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...obiographical self. Extended consciousness builds on core consciousness, requires memory, and is enhanced by language. The discussion of these mechanisms is outside the scope of this article (but see =-=Damasio, 1999-=-). The role of brainstem structures in the generation of consciousness is thus a critical one. This article is dedicated to a review of some of the relevant evidence regarding the functional neuroanat...

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... noradrenergic system is involved in mechanisms underlying attention and learning (Aston-Jones & Bloom, 1981a,Aston-Jones and Bloom, 1981b; Aston-Jones, Rajkowski, Kubiak, Valentino, & Shipley, 1996; =-=Cahill and McGaugh, 1998-=-); the dopaminergic system is involved in motor control and reward mechanisms underlying motivation (Dunnett & Robbins, 1992; Brown & Gershon, 1993; Schultz et al., 1997; Schultz, 1998). Furthermore, ...

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...mulation within the reticular formation in lightly anesthetized non-human mammals, was associated with a desynchronization of the electroencephalogram (EEG) that hallmarks awake and attentive states (=-=Moruzzi & Magoun, 1949-=-; Lindsley, Schreiner, Knowles, Magoun, & Magoun, 1950; French & Magoun, 1952; Magoun, 1952a; Magoun, French, & Von Amerongen, 1952b; French, Verzeano, & Magoun, 1953). It was known by then that the r...

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...lity (for extensive review see Feldman, Meyer, & Quenzer, 1997: Chapter 9, pp. 380±9); the noradrenergic system is involved in mechanisms underlying attention and learning (Aston-Jones & Bloom, 1981a,=-=Aston-Jones and Bloom, 1981-=-b; Aston-Jones, Rajkowski, Kubiak, Valentino, & Shipley, 1996; Cahill and McGaugh, 1998); the dopaminergic system is involved in motor control and reward mechanisms underlying motivation (Dunnett & Ro...

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...rns of synchronization embedded in the global desynchronization (Munk, Roelfsema, KoÈnig, Engel, & Singer, 1996; Herculano-Houzel, Munk, Neuenschwander, & Singer, 1999). Llinas (Llinas & PareÂ, 1991; =-=Llinas, Ribary, Contreras, & Pedroarena, 1998-=-) and colleagues have found that the non-speci®c projections from the thalamus are important for generating a thalamocortical resonance which they suggest is a necessary substrate for consciousness. I...

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...nd dopaminergic nuclei (Moore, 1980). There are direct noradrenergic and serotonergic projections from the locus coeruleus and the rostral raphe complex, respectively, to most of the cortical mantle (=-=Moore & Bloom, 1979-=-). The dopaminergic projections from the substantia nigra and the J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159144 ventral tegmental area project extensively to the putamen, caudate nucleus, nu...

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...rting increased attentiveness and behavioral response to environmental stimuli, is well documented (Clark, Geffen, & Geffen, 1987; Jacobs, Wilkinson, & Fornal, 1990; Azmitia & Whitaker-Azmitia, 1991; =-=Aston-Jones, Chiang, & Alexinsky, 1991-=-; Berridge, Arnsten, & Foote, 1993; Geyer, 1996; Bloom, 1997; Cahill & McGaugh, 1998; Rico & Cavada, 1998). The role of dopaminergic nuclei in the same processes is less well understood although their...

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...n turn project to various cortical regions (Brodal, 1959; Jones & Leavitt, 1974; Edwards & de Olmos, 1976; Jackson & Crossman, 1983; Kaufman & Rosenquist, 1985; Steriade, Pare, Parent, & Smith, 1988; =-=Lavoie & Parent, 1994-=-; Groenewegen & Berendse, 1994; Newman & Ginsberg, 1994). The deep mesencephalic nucleus and, to a lesser extent, the nucleus pontis oralis project to the basal forebrain, from which widespread cholin...

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...lar formation is more complex than simply the desynchronization of its electrophysiological rhythm and leads, in effect, to local patterns of synchronization embedded in the global desynchronization (=-=Munk, Roelfsema, KoÈnig, Engel, & Singer, 1996-=-; Herculano-Houzel, Munk, Neuenschwander, & Singer, 1999). Llinas (Llinas & PareÂ, 1991; Llinas, Ribary, Contreras, & Pedroarena, 1998) and colleagues have found that the non-speci®c projections from ...

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... project to the PAG and the PBN (Hardy & Leichnetz, 1981a,b; Holstege, Meiners, & Tan, 1985; Moga, Herbert, Hurley, Yasui, Gray, & Saper, 1990; Beitz, 1990; Buchanan, Thompson, Maxwell, & Well, 1994; =-=An, Bandler, Ongur, & Price, 1998-=-; Moga et al., 1990). In a recent study, R.J. Morecraft has traced direct projections from the cingulate cortex to the locus coeruleus (personal communication). In conclusion, the state of the organis...

