Dorsolateral prefrontal cortex: comparative cytoarchitectonic analysis in the human and the macaque brain and corticocortical connection patterns

“…defined areas in non-human primates have also been established in the human brain using comparative high-resolution fMRI (see glossary; Baker, Patel, Corbetta, & Snyder, 2005; Koyama et al., 2004; Orban et al., 2006) and cytoarchitectonic (Petrides & Pandya, 1999, 2002) analyses.”

“These prefrontal areas, which in accordance with maps delineated by Petrides and Pandya (1999) would be classified as areas 9 and 8B, are dorsal to the areas 9/46d, 9/46v, 8Ad and 8Av along the middle frontal gyrus, parts of which are customarily referred to as the dorsolateral prefrontal cortex…”

“Although macaque and human brains share a high degree of similarity in terms of cytoarchitecture (Petrides and Pandya, 1999, 2002a; Ongür et al., 2003), functional organization (Rees et al., 2000; Koyama et al., 2004; Petrides et al., 2005; Nakahara et al., 2007), and anatomical connections…”

“It is well established that macaque and human brains share a high degree of similarity in terms of cytoarchitecture (Petrides and Pandya, 1999, 2002; Ongür et al., 2003), functional organization (Rees et al., 2000; Koyama et al., 2004; Petrides et al., 2005; Nakahara et al., 2007), anatomical…”

“1 stainings in good agreement with (87).”

“PFC is not a homogeneous structure but is composed of multiple areas that differ in terms of cytoarchitecture and anatomical connections with other areas (Petrides and Pandya, 1999, 2002; Figure 3).”

“However, several connectional studies in experimental animals concluded that the arcuate fasciculus is not related to language (105, 118).”

“In contrast, VLPFC, which connects with high-level visual areas in the temporal lobe (Barbas, 1988; Carmichael and Price, 1995b; Petrides and Pandya, 1999), should maintain information about pictures and objects (Wilson et al., 1993).”

“2).(73,74) The MDLFC occupies the middle frontal and superior frontal gyri and comprises cytoarchitectonic areas 46 and 9/46 (middle frontal gyrus) and area 9 (superior frontal gyrus).”

“…speculated that based on the different connectivity that the two brain regions have, the multisensory nature of OFC could point to a role in the immediate valuation of items (Rolls, 2000, 2004), while the dlPFC could bemore involved in the execution of choice behavior (Petrides and Pandya, 1999).”

“Rhesus monkeys are an ideal animal model for studying normal aging for many reasons: (1) their brain anatomy closely resembles that of humans, (2) they are vulnerable to age-related cognitive decline, (3) they can perform cognitive tests similar to those administered to humans, and (4) they do not develop neurodegener…”

“…primates (Jacobson and Trojanowski, 1977; Mesulam et al., 1977; Seltzer and Pandya, 1984, 1989, 1994; Webster et al., 1994; Pandya and Yeterian, 1996; Petrides and Pandya, 1999), diffusion tensor imaging studies in humans (Catani et al., 2002, 2004), and the functional and effective connectivity…”

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“This suggests that communication between the MFG and FFA, while most likely polysynaptic (Petrides and Pandya 1999), may make a functional contribution to the task in a loadindependent manner.”

“Our interpretatio hat episodic memory failure in MCI-WMH subjects is s ndary to a more general impairment in executive con rocesses. rontal, retrosplenial, hippocampal circuit may also give o deficits observed in MCI (Morris, Pandya, & Petride 999; Petrides & Pandya, 1999).”

“Area V4 was chosen because it is considered to be a sensory visual area at an intermediary processing stage in the ventral stream and because it is directly connected to parts of the PF cortex (Petrides and Pandya 1999).”

“…hippocampal formation output to lateral prefrontal cortex (Insausti & Munoz, 2001) and the pattern of cortical input to these areas (Insausti et al., 1987; Suzuki & Amaral, 1994a; see Figure 5 in Barbas et al., 1999; Petrides & Pandya, 1999; Petrides & Pandya, 2002; Blatt et al., 2003).”

“DLPFC is important for working memory (Petrides and Pandya, 1999; Muller and Knight, 2006).”

“Specificity for Language 843 lobes (Petrides and Pandya 1999, 2001; Nelissen et al. 2005), the alternative view, whereby brain regions have very broad functions and each region can perform multiple types of computations, is consistent with the connectivity patterns, which suggest extensive sharing…”

“lateral—sensory and motor cortex; orbital—limbic), however, all of these sub-regions are so densely interconnected that any and all information is quickly integrated across sub-regions [41–43].”

