ect that was observed independent of perceptual format. The singular image of the rst LV from the PLS analysis, which re ected the strongest e ect in the data, identi ed brain regions that showed these general rcbf di erences between recognition and encoding (see Fig. 1(d)). Maxima and minima of this image are listed in Table 5. Regions in which maxima were found showed increases in rcbf during recognition as compared to encoding, irrespective of object format, and included pre-frontal and parietal regions but no peak in the MTLs'. Regions associated with minima of singular image 1 showed higher rcbf during encoding than during recognition. They extended predominantly across bilateral temporal cortices and included a peak in left anterior parahippocampal gyrus (uncus) Summary and further examination of e ects in MTL regions The observed patterns of rcbf changes in the MTLs' that were found to be related to object format are summarized in Fig. 2. The gure demonstrates that object format a ected these MTL regions in di erent ways. The peak displayed in Fig. 2(a), which was extracted with LV 2, showed a response to object format that was comparable at encoding and recognition. MTL peaks in Fig. 2(b), which were extracted with LV 3, showed a response to object format that was speci c to encoding. Finally, the peaks in Fig. 2(c), which were extracted with LV 4, showed a formatrelated response that was speci c to recognition. For the peaks displayed in Fig. 2(a) and (b), the two di erent statistical assessments converged in demonstrating that these regions make stable contributions to the patterns identi ed with PLS (bootstrap ratio > 2:0) and show a signi cant e ect (P < 0:001) in the univariate tests coding for the corresponding contrast. For the

13 S. Kohler et al. / Acta Psychologica 105 (2000) 159± Table 5 Maxima and minima of singular image 1 re ecting task-related e ects independent of object format a Region Hem BA x y z Ratio F F prob Maxima (recognition > encoding) Middle/inferior frontal R gyrus R < R 8/ < L 10 ) < L 46 ) < L 8/9 ) Middle temporal gyrus L 21 )64 )32 ) < Retrosplenial cortex L 23/ )6 ) (posterior cingulate gyrus) 29 Inferior parietal sulcus R 39/ 36 ) < L 39/ )36 ) < Parieto-occipital sulcus L 7/19 )12 ) Minima (encoding > recognition) Anterior cingulate gyrus L 24 )10 26 ) < Superior/middle temporal R ) < gyrus (temporal pole) R ) < L 38 )42 14 ) < Insular cortex L ) Anterior parahippocampal L 35/ )30 )10 ) gyrus/uncus 36 Anterior fusiform gyrus R )12 ) L 20 )42 )12 ) < Superior temporal gyrus/ R ) < inferior parietal lobule L 22 )62 ) L 40 )58 ) < Middle temporal gyrus L 37 )50 ) Midline cerebellum R 18 )58 ) L )26 )60 ) < Inferior/middle occipital gyrus R 18/ ) a Note: BA: Brodmann Area; Hem: Hemisphere; Ratio: stability coe cient in bootstrap analysis; degrees of freedom for F ˆ 1; 55; coordinates of representative maxima and minima listed in mm according to Talairach and Tournoux (1988). peak in Fig. 2(c), the univariate test was signi cant at a less stringent level (P < 0:01). The results of additional univariate analyses, which were performed to allow for more speci c comparisons than those re ected in the overall contrasts, are presented in the following summary. The right posterior parahippocampal gyrus region shown in Fig. 2(a) exhibited an rcbf increase during picture as compared to word processing at encoding

14 172 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 (t 11 ˆ3:51; P ˆ 0:005) and at recognition. During recognition, it responded to the format of the stimulus present F 1; 11 ˆ10:71; P ˆ 0:007 but not to the format in which the targets had been encoded (P > 0:20). By contrast, the right anterior parahippocampal region shown in Fig. 2(c) demonstrated a picture-speci c response during recognition that did depend on the format in which the targets had been encoded F 1; 11 ˆ 14:00; P ˆ 0:003. Importantly, this response did not depend on the format of the stimulus present during recognition (P > 0:50). In the left hemisphere, anterior and more posterior parahippocampal regions showed format-speci c activation related to encoding; the rcbf for these regions was higher during picture than word encoding (anterior t 11 ˆ 2:75; P ˆ 0:019; posterior t 11 ˆ 2:33; P ˆ 0:040) with no comparable e ect at recognition (Ps > 0:10; Fig. 2(b)). In addition to these format-speci c e ects, we also found evidence for task-related activation in MTL regions that could be observed across both object formats. A region in left anterior parahippocampal gyrus was part of a network that demonstrated a general rcbf increase during encoding as compared to recognition regardless of whether processing of pictures or words was required (all Ps for formatrelated e ects >0.15). 