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...that C-®bers are also involved in detecting changes in pH, pCO2, pO2, glucose concentration, osmolarity and in signaling the presence of in¯ammatory agents (Moskowitz, 1991; MacIver & Tanelian, 1992; =-=Burnstock & Wood, 1996-=-; see Craig, 1997, for more references). Thus these ®bers carry signals related to the internal state of the organism. Contrary to the traditional view, not all C-®bers are silent in the absence of no...

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...effect, to local patterns of synchronization embedded in the global desynchronization (Munk, Roelfsema, KoÈnig, Engel, & Singer, 1996; Herculano-Houzel, Munk, Neuenschwander, & Singer, 1999). Llinas (=-=Llinas & PareÂ, 1991-=-; Llinas, Ribary, Contreras, & Pedroarena, 1998) and colleagues have found that the non-speci®c projections from the thalamus are important for generating a thalamocortical resonance which they sugges...

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...9 145 subregion of the PBN (Bester, Bourgeais, Villanueva, Besson, & Bernard, 1999), project to the intralaminar thalamic nuclei. Moreover, there are projections from the PBN (Fulwiler & Saper, 1984; =-=Alden, Besson, & Bernard, 1994-=-) and the PAG (Mantyh, 1983; Beitz, 1990; Parent & Steriade, 1981) to the basal forebrain and other brainstem nuclei such as the classical reticular nuclei involved in activating the cerebral cortex. ...

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...4). The deep mesencephalic nucleus and, to a lesser extent, the nucleus pontis oralis project to the basal forebrain, from which widespread cholinergic projections arise aimed at the cerebral cortex (=-=Jones & Yang, 1985-=-). The classical reticular nuclei mentioned above are located in the upper brainstem. However, some structures in the lower brainstem, well below midbrain and upper pons, may also have the anatomical ...

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...of the upper midbrain (Fig. 1A) (Olszewski & Baxter, 1982; Paxinos & Huang, 1995). It is anatomically continuous with the core regions of the spinal cord and extends rostrally into the thalamus (e.g. =-=Martin, 1996-=-). In short, the term reticular formation was assigned to a region of the brainstem when the nuclear heterogeneity of this region was not yet appreciated because of the limited methods of the time. Th...

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... Hand, 1978; Berkley et al., 1986; Wiberg & Blomqvist, 1984; Wiberg et al., 1987). In turn the tectum projects to the nuclei of the pons and midbrain (Shammah-Lagnado, Negrao, Silva, & Ricardo, 1987; =-=Cornwall, Cooper, & Phillipson, 1990-=-). Another motor-related channel to the upper brainstem is via the cerebellum (Brodal, 1959; Boivie, 1988; Rathelot & Padel, 1997). Some other nuclei in the brainstem, such as the J. Parvizi, A. Damas...

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...eus should be considered introceptive rather than only nociceptive (Craig, 1996, 1997). Interestingly, the PAG and the PBN are also major endpoints for projections from the NTS and the area postrema (=-=Beckstead, Morse, & Norgren, 1980-=-; Mantyh, 1982; Fulwiler & Saper, 1984; Herbert, Moga, & Saper, 1990; Ito & Seki, 1998). As mentioned, the NTS receives afferents through cranial nerves such as the vagus, which carry signals pertaini...

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...1980; Brown, 1982; Willis & Coggeshall, 1991: pp. 13±45). Among the brainstem nuclei that receive the majority of these C- and Ad ®berrelated inputs are the PBN and the PAG (Wiberg & Blomqvist, 1984; =-=Bernard & Besson, 1990-=-a; Blomqvist & Berkley, 1992; Barnett et al., 1995; Craig, J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159148 Fig. 4. The afferents to brainstem reticular nuclei. The brainstem reticular nuclei r...

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...ergic ascending projections to the basal ganglia and the intralaminar thalamic nuclei which in turn project to various cortical regions (Brodal, 1959; Jones & Leavitt, 1974; Edwards & de Olmos, 1976; =-=Jackson & Crossman, 1983-=-; Kaufman & Rosenquist, 1985; Steriade, Pare, Parent, & Smith, 1988; Lavoie & Parent, 1994; Groenewegen & Berendse, 1994; Newman & Ginsberg, 1994). The deep mesencephalic nucleus and, to a lesser exte...

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...laminar nuclei of the thalamus, which are the origin of the so-called diffuse thalamocortical projections, since they are not connected in topographical fashion with speci®c sensory or motor regions (=-=Morison & Dempsey, 1942-=-). As a consequence, it was proposed that the brainstem reticular formation is the origin of the ascending reticular activating system that J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159 141 Fig...

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...la, and prefrontal cortex are also known to project to the PAG and the PBN (Hardy & Leichnetz, 1981a,b; Holstege, Meiners, & Tan, 1985; Moga, Herbert, Hurley, Yasui, Gray, & Saper, 1990; Beitz, 1990; =-=Buchanan, Thompson, Maxwell, & Well, 1994-=-; An, Bandler, Ongur, & Price, 1998; Moga et al., 1990). In a recent study, R.J. Morecraft has traced direct projections from the cingulate cortex to the locus coeruleus (personal communication). In c...

