Reward expectancy in primate prefrental neurons

“…1992; Tremblay and Schultzstimuli, the further behavioral improvement might be mainly due to the acquisition of predictive processing of spatial 1995), and 4) modulations of behavior-related activations by predicted reward (Hollerman et al. 1994; Watanabe 1990,positions by other neuronal systems.”

“…of value-based decisions. been described in orbitofrontal and cingulate cortex and the basal ganglia, areas of the brain traditionally associated with reward-seeking behavior (Kawagoe et al. 2004; Lauwereyns et al. 2002; McCoy et al. 2003; Schultz 1992, 1999, 2000; Watanabe 1996; Watanabe et al. ).”

“Furthermore, an arbitrary sensory cue elicits greater responses from dorsolateral prefrontal neurons when it signals the impending delivery of a preferred food item, such as a piece of apple or cabbage, than when it signals a less favourite food such as a raisin (Watanabe, 1996).”

“…and amounts of food and Given the central importance of rewards for survival, reproduction, and competitive gains, it may not be sur-liquid rewards (Watanabe, 1996; Leon and Shadlen, 1999; Tremblay and Schultz, 1999; Hassani et al., 2001) prising that several specialized and only partly…”

“…supplementary eye field (Amador et al. 2000, Apicella et al. 1992, Cromwell & Schultz 2003, Hikosaka et al. 1989, Hollerman et al. 1998, Pratt & Mizumori 2001, Schoenbaum et al. 1998, Schultz et al. 1992, Shidara & Richmond 2002, Tremblay & Schultz 1999, 2000a, Watanabe 1996, Watanabe et al. 2002).”

“…the specific stimulus, movement, and delay-related changes in the millisecond and second range in prefrontal, parietal, and striatal neurons (Watanabe 1996, Hollerman et al. 1998, Platt & Glimcher 1999), suggesting that influences of rewards on task-specific neuronal processes have…”

“Eachfor employing affective information in biasing behavior subject participated in one of three similarly designed experiments(Watanabe, 1996; Davidson and Irwin, 1999).”

“The PFC is likely to play a role in affective working memory, a process critical to anticipating future affective outcomes (see, e.g., Watanabe, 1996).”

“For example, the gain of delay-period activity of spatially tuned units in dorsolateral PFC is modulated by the type of reward anticipated on a particular trial (Watanabe, 1996).”

“It is likely that the PFC plays a key role in this process (see e.g., Watanabe 1996).”

“DA projections from the ventral tegmental area (VTA) convey information on reward value to the prefrontal cortex (PFC; Watanabe, 1996, Tremblay and Schultz, 1999).”

“They are activated by stimuli from all sensory modalitie”

“For example, work by Watanabe and colleagues (Watanabe 1996, Watanabe et al. 2002) showed that PFC neurons involved in working memory display tonically www.annualreviews.org • Motivation and Cognitive Control 97 A nn u.”

“Similar activities have been reported for the dorsolateral prefrontal cortex (Watanabe, 1996).”

“…that reward-selective neurons were also in DLPFC (Amemori & Sawaguchi 2006, Hikosaka & Watanabe 2000, Kobayashi et al. 2002, Leon & Shadlen 1999, Watanabe 1996), and DLPFC neurons showed similar responses to OFC neurons during the performance of stimulus-reward reversal tasks (Wallis & Miller…”

“The dorsal striatum responds to magnitude of primary rewards and reward prediction (Leon & Shadlen, 1999; Rogers et al., 1999) and shows sustained phasic activations during reward expectation (Watanabe, 1996).”

“This is consistent with previous monkey neurophysiology studies that have found neurons in the DLPFC that encode aspects of the decision or the incentive value of stimuli (Watanabe, 1996; Wallis and Miller, 2003).”

“In these tasks, the monkey must keep " in mind " , or actively maintain, a representation of the target stimulus [or of the anticipated probe stimulus (Rainer et al. 1999; Watanabe 1996)] over a short delay.”

“Brain structures activated in decision making and modulated by reward signals include caudate (Samejima et al., 2005; Hikosaka et al., 2006), lateral parietal cortex area LIP (Platt and Glimcher, 1999; Sugrue et al., 2004), and prefrontal cortex (Watanabe, 1996; Roesch and Olson, 2003).”

“It is also consistent with single unit recordings from nonhuman primates demonstrating that the activity of neurons in the principal sulcus depends on the motivational context of a task (Watanabe et al., 2002) and increases during WM delays in anticipation of a preferred reward (Watanabe, 1996).”

“And, indeed, a wide range of operations integral to WM, including preparation, inhibition, or sequencing of motor responses and context-contingencies, have been consistently observed in DL cortex (Funahashi et al. 1993b, 1997; Williams and Goldman-Rakic 1995; Watanabe 1996).”

“However, there have been few physiological studies that manipulated the outcome of an action in terms of reward (its amount or kind) while keeping the subject's actions constan”

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“Research with primates suggests that neurons within the dorsolateral prefrontal cortex are selectively activated when a reward is expected following a behavioral response (Watanabe, 1996).”