3. Discussion The results of the present PET study provide clear evidence for format speci city with respect to the involvement of MTL regions in episodic encoding and recognition. We found that a right posterior MTL region was di erentially activated by picture stimuli irrespective of task (semantic encoding versus recognition); other MTL regions exhibited a di erential involvement in picture processing that was task speci c and appears to be tied to distinct memory processes involved in encoding and recognition. In addition to these format-speci c e ects, we found task-speci c activation in MTL regions that occurred irrespective of object format. Although the focus of the discussion will be on the MTLs', format speci c and general task-related e ects on rcbf were observed in other brain regions as well. These other activations will be discussed to the extent that they place the MTL results into context Format-related e ects comparable at encoding and recognition The network of brain regions identi ed with the second image from the PLS analysis showed a di erential rcbf response to pictures and words that was comparable during encoding and recognition; these brain regions responded di erentially to the stimuli presented during scanning regardless of the particular task demands. During recognition, their response was driven by the format of the stimulus present irrespective of the format in which the target information had been encoded. The most prominent increases in rcbf for pictures over words were found in bilateral visual extrastriate cortex (fusiform and lingual gyrus) and right posterior inferior and medial temporal cortex. Brain regions that showed an increase in rcbf for words

15 S. Kohler et al. / Acta Psychologica 105 (2000) 159± over pictures included portions of the left inferior parietal lobule and of dorsomedial cuneus. Not surprisingly, the extrastriate and parietal regions overlap with those that have previously been activated when the same type of stimuli were presented in the context of perceptual tasks for pictures and words, respectively (Howard et al., 1992; Martin, Wiggs, Ungerleider, & Haxby, 1996; Menard, Kosslyn, Thompson, Alpert, & Rauch, 1996; Vandenberghe, Price, Wise, Josephs, & Frackowiak, 1996). Thus, the di erential rcbf response to pictures and words observed in these regions in the present study most likely re ects the distinct perceptual identi cation processes that were triggered by the two di erent stimulus types during encoding and recognition. Importantly, a right posterior parahippocampal region also showed a di erential response to pictures during encoding and recognition. Grady et al. (1998) found activation in a nearby or identical region (peak: XYZ ˆ ; in present study: XYZ ˆ ) when picture encoding was compared with word encoding in the context of an episodic-memory paradigm. They suggested that pictures more e ectively engage those encoding processes that are supported by MTL regions. The present data bolster the notion that pictures engage right posterior MTL structures more so than words. However, they also suggest that the speci c processes supported by these structures are implicated in encoding and recognition tasks. This nding is corroborated by another recent report of picture-speci c right MTL activation that was observed in relation to encoding and recognition of complex visual scenes (Nyberg et al., 2000). Considered in isolation, the present data do not allow us to rule out that the functional contribution of right posterior MTL structures to picture processing is purely perceptual in nature. However, lesion studies in humans and non-human primates have shown that damage to MTL structures leads to severe episodic memory de cits while leaving perceptual abilities for the materials tested intact (Milner, 1990; Scoville & Milner, 1957). Studies in patients who underwent unilateral temporal lobectomy have demonstrated that the degree of impairment in memory for the names of previously studied objects is positively associated with the posterior extent of the lesion in right-sided MTL regions (Nunn et al., 1999; Smith & Milner, 1981). Further, recent evidence from a functional magnetic resonance-imaging study in healthy individuals suggests that the degree of activation in posterior MTL structures during encoding of complex pictures predicts subsequent performance accuracy on a memory task for these pictures (Brewer, Zhao, Desmond, Glover, & Gabrieli, 1998). In light of these ndings, it appears unlikely that right posterior MTL structures play a functional role that is limited to perceptual processing. Our data are open to the interpretation that right posterior MTL structures support encoding and recognition processes for pictorial stimuli (Kohler, McIntosh, Moscovitch, & Winocur, 1998). During encoding, this region may be involved in setting up a representation of the perceptual information that comprises the picture by directing this information to storage sites in posterior neocortex and by indexing the storage sites (Teyler & DiScenna, 1986). By contrast, during recognition, it may play a role in guiding recovery processes based on the picture presented as a cue at recognition (see Moscovitch, 1995). Clearly, further research is necessary in order to determine the functional importance of the involvement of right posterior MTL