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...ornwall et al., 1990). These rostral structures and the extended amygdala, cingulate gyrus, insula, and prefrontal cortex are also known to project to the PAG and the PBN (Hardy & Leichnetz, 1981a,b; =-=Holstege, Meiners, & Tan, 1985-=-; Moga, Herbert, Hurley, Yasui, Gray, & Saper, 1990; Beitz, 1990; Buchanan, Thompson, Maxwell, & Well, 1994; An, Bandler, Ongur, & Price, 1998; Moga et al., 1990). In a recent study, R.J. Morecraft ha...

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..., Villanueva, Besson, & Bernard, 1999), project to the intralaminar thalamic nuclei. Moreover, there are projections from the PBN (Fulwiler & Saper, 1984; Alden, Besson, & Bernard, 1994) and the PAG (=-=Mantyh, 1983-=-; Beitz, 1990; Parent & Steriade, 1981) to the basal forebrain and other brainstem nuclei such as the classical reticular nuclei involved in activating the cerebral cortex. Thus the PBN and the PAG ha...

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... Feldman, Meyer, & Quenzer, 1997: Chapter 9, pp. 380±9); the noradrenergic system is involved in mechanisms underlying attention and learning (Aston-Jones & Bloom, 1981a,Aston-Jones and Bloom, 1981b; =-=Aston-Jones, Rajkowski, Kubiak, Valentino, & Shipley, 1996-=-; Cahill and McGaugh, 1998); the dopaminergic system is involved in motor control and reward mechanisms underlying motivation (Dunnett & Robbins, 1992; Brown & Gershon, 1993; Schultz et al., 1997; Sch...

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...ta (Muller, Lewandowski, & Singer, 1993). The reticular nucleus of the thalamus projects to other thalamic nuclei (Scheibel & Scheibel, 1966), and inhibits their activity (Steriade & Deschenes, 1984; =-=Barth & MacDonald, 1996-=-), thereby functioning as a pacemaker for the thalamic spindle oscillations which hallmark deep sleep (Steriade & Deschenes, 1984; Steriade, McCormick, & Sejnowski, 1993). The activity of the brainste...

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...linated Ad ®bers which convey signals related to pain and temperature. The phylogenetically old C- and Ad ®bers have free endings, unlike other sensory ®bers which have specialized sensory receptors (=-=Cervero & Iggo, 1980-=-; Brown, 1982; Willis & Coggeshall, 1991: pp. 13±45). Among the brainstem nuclei that receive the majority of these C- and Ad ®berrelated inputs are the PBN and the PAG (Wiberg & Blomqvist, 1984; Bern...

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...gnition 79 (2001) 135±159 145 subregion of the PBN (Bester, Bourgeais, Villanueva, Besson, & Bernard, 1999), project to the intralaminar thalamic nuclei. Moreover, there are projections from the PBN (=-=Fulwiler & Saper, 1984-=-; Alden, Besson, & Bernard, 1994) and the PAG (Mantyh, 1983; Beitz, 1990; Parent & Steriade, 1981) to the basal forebrain and other brainstem nuclei such as the classical reticular nuclei involved in ...

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...associated with a desynchronization of the electroencephalogram (EEG) that hallmarks awake and attentive states (Moruzzi & Magoun, 1949; Lindsley, Schreiner, Knowles, Magoun, & Magoun, 1950; French & =-=Magoun, 1952-=-; Magoun, 1952a; Magoun, French, & Von Amerongen, 1952b; French, Verzeano, & Magoun, 1953). It was known by then that the reticular formation projects to the intralaminar nuclei of the thalamus, which...

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...±45). Among the brainstem nuclei that receive the majority of these C- and Ad ®berrelated inputs are the PBN and the PAG (Wiberg & Blomqvist, 1984; Bernard & Besson, 1990a; Blomqvist & Berkley, 1992; =-=Barnett et al., 1995-=-; Craig, J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159148 Fig. 4. The afferents to brainstem reticular nuclei. The brainstem reticular nuclei receive afferents from various sources. The state o...

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...ctive, that of life regulation, and that the new perspective may help explain why and how brainstem nuclei exert their varied in¯uences on structures located rostrally, namely on the cerebral cortex (=-=Damasio, 1998-=-, 1999). 1.1. A brief summary of the new proposal Some nuclei of the brainstem have long been linked to the regulation of life, along with nuclei in the nearby hypothalamus, but a link between nuclei ...

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...tral structures and the extended amygdala, cingulate gyrus, insula, and prefrontal cortex are also known to project to the PAG and the PBN (Hardy & Leichnetz, 1981a,b; Holstege, Meiners, & Tan, 1985; =-=Moga, Herbert, Hurley, Yasui, Gray, & Saper, 1990-=-; Beitz, 1990; Buchanan, Thompson, Maxwell, & Well, 1994; An, Bandler, Ongur, & Price, 1998; Moga et al., 1990). In a recent study, R.J. Morecraft has traced direct projections from the cingulate cort...

