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...i that were inserted into the presentation sequence of stimuli, all other aspects of the tactile aversive condition were identical to the tactile control condition. This condition was always presented last in the scanning session. Data Analysis The CBF images were reconstructed and blurred with a Gaussian filter to a resolution of 12-mm, normalized for differences in global CBF, and coregistered with the individual MRIs [Woods et al., 1993]. The MRI volumes were then transformed into the Montreal Neurological Institute (MNI) standardized stereotaxic space that is based on the Talairach space [Talairach and Tournoux, 1988] by means of an automated feature-matching algorithm [Collins et al., 1994]. The statistical significance of focal changes was tested with a method based on 3D Gaussian random field theory. For an exploratory search involving all peaks within the grey matter volume of 600 ml, the threshold for reporting a peak as significant was set at t 5 4.41, corresponding to a corrected probability of P < 0.05. For a priori regions of interest within the orbitofrontal region, the threshold for significance was set at t 5 3.10, corresponding to a corrected probability of P < 0.05 based on a 2 cm diameter s...

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... with aversive stimuli. It should be noted that areas 11 and 13 are heavily connected to all the other areas of the orbitofrontal cortex [Carmichael and Price, 1996] as well as many areas of the lateral and medial prefrontal cortex [Barbas, 1993; Carmichael and Price, 1996]. These orbitofrontal areas, therefore, are in a unique position to mediate between the higher level control processes that depend on lateral prefrontal cortex (e.g. monitoring of information in working memory or active retrieval of information from memory) and the memory systems centered in the medial temporal lobe region [Petrides, 1996]. REFERENCES Aggleton JP, Burton MJ, Passingham RE (1980): Cortical and subcortical afferents to the amygdala of the rhesus monkey (macaca mulatta). Brain Res 190:347–368. r Encoding Touch and the Orbitofrontal Cortex r r 657 r Alexander MP, Freedman M (1984): Amnesia after anterior communicating artery aneurysm rupture. Neurology 34:752–757. Amaral DG, Price JL (1984): Amygdalo-cortical projections in the monkey (Macaca fascicularis). J Comp Neurol 230:465–496. Amaral DG, Price JL, Pitkanen A, Carmichael ST (1992):Anatomical organization of the primate amygdaloid complex. In: Aggleton JP, e...

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...Barbas, 1998] can produce autonomic changes, such as modifications of heart rate, respiration, and gastric motility [Bailey and Sweet, 1940; Delgado and Livingston, 1948; Hall and Cornish, 1977; Hall et al., 1977; Livingston et al., 1948]. Furthermore, there is a large body of evidence from lesion and electrophysiological studies demonstrating that the orbitofrontal cortex is involved in the processing of emotionally charged stimuli [Rolls, 2000]. In particular, there are orbitofrontal neurons that respond to deviations from expectation of reward [Rosenkilde et al., 1981; Thorpe et al., 1983; Tremblay and Schultz, 2000] which would be important in alerting the organism to potentially harmful stimuli. Additionally, there are studies which link lesions of the orbitofrontal cortex in monkeys [Butter, 1964; Butter et al., 1970; Izquierdo et al., 2005; Ruch and Shenkin, 1943] and patients [Hornak et al., 1996; Rolls et al., 1994] with abnormal behavioral responses to emotional stimuli, and functional neuroimaging data linking the orbitofrontal cortex in the human brain with emotional responses to stimuli [Angrilli et al., 1999; Blood et al., 1999; Dolan et al., 1996; Sarazin et al., 1998]. Several functional ima...

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...7; Accepted 27 November 2007 DOI: 10.1002/hbm.20532 Published online 2 January 2008 in Wiley InterScience (www. interscience.wiley.com). VC 2008 Wiley-Liss, Inc. r Human Brain Mapping 30:650–659 (2009) r Anatomical studies have demonstrated that the majority of connections between the medial temporal lobe and the frontal lobe are with the orbitofrontal cortex [Barbas and Blatt, 1995; Insausti et al., 1987]. In line with these facts, recent functional neuroimaging studies have demonstrated that the orbitofrontal cortex is involved in the encoding of visual [Frey and Petrides, 2000, 2002, 2003; Petrides et al., 2002], and auditory information [Frey et al., 2000, 2004]. Several functional imaging studies have also implicated the orbitofrontal cortex in various aspects of somatosensory processing [Francis et al., 1999; Hagen et al., 2002, Rolls et al., 2003]. In addition, the posterior agranular and dysgranular region of the orbitofrontal cortex has strong bidirectional connections with structures such as the amygdala [Amaral and Price, 1984; Barbas and de Olmos, 1990; Carmichael and Price, 1995; Pandya and Yeterian, 2001; Porrino et al., 1981] and the hypothalamus [Ongur et al., 1998; Rempel-Clower and ...