16 174 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 structures in picture processing with more certainty. Regardless of the outcome of this research, however, the present ndings are important in that they demonstrate that the di erential involvement of right MTL structures in picture over word processing is not limited to encoding tasks Format-related e ects speci c to encoding With the third image of the PLS analysis, we identi ed a more speci c e ect of perceptual format on brain activation associated with episodic memory for objects. The brain regions that showed this e ect were di erentially involved in processing of pictures as opposed to words during encoding but showed no parallel di erence in rcbf across the recognition tasks. We found evidence for this type of format speci city in extrastriate cortex. Left medial aspects of the cuneus and lingual gyrus exhibited a higher rcbf response during word than picture encoding; by contrast, bilateral aspects of lateral fusiform gyrus and left middle occipital gyrus regions demonstrated a higher rcbf response during picture than word encoding. The activated regions in extrastriate cortex overlap with those that have been reported to participate in perceptual processing of objects presented as words and pictures, respectively (Bookheimer, Ze ro, Blaxton, Gaillard, & Theodore, 1995; Martin et al., 1996; Menard et al., 1996; Petersen, Fox, Snyder, & Raichle, 1990; Zelkowicz, Herbster, Nebes, Mintun, & Becker, 1998). Given that the recognition tasks of the present study had similar perceptual demands as the encoding tasks, our results suggest that parts of extrastriate cortex may support format-speci c processes of encoding that go over and beyond the perceptual analysis of objects. These processes may have to do with setting up records of the objects encountered. The e ect cannot be explained in terms of the perceptual novelty of the stimuli at encoding. Two of the four recognition tasks also involved the presentation of stimuli in a novel perceptual format (RPEW and RWEP); yet, the brain regions that showed format-speci c e ects during encoding exhibited no di erence in rcbf between these and the other two recognition tasks. The idea that parts of extrastriate cortex may contribute to establishing memory representations of the objects encountered during encoding is strengthened by our nding that there were MTL structures that showed parallel e ects, i.e. formatspeci c e ects that were speci c to the encoding tasks. Two distinct regions in left anterior and posterior parahippocampal gyrus were found to show a di erential rcbf increase during picture processing only at encoding. This result con rms those of three previous functional imaging studies that have shown an involvement of left MTL regions in picture encoding (Grady et al., 1998; Kelley et al., 1998; Martin et al., 1997). In each of these studies, left MTL structures were co-activated with right MTL structures. As discussed in the previous section, we also found format-speci c activation in right posterior MTL regions in relation to picture processing. However, our data indicate that this right MTL involvement is not restricted to encoding tasks. Thus, our results suggest that MTL structures in the left and right hemispheres may make distinct contributions to memory processing of pictures. Left-sided anterior MTL structures may speci cally be involved in establishing episodic memory