“V(s(t)) V(s(t-1)) are dependent on the amount and type of the reward associated with the memorized items (Watanabe, 1996; Tremblay & Schultz, 2000).”

“…Fuster, 1999; Carlson, Rämä, Tanila, Linnankoski, & Mansikka, 1997; Fuster et al., 1982) and/or the reward expected or resulting from motor action (Ichihara-Takeda & Funahashi, 2008; Hikosaka & Watanabe, 2000; Schultz, Tremblay, & Hollerman, 2000; Watanabe, 1996; Rosenkilde, Bauer, & Fuster, 1981).”

“After the pioneering study by Watanabe [21] on reward-discriminating prefrontal activity, this work makes the next step by investigating the actual gain in neural information derived from the expectation of reward.”

“The participation of the dorsolateral prefrontal cortex (DLPFC) in WM is well established (Friedman and Goldman-Rakic 1994; Petrides et al 1993b) and most neuroimaging studies of WM in schizophrenia demonstrate aberrant DLPFC activation (Manoach et al 1999b; Weinberger and Berman 1996).”

“Neurophysiological studies using spatial working memory tasks such as the delayed-response task have revealed that many neurons in the prefrontal cortex exhibit delay-period activity (Fuster, 1973; Niki, 1974; Niki and Watanabe, 1976; Kojima and Goldman-Rakic, 1982, 1984; Funahashi et al., 1989; Watanabe, 1996; Carlso…”

“Moreover, OFC activity is correlated with the preference ranking of different rewards (Arana et al., 2003; Tremblay & Schultz, 1999; Watanabe, 1996) and the combination of reward preference and magnitude (Padoa-Schioppa & Assad, 2006, 2008).”

“(Watanabe, 1996) and actions (Quintana and Fuster, 1992; Asaad”

“Several recent studies have extended the pioneering work by examining the role of PF in processing information about stimuli, responses, rewards, errors, and other behaviorally significant events (Hasegawa et al. 1998; Lecas 1995; Miller et al. 1996; Rao et al. 1997; Rolls and Baylis 1994; Rolls et al. 1996a, 1996b; S…”

“This association is consistent with the known role of the OFC in processing expectation of food reward (Watababe, 1996).”

“The integration of these signals reflects more than just additive contributions of cognitive demands and reward value, but actually enhances functional coding within PFC, such as by maximizing signal-to-noise ratio, enhancing discriminability of visuospatial signals, and increasing the amount of information transmitte…”

“…pation of the expected stimulus, action, or reward and structuring information flow among different brain (Asaad et al., 1998; Rainer et al., 1999; Watanabe, 1996), regions (Diesmann et al., 1999; Salinas and Sejnowwhereas activity early during the delay tends to reflect ski, 2001). retrospective…”

“It receives highly processed information from several brain areas and is involved in complex cognitive and behavioral processes (Bubser and Schmidt, 1990; Granon et al., 1994; Kolb, 1984; Muir et al., 1996; Watanabe, 1996).”

“In monkeys performing spatial-delayed response tasks, DLPFC neurons involved in retaining information in WM are responsive to reward; they show more activity during delay periods when a preferred reward is anticipated than when it is not (Watanabe 1996).”

“Allrelated sustained activity in a spatial delayed response task (Watanabe 1996).”

“These activations can be dependent on the reward type (Watanabe 1996) or magnitude (Leon and Shadlen 1999).”

“…cognitive control enhancements that occur with the provision of incentives in both animals (Kobayashi et al., 2006; Krawczyk et al., 2007; Sakagami and Watanabe, 2007; Watanabe, 1996), and humans (Beck et al., 2010; Jimura et al., 2010; Savine and Braver, 2010; Tsujimoto and Sawaguchi, 2005).”

“with several fixed rewards — has been already described in structures interconnected with the ACC, including prefrontal cortex (Watanabe, 1996; Leon and Shadlen, 1999; Hikosaka and Watanabe, 2000; Watanabe et al., 2002; Wallis and Miller, 2003), parietal cortex (Platt and Glimcher, 1999; Sugrue et al.”

“Reward Expectancy-related Neuronal Activity in the LPFC To compare characteristics of delay activity of OFC and LPFC neurons in relation to the response outcome, we introduce here reward expectancy-related neuronal activities of the LPFC which were reported previously (Watanabe, 1996).”

“Previous studies have found that the DLPFC might provide several different types of signals necessary for updating the value functions during the matching pennies task used in the present study (Barraclough et al., 2004; Seo et al., 2007).”

“frontal neurons (Watanabe, 1996) are preferentially activated by”

“It receives highly processed information from several brain areas, and is involved in complex cognitive and behavioral processes (Bubser and Schmidt, 1990; Granon et al., 1994; Kolb, 1984; Muir et al., 1996; Watanabe, 1996).”

“More recently (Sutton & Davidson 1997) we showed that scores on a self-report measure designed to operationalize Gray’s concepts of the BIS/BAS scales (Carver & White 1994) were even more strongly predicted by electrophysiological measures of prefrontal asymmetry than were scores on the PANAS (see figure 2).”