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...vada, 1998). The role of dopaminergic nuclei in the same processes is less well understood although their central role in motor control and reward mechanisms underlying motivation is widely accepted (=-=Dunnett & Robbins, 1992-=-; Brown & Gershon, 1993; Schultz, Dayan, & Montague, 1997; Schultz, 1998). The above-mentioned monoaminergic nuclei are located within the upper reticular formation. Monoaminergic nuclei in the lower ...

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...ding rather than ascending projections (Moore, 1980). 3. The cholinergic nuclei which include the laterodorsal tegmental nucleus and the cholinergic portion of the pedunculopontine tegmental nucleus (=-=Mesulam, Geula, Bothwell, & Hersh, 1989-=-). These cholinergic nuclei are also located in the upper brainstem. They project to several thalamic nuclei including the reticular nucleus of the thalamus (Pare, Smith, Parent, & Steriade, 1988; Ste...

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...oral response to environmental stimuli, is well documented (Clark, Geffen, & Geffen, 1987; Jacobs, Wilkinson, & Fornal, 1990; Azmitia & Whitaker-Azmitia, 1991; Aston-Jones, Chiang, & Alexinsky, 1991; =-=Berridge, Arnsten, & Foote, 1993-=-; Geyer, 1996; Bloom, 1997; Cahill & McGaugh, 1998; Rico & Cavada, 1998). The role of dopaminergic nuclei in the same processes is less well understood although their central role in motor control and...

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...eticular formation. They send presumably glutamatergic ascending projections to the basal ganglia and the intralaminar thalamic nuclei which in turn project to various cortical regions (Brodal, 1959; =-=Jones & Leavitt, 1974-=-; Edwards & de Olmos, 1976; Jackson & Crossman, 1983; Kaufman & Rosenquist, 1985; Steriade, Pare, Parent, & Smith, 1988; Lavoie & Parent, 1994; Groenewegen & Berendse, 1994; Newman & Ginsberg, 1994). ...

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...cular formation in lightly anesthetized non-human mammals, was associated with a desynchronization of the electroencephalogram (EEG) that hallmarks awake and attentive states (Moruzzi & Magoun, 1949; =-=Lindsley, Schreiner, Knowles, Magoun, & Magoun, 1950-=-; French & Magoun, 1952; Magoun, 1952a; Magoun, French, & Von Amerongen, 1952b; French, Verzeano, & Magoun, 1953). It was known by then that the reticular formation projects to the intralaminar nuclei...

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...l activity of cortex, and in supporting increased attentiveness and behavioral response to environmental stimuli, is well documented (Clark, Geffen, & Geffen, 1987; Jacobs, Wilkinson, & Fornal, 1990; =-=Azmitia & Whitaker-Azmitia, 1991-=-; Aston-Jones, Chiang, & Alexinsky, 1991; Berridge, Arnsten, & Foote, 1993; Geyer, 1996; Bloom, 1997; Cahill & McGaugh, 1998; Rico & Cavada, 1998). The role of dopaminergic nuclei in the same processe...

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...ted the term reticular formation. They send presumably glutamatergic ascending projections to the basal ganglia and the intralaminar thalamic nuclei which in turn project to various cortical regions (=-=Brodal, 1959-=-; Jones & Leavitt, 1974; Edwards & de Olmos, 1976; Jackson & Crossman, 1983; Kaufman & Rosenquist, 1985; Steriade, Pare, Parent, & Smith, 1988; Lavoie & Parent, 1994; Groenewegen & Berendse, 1994; New...

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...als, was associated with a desynchronization of the electroencephalogram (EEG) that hallmarks awake and attentive states (Moruzzi & Magoun, 1949; Lindsley, Schreiner, Knowles, Magoun, & Magoun, 1950; =-=French & Magoun, 1952-=-; Magoun, 1952a; Magoun, French, & Von Amerongen, 1952b; French, Verzeano, & Magoun, 1953). It was known by then that the reticular formation projects to the intralaminar nuclei of the thalamus, which...

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...ormed by several families of nuclei (Fig. 3). Accordingly, several studies have con®rmed that bilateral single lesions to some of the brainstem nuclei mentioned above are not suf®cient to cause coma (=-=Jones et al., 1973-=-; Kitsikis & Steriade, 1981; Webster & Jones, 1988; Lai, Shalita, Hajnik, Wu, Kuo, Chia, & Siegel, 1999). Third, it also indicates that the notion of ªmesencephalicº reticular formation as the sole pl...