“Taken together, these data suggest that DA receptor activa- tion increases interneuron excitability and spontaneous spike firing via a D1 receptor mechanism without altering action potential--independent release of GABA.”

“This is probably not surprising, since the DLPFC is heavily interconnected with the ACC (Mayberg et al., 2005; Petrides and Pandya, 1999).”

“The connections in this model were based on known primate and rodent anatomy (Amaral et al., 2003; Burman et al., 2009; Petrides & Pandya, 1999; Rosene and Van Hoesen, 1977) and were further refined to ensure stability of the model.”

“Identification of frontal areas of the human (A) and macaque monkey (B) functional FEF network based on cytoarchitectonic parcellation (Petrides and Pandya 1999, 2002).”

“…environment and internal milieu, the primate frontal cortex, with its close anatomical connections with high-order sensorimotor cortical areas (Petrides and Pandya, 1999, 2002) and subcortical limbic system (Öngür and Price, 2000), is likely to be involved in various aspects of decision…”

“The model was based on known anatomical connections in human and non-human primates (Middleton and Strick, 1994, 2000, 2001; Petrides and Pandya, 1999; Clower et al., 2001) and on data from published functional imaging studies (Kim et al., 1994; Andreasen et al., 1998).”

“Nevertheless, parts of the caudal PFC, such as the frontal eye fields, also receive information from the “ventral visual stream,” which processes colors, shapes, and textures of visual stimuli (Petrides & Pandya, 1999; Webster, Bachevalier & Ungerleider, 1994).”

“In addition to the caudal-rostral axis, there is a dorsal-ventral axis of organization of the mid-lateral PFC (DLPFC: BA46 and BA9/46; Petrides and Pandya, 1999; Petrides, 2005b).”

“Besides the widely discussed homology between area F5 and Broca’s area (see Von Bonin and Bailey 1949; Petrides and Pandya 1999 for reviews; also Rizzolatti et al. 2001), the functional analogy is stressed best by the finding that Broca’s area is activated during hand movements (Krams et al. 1998; Binkofski et al. 199…”

“Instead it derives its cortical inputs from the parietal and temporal cortex [Petrides and Pandya, 1999, 2002].”

“…strong anatomical evidence for reciprocal connections between these regions based on studies in nonhuman primates (Bates and Goldman-Rakic, 1993; Petrides and Pandya, 1999), a natural hypothesis is that monitoring-related activity in dmPFC engages regulatory processes in the dlPFC that shape…”

“The left IFS (extending onto MFG), a subregion of DLPFC, was the only region showing increased effective connectivity with face and house regions that (1) was modulated by sensory evidence levels during the perceptual decision; (2) showed greatest activity during the decision stage; and (3) was activated above baselin…”

“Others, based on monkey studies, have argued that the border zone is constituted by the middle frontal sulcus (as the analogue of the monkey’s sulcus principalis)—a region within the middle frontal gyrus (Petrides and Pandya, 1999; see Fig.”

“…RIFG, LHC and LIPL was based upon known primate neuroanatomy (Clower et al., 2001; Felleman and van Essen, 1991; Insausti et al., 1987; Insausti and Munoz, 2001; Mesulam et al., 1977; Petrides and Pandya, 1984, 1999; Rockland and Van Hoesen, 1999; Schall et al., 1995; Suzuki and Amaral, 1994).”

“Within Brodmann ’ s original work, the SFG was divided into four distinct regions, including areas 6, 8, 9, and 32 (Petrides and Pandya, 1999, 2002).”

“As in the human, macaque area 10 occupies the dorsal, medial, and ventral aspects of the macaque polar region (4, 66, 67).”

“This dissociation in humans has been corroborated in investigations of non-human primates revealing important cytoarchitectural differences between VLPFC and DLPFC [Owen, 1997; Petrides, 1996; Petrides and Pandya, 1999].”

“Because the medial PFC and the lateral PFC have reciprocal dense connections [43–48], these two prefrontal areas are likely to interact to support cognitive functions.”

“The DLPFC, which is connected to the striatum, the anterior cingulate cortex, and the thalamus (Barbas, 2000; Paus et al., 2001; Petrides and Pandya, 1999), is the most common target for rTMS.”