17 S. Kohler et al. / Acta Psychologica 105 (2000) 159± representations of objects based on their meaning (Martin et al., 1997). They do not appear to participate in accessing stored episodic information about objects, based on pictorial information, when processing of their meaning is not required Format-related e ects speci c to recognition We found only equivocal evidence for e ects of the perceptual format of objects at encoding on rcbf during recognition. Although the PLS analysis did extract an image with brain regions that showed such an e ect, this image as a whole was only marginally signi cant when examined by permutation test P ˆ 0:08. Moreover, none of the individual regions identi ed with this image showed changes in rcbf that could be con rmed with univariate statistical tests at a level commonly accepted in functional imaging research (P < 0:001 uncorrected). However, it is noteworthy that the image did include a region in right anterior parahippocampal gyrus that just failed to reach this statistical criterion P ˆ 0:0051. It showed higher rcbf during recognition of objects encoded as pictures than of objects encoded as words. Given that this di erence was observed irrespective of the format of the stimulus actually present at recognition, these results hint that portions of right anterior parahippocampal gyrus could play a speci c role in processes related to accessing picture-speci c information in episodic memory. This particular aspect of our data, although requiring replication in further neuroimaging studies, is in good accord with the results of a lesion study in humans (Jaccarino, 1975). Jaccarino's study showed that right-sided anterior temporal lobectomies can produce de cits in delayed recall of objects that were studied as pictures, even when pictorial cues are completely absent during recovery. The subtlety of the e ect of object format at encoding on brain activity during recognition is in keeping with the results from the PET study on cued recall of pictures and words by Buckner et al. (1996), which were summarized in the Introduction. Buckner et al. obtained these results with a di erent encoding task that speci cally required participants to focus their attention on perceptual attributes of the objects. Thus, the subtlety of the e ect seems to hold irrespective of the degree of perceptual processing demands at encoding. However, it may still be premature to conclude from these results that object format at encoding has generally only slight, if any, e ects on brain activity related to recovery of object information from episodic memory. It is important to consider that the recognition tasks of the present study did not require participants to decide whether the objects presented were originally seen as pictures or as words. Instead, participants were asked to decide whether they had encountered the objects previously irrespective of format. Recovery of information about encoding format may be associated with more pronounced di erences in patterns of rcbf that re ect whether objects were encountered as words or as pictures originally Task-related e ects independent of object format Interestingly, the present data provide no evidence for format-speci c increases in activation that were related to processing of words in MTL regions. This observation

18 176 S. Kohler et al. / Acta Psychologica 105 (2000) 159±179 may re ect the fact that the visual features of words are less discriminable than those of pictures (Nelson et al., 1976). However, this is not to say that words did not activate MTL structures in relation to memory processing in our study. We found evidence that left-sided MTL regions were part of a network of brain regions, identi ed with the rst singular image of the PLS analysis, that showed a general increase in rcbf during encoding as compared to recognition irrespective of object format. Our results con rm those of another study, in which semantic processing of object information, which can be conceptualized as episodic encoding (Kapur et al., 1994; Tulving, Kapur, Craik, Moscovitch, & Houle, 1994), was associated with higher rcbf in left parahippocampal gyrus than episodic retrieval of that same type of information (Wiggs, Weisberg, & Martin, 1999). Moreover, they converge with data reported by Vandenberghe et al. (1996), which showed that MTL structures are part of a left-lateralized network of regions that supports semantic processes across pictures and words. However, in conjunction with our results on format speci city discussed in the previous sections, the present ndings suggest functional specialization within the left MTL with respect to semantic processing; there appear to be some portions of left parahippocampal gyrus that participate similarly in semantic encoding of pictures and words whereas other subregions appear to make contributions that are picture speci c. 4. Conclusions The present data show that right MTL structures are di erentially involved in processing pictures of objects in episodic memory during encoding and recognition. A posterior right parahippocampal-gyrus region showed higher activation in response to the presentation of pictures than to words across all conditions. At encoding, this region may be involved in setting up memory representations. At