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...the serotonergic and noradrenergic systems in modulating the global activity of cortex, and in supporting increased attentiveness and behavioral response to environmental stimuli, is well documented (=-=Clark, Geffen, & Geffen, 1987-=-; Jacobs, Wilkinson, & Fornal, 1990; Azmitia & Whitaker-Azmitia, 1991; Aston-Jones, Chiang, & Alexinsky, 1991; Berridge, Arnsten, & Foote, 1993; Geyer, 1996; Bloom, 1997; Cahill & McGaugh, 1998; Rico ...

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...muli, is well documented (Clark, Geffen, & Geffen, 1987; Jacobs, Wilkinson, & Fornal, 1990; Azmitia & Whitaker-Azmitia, 1991; Aston-Jones, Chiang, & Alexinsky, 1991; Berridge, Arnsten, & Foote, 1993; =-=Geyer, 1996-=-; Bloom, 1997; Cahill & McGaugh, 1998; Rico & Cavada, 1998). The role of dopaminergic nuclei in the same processes is less well understood although their central role in motor control and reward mecha...

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... changes in the position and the movement of the head in space. There are major projections from the vestibular nuclei to other brainstem nuclei such as the PBN in the upper brainstem (Balaban, 1996; =-=Balaban & Porter, 1998-=-). These projections are involved in mediating adjustments in cardiovascular, respiratory, and gastroenteric functions needed when the position of the body is changed in space. 4. The state of the mus...

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...ee Willis and Coggeshall, 1991: pp. 265±306). In turn, there are distinct projections from the dorsal column nuclei to rostral regions such as in the midbrain, thalamus, zona incerta, and cerebellum (=-=Berkley, Budell, Blomqvist, & Bull, 1986-=-). Interestingly, the regions that receive primary cutaneous afferents project, in somatotopical order, to the thalamic relay nuclei whereas the upper brainstem receives projections from neurons that ...

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...tex. For instance, both PAG (Jones & Yang, 1985; Kaufman & Rosenquist, 1985; Pare et al., 1988) and the internal lateral J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159 145 subregion of the PBN (=-=Bester, Bourgeais, Villanueva, Besson, & Bernard, 1999-=-), project to the intralaminar thalamic nuclei. Moreover, there are projections from the PBN (Fulwiler & Saper, 1984; Alden, Besson, & Bernard, 1994) and the PAG (Mantyh, 1983; Beitz, 1990; Parent & S...

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... & Coggeshall, 1991: pp. 13±45). Among the brainstem nuclei that receive the majority of these C- and Ad ®berrelated inputs are the PBN and the PAG (Wiberg & Blomqvist, 1984; Bernard & Besson, 1990a; =-=Blomqvist & Berkley, 1992-=-; Barnett et al., 1995; Craig, J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159148 Fig. 4. The afferents to brainstem reticular nuclei. The brainstem reticular nuclei receive afferents from variou...

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...n, 1996, for more references). As Craig has suggested, the ascending pathways from lamina I and the caudal spinal trigeminal subnucleus should be considered introceptive rather than only nociceptive (=-=Craig, 1996-=-, 1997). Interestingly, the PAG and the PBN are also major endpoints for projections from the NTS and the area postrema (Beckstead, Morse, & Norgren, 1980; Mantyh, 1982; Fulwiler & Saper, 1984; Herber...

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... that hallmarks awake and attentive states (Moruzzi & Magoun, 1949; Lindsley, Schreiner, Knowles, Magoun, & Magoun, 1950; French & Magoun, 1952; Magoun, 1952a; Magoun, French, & Von Amerongen, 1952b; =-=French, Verzeano, & Magoun, 1953-=-). It was known by then that the reticular formation projects to the intralaminar nuclei of the thalamus, which are the origin of the so-called diffuse thalamocortical projections, since they are not ...

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...the PAG and the PBN are also major endpoints for projections from the NTS and the area postrema (Beckstead, Morse, & Norgren, 1980; Mantyh, 1982; Fulwiler & Saper, 1984; Herbert, Moga, & Saper, 1990; =-=Ito & Seki, 1998-=-). As mentioned, the NTS receives afferents through cranial nerves such as the vagus, which carry signals pertaining to the visceral state. While the NTS constructs a neural map of the viscera, the ar...

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...s to the basal ganglia and the intralaminar thalamic nuclei which in turn project to various cortical regions (Brodal, 1959; Jones & Leavitt, 1974; Edwards & de Olmos, 1976; Jackson & Crossman, 1983; =-=Kaufman & Rosenquist, 1985-=-; Steriade, Pare, Parent, & Smith, 1988; Lavoie & Parent, 1994; Groenewegen & Berendse, 1994; Newman & Ginsberg, 1994). The deep mesencephalic nucleus and, to a lesser extent, the nucleus pontis orali...

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...colleagues, the stimulation of the PBN was found to induce maximal changes in the electrophysiological activity of cortex. In a series of studies by Moruzzi (Moruzzi, Magni, Rossi, & Zanchetti, 1959; =-=Moruzzi, 1963-=-) and others (Batini, Moruzzi, Palestini, Rossi, & Zanchetti, 1959) it was found that another component of the brainstem autonomic system, the nucleus tractus solitarius (NTS) in the medulla, can stro...

