Reward expectancy in primate prefrental neurons

“…studies have identified neural signals related to reward values in different areas of the primate brain, including the lateral prefrontal cortex (Watanabe, 1996; Leon and Shadlen, 1999; Kobayashi et al., 2002; Barraclough et al., 2004; Seo et al., 2007), the orbitofrontal cortex (Tremblay and…”

“Such sustained activity is often specific to a particular type of task-relevant information such as stimuli, locations, rewards, and even abstract task rules (Watanabe 1996; Asaad et al. 1998; Rainer et al. 1998; Wallis et al. 2001).”

“It is also known that LPFC neurons encode reward value (Amemori and Sawaguchi, 2006b; Kim et al., 2008) and show differential activity depending on whether or not a monkey expects and/or obtains a more preferable outcome (Watanabe, 1996; Leon and Shadlen, 1999; Hikosaka and Watanabe, 2000).”

“Cells coding for expected reward outcomes are found in orbitofrontal cortex (OFC), and in regions of the PFC and amygdala [11,48,103,133,137,145,153], i.”

“On the other hand, reward expectations can apparently also be processed in a qualityspecific manner (e.g., Dickinson and Balleine, 1994; Watanabe, 1996).”

“Although PFC neurons represent both received and predicted rewarding outcomes in monkeys (Watanabe, 1996; Mansouri et al., 2006; Seo et al., 2007; Received Feb. 16, 2017; revised June 2, 2017; accepted June 9, 2017.”

“The expected value of a reward has been shown to modulate activity during the performance of a task (Ikeda and Hikosaka 2003; Platt and Glimcher 1999; Shidara and Richmond 2002; Watanabe 1996).”

“Electrophysiological studies identified NCL neurons in that—comparable to PFC neurons—show delay- and reward expectancy-related activity (Fuster, 1989; Kalt, Diekamp, & Güntürkün, 1999; Watanabe, 1996).”

“Neurons in primate dorsolateral prefrontal cortex also modulate with reward expectancy (Watanabe, 1996).”

“As in the work of Husbands et al. (1998), the diffusion of neuromodulators was evolved together with the network architecture.”

“Activity in the dlPFC is enhanced especially in response to motivationally intriguing goals (Watanabe, 1996; Amemori and Sawaguchi, 2006; Kennerley and Wallis, 2009).”

“The dlPFC, on the other hand, represents the expected rewards (48) and the task rules (49).”

“…include coding of reward expectancy or registering the lack of an expected reward (when seen on incorrect trials) comparable with results from visual memory tasks in PFC (Watanabe, 1996; Leon and Shadlen, 1999; Hikosaka and Watanabe, 2000; Wallis and Miller, 2003; Kennerley and Wallis, 2009).”

“Several studies have suggested that single neurons in the PFC show directional delay-period activity that is modulated by predicted reward (Watanabe, 1996; Leon and Shadlen, 1999; Kobayashi et al., 2002).”

“…and rodents have revealed that PFC neurons are responsive during goal-directed behaviors for liquid or food rewards (Niki and Watanabe, 1976, 1979; Watanabe, 1996; Tremblay and Schultz, 1999, 2000; Watanabe et al., 2002; Schoenbaum et al., 1998; Takenouchi et al., 1998; Hikosaka and Watanabe,…”

“…for optimal decision making such as sensory evidence (Romo and Salinas, 2003; Heekeren et al., 2004; Philiastides et al., 2011), categories (Seger and Miller, 2010), rules (Wallis et al., 2001), and previous choices and rewards (Watanabe, 1996; Barraclough et al., 2004; Lee and Seo, 2007).”

“Second, the PFC plays an important role in decision making by encoding a particular state of the environment and the desirability of the outcome expected from such a state (13, 14).”

“, 2012) and is active during the presentation of reward or reward-predictive cues (Watanabe, 1996; Matsumoto et al., 2007; Moorman and Aston-Jones, 2015; Pinto and Dan, 2015).”