recognition, it could play a role in retrieval processes that are driven by the perceptual input; these processes appear to be triggered by picture cues regardless of whether stored information is available in picture format. By contrast, another more anterior right parahippocampal region may be speci cally involved in accessing stored pictorial representations in episodic memory irrespective of whether the retrieval cue is pictorial in nature. The involvement of this region in object recognition appears to depend on the degree to which picture-speci c information needs to be accessed for the memory decision at hand. Working in concert, the right MTL structures identi ed in the present study would be in a position to support memory processing, encoding and recognition, of the distinct perceptual features that distinguish pictures from words in episodic memory. Our results also indicate that the MTL involvement in picture processing is not limited to the right hemisphere. During encoding of pictures, right MTL structures were found to be co-activated with left MTL structures. It is likely that these leftsided structures play a role in setting up an episodic record of the objects encountered based on their meaning rather than based on their visual appearance. Some regions in the left MTL show an involvement in semantic encoding of objects that is

19 S. Kohler et al. / Acta Psychologica 105 (2000) 159± picture speci c; others appear to show a more general involvement across pictures and words. Finally, taken together our ndings add further support to the general notion emerging from recent reviews of functional neuroimaging data (Lepage, Habib, & Tulving, 1998; Schacter & Wagner, 1999) that di erent MTL regions, across as well as within the two hemispheres, make distinct functional contributions to memory processing. Acknowledgements This research was supported by a Medical Research Council (MRC) of Canada grant to G.W. and M.M. and a Woman's Auxiliary Alzheimer Research Fellowship of Baycrest Centre for Geriatric Care to S.K. During the course of writing, S.K. was supported by a research fellowship at the Montreal Neurological Institute granted by Dr. Brenda Milner. We would like to thank Reza Habib for programming of the behavioral experiment, the sta at the PET Centre for administration of the scanning procedures, and Drs. Brenda Milner, Lars Nyberg, and Martin Lepage for helpful comments on this manuscript. References Amaral, D. G. (1999). Introduction: what is where in the medial temporal lobe?. Hippocampus, 9(1), 1±6. Bookheimer, S. Y., Ze ro, T. A., Blaxton, T., Gaillard, W., & Theodore, W. (1995). Regional cerebral bood ow during object naming and word reading. Human Brain Mapping, 3, 93±106. Brewer, J. B., Zhao, Z., Desmond, J. E., Glover, G. H., & Gabrieli, J. D. (1998). Making memories: brain activity that predicts how well visual experience will be remembered. Science, 281(5380), 1185±1187. Buckner, R. L., Raichle, M. E., Miezin, F. M., & Petersen, S. E. (1996). Functional anatomic studies of memory retrieval for auditory words and visual pictures. Journal of Neuroscience, 16(19), 6219±6235. Cabeza, R., Grady, C. L., Nyberg, L., McIntosh, A. R., Tulving, E., Kapur, S., Jennings, J. M., Houle, S., & Craik, F. I. (1997). Age-related di erences in neural activity during memory encoding and retrieval: a positron emission tomography study. Journal of Neuroscience, 17(1), 391±400. Friston, K. J., Holmes, A. P., Worsley, K. J., Poline, J. B., Frith, C. D., & Frackowiak, R. S. J. (1995). Statistical parametric maps in functional imaging: a general linear approach. Human Brain Mapping, 2, 189±210. Grady, C. L., McIntosh, A. R., Rajah, M. N., & Craik, F. I. (1998). Neural correlates of the episodic encoding of pictures and words. Proceedings of the National Academy of Sciences of the United States of America, 95(5), 2703±2708. Howard, D., Patterson, K., Wise, R., Brown, W. D., Friston, K., Weiller, C., & Frackowiak, R. (1992). The cortical localization of the lexicons. Positron emission tomography evidence. Brain, 115(Pt 6), 1769±1782. Incisa della Rocchetta, A. (1986). Classi cation and recall of pictures after unilateral frontal or temporal lobectomy. Cortex, 22, 189±211. Jaccarino, G. (1975). Dual encoding in memory: vidence from temporal-lobe lesions in man. Montreal: McGill University. Unpublished M.A. Thesis. Kapur, S., Craik, F. I., Tulving, E., Wilson, A. A., Houle, S., & Brown, G. M. (1994). Neuroanatomical correlates of encoding in episodic memory: levels of processing e ect. Proceedings of the National Academy of Sciences of the United States of America, 91(6), 2008±2011.

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