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...on occupies most of the central and dorsal part of the brainstem extending from J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159140 the lower medulla to the level of the upper midbrain (Fig. 1A) (=-=Olszewski & Baxter, 1982-=-; Paxinos & Huang, 1995). It is anatomically continuous with the core regions of the spinal cord and extends rostrally into the thalamus (e.g. Martin, 1996). In short, the term reticular formation was...

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...Gershon, 1993; Schultz et al., 1997; Schultz, 1998). Furthermore, classical reticular nuclei such as the nucleus cuneiforme and the pedunculopontine tegmental nucleus are also involved in locomotion (=-=Allen, Inglis, & Winn, 1996-=-). The pedunculopontine tegmental nucleus also plays an important role in mechanisms underlying attention and learning (Allen et al., 1996), and in subserving the rewarding effect of opiates (Bechara ...

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...ed in detecting changes in the position and the movement of the head in space. There are major projections from the vestibular nuclei to other brainstem nuclei such as the PBN in the upper brainstem (=-=Balaban, 1996-=-; Balaban & Porter, 1998). These projections are involved in mediating adjustments in cardiovascular, respiratory, and gastroenteric functions needed when the position of the body is changed in space....

Citation Context

... of the PBN was found to induce maximal changes in the electrophysiological activity of cortex. In a series of studies by Moruzzi (Moruzzi, Magni, Rossi, & Zanchetti, 1959; Moruzzi, 1963) and others (=-=Batini, Moruzzi, Palestini, Rossi, & Zanchetti, 1959-=-) it was found that another component of the brainstem autonomic system, the nucleus tractus solitarius (NTS) in the medulla, can strongly modulate the global activity of the cerebral cortex. In these...

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...cal reticular nuclei such as the nucleus cuneiforme are examples of other nuclei that receive this kind of spinal afferents (Wiberg & Blomqvist, 1984; Blomqvist & Berkley, 1992; Barnett et al., 1995; =-=Craig, 1995-=-; Willis and Westlund, 1997). Projections from the super®cial dorsal horn of the spinal cord and the caudal spinal trigeminal subnucleus provide the anatomical means for relaying to the upper brainste...

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...ed in detecting changes in pH, pCO2, pO2, glucose concentration, osmolarity and in signaling the presence of in¯ammatory agents (Moskowitz, 1991; MacIver & Tanelian, 1992; Burnstock & Wood, 1996; see =-=Craig, 1997-=-, for more references). Thus these ®bers carry signals related to the internal state of the organism. Contrary to the traditional view, not all C-®bers are silent in the absence of noxious stimuli (e....

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...peci®c lesion in the core of the midbrain ± that spared both ascending pathways originated below the midbrain and local connections within the midbrain ± did not cause alterations in the EEG pattern (=-=Denoyer, Sallanon, Kitahama, & Jouvet, 1991-=-). J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159146 3. A functional context for the ascending reticular activating system In the introduction to this article, we noted that it is important to u...

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...mmah-Lagnado et al., 1987; Cornwall et al., 1990). These rostral structures and the extended amygdala, cingulate gyrus, insula, and prefrontal cortex are also known to project to the PAG and the PBN (=-=Hardy & Leichnetz, 1981-=-a,b; Holstege, Meiners, & Tan, 1985; Moga, Herbert, Hurley, Yasui, Gray, & Saper, 1990; Beitz, 1990; Buchanan, Thompson, Maxwell, & Well, 1994; An, Bandler, Ongur, & Price, 1998; Moga et al., 1990). I...

Citation Context

...gic systems in modulating the global activity of cortex, and in supporting increased attentiveness and behavioral response to environmental stimuli, is well documented (Clark, Geffen, & Geffen, 1987; =-=Jacobs, Wilkinson, & Fornal, 1990-=-; Azmitia & Whitaker-Azmitia, 1991; Aston-Jones, Chiang, & Alexinsky, 1991; Berridge, Arnsten, & Foote, 1993; Geyer, 1996; Bloom, 1997; Cahill & McGaugh, 1998; Rico & Cavada, 1998). The role of dopami...

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...ilies of nuclei (Fig. 3). Accordingly, several studies have con®rmed that bilateral single lesions to some of the brainstem nuclei mentioned above are not suf®cient to cause coma (Jones et al., 1973; =-=Kitsikis & Steriade, 1981-=-; Webster & Jones, 1988; Lai, Shalita, Hajnik, Wu, Kuo, Chia, & Siegel, 1999). Third, it also indicates that the notion of ªmesencephalicº reticular formation as the sole platform for modulating the g...

Citation Context

...ve rather than only nociceptive (Craig, 1996, 1997). Interestingly, the PAG and the PBN are also major endpoints for projections from the NTS and the area postrema (Beckstead, Morse, & Norgren, 1980; =-=Mantyh, 1982-=-; Fulwiler & Saper, 1984; Herbert, Moga, & Saper, 1990; Ito & Seki, 1998). As mentioned, the NTS receives afferents through cranial nerves such as the vagus, which carry signals pertaining to the visc...