“More simply, Shallice et al. (2008b) took each Petrides and Pandya (1999) region of frontal cortex, and compared the performance of patients with damage to that region with that of all the other patients with frontal lesions.”

“In macaques, the label was in the regions of the LIP (see also Andersen et al., 1985; Petrides and Pandya, 1999) and VIP areas of the intraparietal sulcus, and comparable regions were labeled in owl monkeys and squirrel monkeys, although these regions were in the caudal extension of the lateral…”

“The regional boundaries seen in the coronal plane were then translated into a stretched-out surface view using methodology similar to that of Barbas and Pandya (1987) and Petrides and Pandya (1999).”

“intermediate frontal sulcus is the most likely human homologue of the principal sulcus, areas 46 and 9/46 (Petrides and Pandya, 1999; Rajkowska and Goldman-Rakic, 1995).”

“Our results also indicate that the marmoset has a region equivalent to area 46 described in Old World monkeys and humans (Walker, 1940; Petrides and Pandya, 1999), but not in Galago (Preuss and Goldman-Rakic, 1991a).”

“Petrides M, Pandya DN (1999) Dorsolateral prefrontal cortex: comparative cytoarchitectonic analysis in the human and the macaque brain and corticocortical connection patterns.”

“It is important to note that the human homolog of the monkey sulcus prin-cipalis seems to be the intermediate frontal sulcus (Petrides and Pandya, 1999), which shows large individual variation in folding in the MFG.”

“This finding corroborates the hypothesis of connectivity between dorsal (BA45/46) and ventral aspects (BA47) of the frontal lobes (Petrides and Pandya, 1999).”

“There are anatomical and functional connections between the dorsolateral PFC (DLPFC) and medial PFC, including the ACC and the OFC [23–26].”

“The executive component of working memory is generally associated with Brodmann Areas 9 and 46 in the dorsolateral section of the prefrontal cortex (Gazzaniga, Ivry, and Mangun, 2002; Goldman-Rakic, 1987; Henson, 2001; McCarthy et al., 1994, Petrides and Pandya, 1999; Smith and Jonides, 1999).”

“1 3 The dlPFC is richly connected with a variety of proximal and distant brain areas (Lu et al. 1994; Petrides and Pandya 1999; Miller and Cohen 2001), and is thought to be a key region for information integration (Hoshi 2006).”

“Wisconsin Card Test, perseverations 11 10 13 17 11 16 14 13 (3) 20 (9)”

“in area 8, [79]).”

“Parietal cortex and PFC are reciprocally anatomically connected (Divac et al., 1977; Petrides and Pandya, 1999), and in the non-human primate seem to be involved in a mutually dependent neural circuit during the delay period of delayed response tasks (Chafee and Goldman-Rakic, 2000; Quintana et…”

“[83] Petrides M, Pandya DN.”

“No direct projecions have been described from the inferotemporal corex to the premotor cortex (Boussaoud et al., 1996; Petrides and Pandya, 1999).”

“This is supported by studies of the macaque, in which there are dense connections between lateral RoPFC and caudal DLPFC (Petrides and Pandya, 2007), which in turn is strongly connected with parietal cortex (Petrides and Pandya, 1999).”

“One way in which the relationship between affective and cognitive processing may be facilitated is through reciprocal neural connections between dorsal and ventral prefrontal brain regions (Cavada, Compañy, Tejedor, CruzRizzolo, & Reinoso-Suárez, 2000; Petrides & Pandya, 1999).”

“Reciprocal connections between the DLFC and ACC (Barbas and Pandya, 1989; Preuss and Goldman-Rakic, 1989; Barbas 1992; Petrides and Pandya, 1999) are part of a frontolimbic network (Arikuni et al., 1994; Barbas 2000).”

“The known cytoarchitecture of the insula cortex and its connectivity to the prefrontal cortex have therefore been proposed to support propagation of slow waves between these potential generator regions (Petrides and Pandya, 1999; van der Kooy et al., 1982).”

“The second module is the ventrolateral prefrontal cortex (vlPFC), which co-activates with dlPFC and PPC to coordinate complex cognitive control processes as well as reappraisal [5,12,14,24,37,57,86,87,94].”

“It should be noted that determination of exact anatomical borders of BA 9 and BA 46 is difficult (Petrides & Pandya, 1999) and that other SOAs, ISIs, or another task design might be necessary to probe the BA 9–M1 connection.”