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...n 28 May 2007; Revised 11 October 2007; Accepted 27 November 2007 DOI: 10.1002/hbm.20532 Published online 2 January 2008 in Wiley InterScience (www. interscience.wiley.com). VC 2008 Wiley-Liss, Inc. r Human Brain Mapping 30:650–659 (2009) r Anatomical studies have demonstrated that the majority of connections between the medial temporal lobe and the frontal lobe are with the orbitofrontal cortex [Barbas and Blatt, 1995; Insausti et al., 1987]. In line with these facts, recent functional neuroimaging studies have demonstrated that the orbitofrontal cortex is involved in the encoding of visual [Frey and Petrides, 2000, 2002, 2003; Petrides et al., 2002], and auditory information [Frey et al., 2000, 2004]. Several functional imaging studies have also implicated the orbitofrontal cortex in various aspects of somatosensory processing [Francis et al., 1999; Hagen et al., 2002, Rolls et al., 2003]. In addition, the posterior agranular and dysgranular region of the orbitofrontal cortex has strong bidirectional connections with structures such as the amygdala [Amaral and Price, 1984; Barbas and de Olmos, 1990; Carmichael and Price, 1995; Pandya and Yeterian, 2001; Porrino et al., 1981] and the hypothalamus [Ongu...

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...2 Published online 2 January 2008 in Wiley InterScience (www. interscience.wiley.com). VC 2008 Wiley-Liss, Inc. r Human Brain Mapping 30:650–659 (2009) r Anatomical studies have demonstrated that the majority of connections between the medial temporal lobe and the frontal lobe are with the orbitofrontal cortex [Barbas and Blatt, 1995; Insausti et al., 1987]. In line with these facts, recent functional neuroimaging studies have demonstrated that the orbitofrontal cortex is involved in the encoding of visual [Frey and Petrides, 2000, 2002, 2003; Petrides et al., 2002], and auditory information [Frey et al., 2000, 2004]. Several functional imaging studies have also implicated the orbitofrontal cortex in various aspects of somatosensory processing [Francis et al., 1999; Hagen et al., 2002, Rolls et al., 2003]. In addition, the posterior agranular and dysgranular region of the orbitofrontal cortex has strong bidirectional connections with structures such as the amygdala [Amaral and Price, 1984; Barbas and de Olmos, 1990; Carmichael and Price, 1995; Pandya and Yeterian, 2001; Porrino et al., 1981] and the hypothalamus [Ongur et al., 1998; Rempel-Clower and Barbas, 1998] that are known to regulate emoti...

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...ic and autonomic regions of the brain, was activated when subjects explored novel aversive tactile stimuli. These results suggest that the orbitofrontal cortex, through its connections with the limbic areas of the medial temporal lobe, influences the processing of incoming information and thus contributes to its encoding. Hum Brain Mapp 30:650– 659, 2009. VC 2008 Wiley-Liss, Inc. Key words: somatosensory; tactile tasks; memory; positron emission tomography; human INTRODUCTION At the cortical level, tactile information is first coded in the primary somatosensory cortex (SI) as shown by lesion [Randolph and Semmes, 1974] and electrophysiological studies [Nelson et al., 1980] in the monkey. Further processing takes place in the secondary somatosensory cortex (SII) [Murray and Mishkin, 1984; Ridley and Ettlinger, 1976], as well as in the insular cortex [Friedman et al., 1986], leading to the perception of touch. Several investigators have described a cortico-limbic pathway for tactile memory, linking SI to insular areas via SII and then the insula to limbic structures in the medial temporal lobe, such as the entorhinal and perirhinal cortex, the amygdala, and the hippocampus [Friedman et al., 1986; Mishkin, 19...

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...orpe et al., 1983; Tremblay and Schultz, 2000] which would be important in alerting the organism to potentially harmful stimuli. Additionally, there are studies which link lesions of the orbitofrontal cortex in monkeys [Butter, 1964; Butter et al., 1970; Izquierdo et al., 2005; Ruch and Shenkin, 1943] and patients [Hornak et al., 1996; Rolls et al., 1994] with abnormal behavioral responses to emotional stimuli, and functional neuroimaging data linking the orbitofrontal cortex in the human brain with emotional responses to stimuli [Angrilli et al., 1999; Blood et al., 1999; Dolan et al., 1996; Sarazin et al., 1998]. Several functional imaging studies have investigated the orbitofrontal cortex during somatosensory processing and have reported bilateral activations throughout the orbitofrontal cortex [Francis et al., 1999; Hagen et al., 2002; Rolls et al., 2003]. Although the findings of the above studies were not related to mnemonic functioning but to attending to tactile stimuli, some of which contained hedonic information (e.g., pleasant and painful stimuli; see Francis et al., 1999; Rolls et al., 2003), these activity changes in the orbitofrontal cortex are consistent with our present results, sugges...