Citation Context

... documented (Clark, Geffen, & Geffen, 1987; Jacobs, Wilkinson, & Fornal, 1990; Azmitia & Whitaker-Azmitia, 1991; Aston-Jones, Chiang, & Alexinsky, 1991; Berridge, Arnsten, & Foote, 1993; Geyer, 1996; =-=Bloom, 1997-=-; Cahill & McGaugh, 1998; Rico & Cavada, 1998). The role of dopaminergic nuclei in the same processes is less well understood although their central role in motor control and reward mechanisms underly...

Citation Context

... convey signals related to pain and temperature. The phylogenetically old C- and Ad ®bers have free endings, unlike other sensory ®bers which have specialized sensory receptors (Cervero & Iggo, 1980; =-=Brown, 1982-=-; Willis & Coggeshall, 1991: pp. 13±45). Among the brainstem nuclei that receive the majority of these C- and Ad ®berrelated inputs are the PBN and the PAG (Wiberg & Blomqvist, 1984; Bernard & Besson,...

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...mmah-Lagnado et al., 1987; Cornwall et al., 1990). These rostral structures and the extended amygdala, cingulate gyrus, insula, and prefrontal cortex are also known to project to the PAG and the PBN (=-=Hardy & Leichnetz, 1981-=-a,b; Holstege, Meiners, & Tan, 1985; Moga, Herbert, Hurley, Yasui, Gray, & Saper, 1990; Beitz, 1990; Buchanan, Thompson, Maxwell, & Well, 1994; An, Bandler, Ongur, & Price, 1998; Moga et al., 1990). I...

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...synchronization of its electrophysiological rhythm and leads, in effect, to local patterns of synchronization embedded in the global desynchronization (Munk, Roelfsema, KoÈnig, Engel, & Singer, 1996; =-=Herculano-Houzel, Munk, Neuenschwander, & Singer, 1999-=-). Llinas (Llinas & PareÂ, 1991; Llinas, Ribary, Contreras, & Pedroarena, 1998) and colleagues have found that the non-speci®c projections from the thalamus are important for generating a thalamocorti...

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...dies have con®rmed that bilateral single lesions to some of the brainstem nuclei mentioned above are not suf®cient to cause coma (Jones et al., 1973; Kitsikis & Steriade, 1981; Webster & Jones, 1988; =-=Lai, Shalita, Hajnik, Wu, Kuo, Chia, & Siegel, 1999-=-). Third, it also indicates that the notion of ªmesencephalicº reticular formation as the sole platform for modulating the global activity of the cerebral cortex is incorrect because many of the relev...

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...nis, Pieribone, Nickell, & Shipley, 1986; Van Bockstaele & Aston-Jones, 1992, 1995). 2. The monoaminergic nuclei of the brainstem which encompass noradrenergic, serotonergic, and dopaminergic nuclei (=-=Moore, 1980-=-). There are direct noradrenergic and serotonergic projections from the locus coeruleus and the rostral raphe complex, respectively, to most of the cortical mantle (Moore & Bloom, 1979). The dopaminer...

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...gly, in a recent study by Munk (1996) and colleagues, the stimulation of the PBN was found to induce maximal changes in the electrophysiological activity of cortex. In a series of studies by Moruzzi (=-=Moruzzi, Magni, Rossi, & Zanchetti, 1959-=-; Moruzzi, 1963) and others (Batini, Moruzzi, Palestini, Rossi, & Zanchetti, 1959) it was found that another component of the brainstem autonomic system, the nucleus tractus solitarius (NTS) in the me...

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...thin the brainstem nuclei. For abbreviations see Fig. 2. 1995; Willis and Westlund, 1997). In fact it is estimated that the lamina I projects three times more densely to the PAG than to the thalamus (=-=Mouton & Holstege, 1998-=-). The noradrenergic nuclei such as the locus coeruleus, and the classical reticular nuclei such as the nucleus cuneiforme are examples of other nuclei that receive this kind of spinal afferents (Wibe...

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...959; Jones & Leavitt, 1974; Edwards & de Olmos, 1976; Jackson & Crossman, 1983; Kaufman & Rosenquist, 1985; Steriade, Pare, Parent, & Smith, 1988; Lavoie & Parent, 1994; Groenewegen & Berendse, 1994; =-=Newman & Ginsberg, 1994-=-). The deep mesencephalic nucleus and, to a lesser extent, the nucleus pontis oralis project to the basal forebrain, from which widespread cholinergic projections arise aimed at the cerebral cortex (J...

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...and aggression, whereas stimulation of ventrolateral columns of the PAG, on the other hand, produces a passive coping strategy with hyporeactivity, hypotension, bradycardia, freezing, and immobility (=-=Bandler & Shipley, 1994-=-). Evidence from functional imaging studies also supports the notion that the upper brainstem nuclei are involved in a broad range of functions. For instance, Maquet and colleagues (Maquet, Dive, Salm...