“…et al. 2000; Coe et al. 2002), dorsolateral prefrontal cortex (Kobayashi et al. 2002, 2007; Leon and Shadlen 1999; Tsujimoto and Sawaguchi 2005; Watanabe 1996), parietal cortex (Platt and Glimcher 1999; Sugrue et al. 2004), caudate nucleus (Cromwell and Schultz 2003; Hollerman et al. 1998;…”

“, 2000), expected reward (Watanabe, 1996; Leon and Shadlen, 1999; Kobayashi et al., 2002; Watanabe et al., 2002), numerical quantity (Nieder et al.”

“This might be expected given the role of prefrontal cortex in reward expectancy (Watanabe, 1996).”

“…cortex may evaluate the subjective values of various objects (Padoa-Schioppa 2011), whereas the dorsolateral prefrontal cortex may be more involved in selecting the most appropriate actions and monitoring their outcomes (Watanabe 1996; Barraclough et al. 2004; Seo and Lee 2009; Kim et al. 2009).”

“This could be useful in modulating network response as a function of attention (Reynolds and Chelazzi, 2004) or reward (Schultz et al., 1993; Watanabe, 1996; Leon and Shadlen, 1999; Amemori and Sawaguchi, 2006).”

“It even might have monitored the outcome of goal-directed behaviour (Watanabe, 1996).”

“…cellular activity in the prefrontal cortex during cue-presentation and/or delay period activity can convey information about the upcoming reward (Watanabe, 1986, 1996; Watanabe et al., 2002; Leon and Shadlen, 1999; Tremblay and Schultz, 2000; Kobayashi et al., 2002; Wallis and Miller, 2003),…”

“basis of reward prediction delivered by the PFC (Watanabe, 1996).”

“It is possible that specific outcome expectancies were private events such as differential secretion of salivation (Goeters et al., 1992) or central neuron activities (cf. Watanabe, 1996).”

“In the present study, rabies injections weremade into area 46 of the prefrontal cortex of macaques that is characterized by cognitive functions, such as reward (13), spatial working memory (14), temporal processing (15, 16), evaluation of self-generated decision making (17), categorization (18), and motor learning of…”

“…such as the prefrontal cortex, may provide the signals related to the actions chosen by the animal and their value functions (Baeg et al. 2003; Barraclough et al. 2004; Lee et al. 2007; Leon and Shadlen 1999;Watanabe 1996), whereas dopamine neurons provide reward prediction errors (Schultz 1998).”

“There is evidence that the PFC integrates motivational and cognitive processes, since reward expectations increasedWMmaintenance in the lateral PFC in macaques (Leon & Shadlen, 1999; Watanabe, 1996).”

“REVIEW ARTICLE published: 22 November 2011 doi: 10.3389/fnins.2011.00124 Challenges of interpreting frontal neurons during value-based decision-making”

“Recent neurophysiological work suggests that this kind of expectation is established by activation of the prefrontal cortex (Watanabe, 1996), an area known to play an essential role in working memory.”

“Single-unit recordings from orbital prefrontal cortex reveal neuronal firing associated with stimulus and reward value (Watanabe, 1996; Roesch and Olson, 2004).”

“…studies have shown changes of visual responses by the expectation of reward in many brain areas (Apicella et al. 1992; Hikosaka et al. 1989b; Kawagoe et al. 1998; Kobayashi et al. 2002; Leon and Shadlen 1999; Platt and Glimcher 1999; Schultz et al. 1992; Tremblay and Schultz 2000; Watanabe 1996).”

“The directional activities of a subset of LPFC neurons are modulated by the amount of reward expected at the end of the trials (Watanabe, 1996; Leon and Shadlen, 1999; Kobayashi et al., 2002).”

“Additional evidence indicates that firing rates of PFC neurons during the response and reward phase of these tasks may reflect decision-related or reward/response outcome evaluation [12,13,16,17].”

“I N T R O D U C T I O N Graded neuronal activity that steadily increases or decreases during a delay period is typically seen in the prefrontal cortex when animals are anticipating sensory cues (Rainer et al. 1999), motor responses (Hoshi et al. 2000), or rewards (Watanabe 1996).”