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...Besson, & Bernard, 1999), project to the intralaminar thalamic nuclei. Moreover, there are projections from the PBN (Fulwiler & Saper, 1984; Alden, Besson, & Bernard, 1994) and the PAG (Mantyh, 1983; =-=Beitz, 1990-=-; Parent & Steriade, 1981) to the basal forebrain and other brainstem nuclei such as the classical reticular nuclei involved in activating the cerebral cortex. Thus the PBN and the PAG have the anatom...

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...jections from neurons that receive nonprimary or muscle afferents (Berkley et al., 1986; Wiberg, Westman, & Blomqvist, 1987). In the midbrain, the tectum is among the recipients of these projections (=-=Berkley & Hand, 1978-=-; Berkley et al., 1986; Wiberg & Blomqvist, 1984; Wiberg et al., 1987). In turn the tectum projects to the nuclei of the pons and midbrain (Shammah-Lagnado, Negrao, Silva, & Ricardo, 1987; Cornwall, C...

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...he pons and midbrain (Shammah-Lagnado, Negrao, Silva, & Ricardo, 1987; Cornwall, Cooper, & Phillipson, 1990). Another motor-related channel to the upper brainstem is via the cerebellum (Brodal, 1959; =-=Boivie, 1988-=-; Rathelot & Padel, 1997). Some other nuclei in the brainstem, such as the J. Parvizi, A. Damasio / Cognition 79 (2001) 135±159150 lateral reticular nucleus, also receive motor related projections dir...

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...us cortical regions (Brodal, 1959; Jones & Leavitt, 1974; Edwards & de Olmos, 1976; Jackson & Crossman, 1983; Kaufman & Rosenquist, 1985; Steriade, Pare, Parent, & Smith, 1988; Lavoie & Parent, 1994; =-=Groenewegen & Berendse, 1994-=-; Newman & Ginsberg, 1994). The deep mesencephalic nucleus and, to a lesser extent, the nucleus pontis oralis project to the basal forebrain, from which widespread cholinergic projections arise aimed ...

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...nce of noxious stimuli (e.g. Schaible and Schmidt, 1983). Moreover, only a portion of cells in the super®cial dorsal horn of the spinal cord are speci®c to noxious stimuli (Zhang, Han, & Craig, 1993; =-=Han, Zhang, & Craig, 1998-=-). Other studies have con®rmed that there are both nociceptive and non-nociceptive C-®bers (e.g. Vallbo et al., 1993; or see Lawson, 1996, for more references). As Craig has suggested, the ascending p...

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...or sends out primary efferents to, a speci®c cranial nerve. The fact that the brainstem has a nuclear organization was established more than a century ago (e.g. KoÈlliker, 1854; RamoÂn y Cajal, 1894; =-=Jacobsohn, 1909-=-). However, due to the lack of techniques such as immunohistochemical markers, tracing agents, and novel neurophysiological probes, many brainstem nuclei were de®ned on the basis of cytoarchitectural ...

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...ither receives primary afferents from, or sends out primary efferents to, a speci®c cranial nerve. The fact that the brainstem has a nuclear organization was established more than a century ago (e.g. =-=KoÈlliker, 1854-=-; RamoÂn y Cajal, 1894; Jacobsohn, 1909). However, due to the lack of techniques such as immunohistochemical markers, tracing agents, and novel neurophysiological probes, many brainstem nuclei were de...

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...®c to noxious stimuli (Zhang, Han, & Craig, 1993; Han, Zhang, & Craig, 1998). Other studies have con®rmed that there are both nociceptive and non-nociceptive C-®bers (e.g. Vallbo et al., 1993; or see =-=Lawson, 1996-=-, for more references). As Craig has suggested, the ascending pathways from lamina I and the caudal spinal trigeminal subnucleus should be considered introceptive rather than only nociceptive (Craig, ...

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... noxious stimuli, recent evidence suggests that C-®bers are also involved in detecting changes in pH, pCO2, pO2, glucose concentration, osmolarity and in signaling the presence of in¯ammatory agents (=-=Moskowitz, 1991-=-; MacIver & Tanelian, 1992; Burnstock & Wood, 1996; see Craig, 1997, for more references). Thus these ®bers carry signals related to the internal state of the organism. Contrary to the traditional vie...

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...clei including the reticular nucleus of the thalamus (Pare, Smith, Parent, & Steriade, 1988; Steriade, McCormick, & Sejnowski, 1993), and to basal forebrain regions such as the substantia innominata (=-=Muller, Lewandowski, & Singer, 1993-=-). The reticular nucleus of the thalamus projects to other thalamic nuclei (Scheibel & Scheibel, 1966), and inhibits their activity (Steriade & Deschenes, 1984; Barth & MacDonald, 1996), thereby funct...