“In monkeys, expectancy of a reward subsequent to a behavioral response was preferentially associated with activity of dorsolateral prefrontal cortex (DLPFC) neurons (Watanabe, 1996).”

“More broadly, neural coding in the sensory cortex is sensitive to the effect of reward [51 – 53].”

“Although a number of studies had previously investigated neural responses to reward expectation (Watanabe, 1996; Schultz et al., 1997), the first experiment to record probability-related activity of single neurons from an economic point of view was probably conducted by Platt and Glimcher (1999).”

“Although PFC delay activity is thought to represent a memory trace, PFC neurons have also been shown to alter their firing rates on tasks that impose little or no memory demand, such as immediate reward tasks (Watanabe, 1996).”

“…the domain featuring the full range of motivational, hedonic, state-, task-, and goal-related biasses (Jacobsen, 1935; Nauta, 1971; Fuster, 1980; Watanabe, 1996; Drevets and Raichle, 1994; Wallis and Rolls, 1997; Price, 1999; Asaad et al., 2000; Rolls, 2000; Liu et al., 2000; Miller and Cohen,…”

“17 Watanabe, M. (1996) Reward expectancy in primate prefrontal”

“Previous studies have shown that neurons in the PFC participate in many aspects of cognitive behaviors based on reward [29], evaluating self-generated decisions [30], categorization [31], procedural learning [32], functional separation of ‘‘what’’ and ‘‘when’’ [33], and time prediction and detection [34].”

“Since reward plays a central role in shaping behavior [6], its representation reflects the basis of goal-directed actions [58].”

“First, neurons in the DLPFC encode the value of expected reward (Roesch and Olson, 2003; Kennerley and Wallis, 2009), often for specific actions (Watanabe, 1996; Leon and Shadlen, 1999; Barraclough et al., 2004; Kim et al., 2008).”

“However, in macaques, cells in the LPFC have been shown to change activity during delay periods in tasks in which food rewards were visible or associated but not seen (Watanabe, 1996; Hikosaka and Watanabe, 2000).”

“, 1998), in which the prefrontal cortex plays a crucial role (Rainer et al., 1999; Watanabe, 1996).”

“…while other studies have reported that DLPFC neurons in monkeys show activity related to reward and/or expectancy of reward (Niki and Watanabe, 1979; Watanabe, 1989, 1996; Leon and Shadlen, 1999; Kobayashi et al., 2002; Watanabe et al., 2002; Wallis and Miller, 2003b; Roesch and Olson, 2003).”

“Sim - ilarly , activity of neurons coding the metrics of a single move - ment is often modulated by expected reward in various cortical and subcortical areas ( Watanabe , 1996 ; Kawagoe et al .”

“For example, data from recent fMRI studies examining cognitive control across a range of task domains have confirmed earlier single-unit recording studies in primates (Leon & Shadlen, 1999; Watanabe, 1996) indicating that specific regions within lateral prefrontal cortex (and some associated regions, including the ACC…”

“Single neurons in the two areas were able to encode reward information related to the reward type, amount and availability that was indicated by conditioned visual stimuli (Watanabe 1996; Kawagoe et al. 1998; Kobayashi et al. 2006).”

“It is known that the prefrontal cortex (PFC) and the striatum are involved in reward prediction processes (Watanabe, 1996; Hollerman et al., 1998; O’Doherty et al., 2003a; Roesch and Olson, 2003; Samejima et al., 2005).”

“…about outcomes, whereas dorsolateral premotor cortex (Okano and Tanji, 1987; Romo and Schultz, 1987; Kurata and Wise, 1988), prefrontal cortex (Watanabe, 1996), and distinct regions of the striatum (Schultz and Romo, 1988) have been implicated in translating this temporal information into…”

“…varies over the course of a delay period, often ramping up or down (Kojima and Goldman-Rakic, 1982; Quintana and Fuster, 1992; Miller et al., 1996; Watanabe, 1996; Heinen and Liu, 1997; Chafee and Goldman-Rakic, 1998; Kawagoe et al., 1998; Rainer et al., 1999; Hassani et al., 2001; Brody et al.,…”

“nonpreferred) [41,47].”