Rethinking the Emotional Brain

“In spite of beingwell known andwidely applied in research, the basic emotions point of view has been challenged on various grounds (e.g., Averill, 1980; Ortony and Turner, 1990; Russell, 1980, 2003; Barrett, 2006a; Barrett et al., 2007).”

“…and recognized in people around the world, conserved in our close animal ancestors, and supposedly hard-wired into brain circuits by evolution (Darwin, 1872; Tomkins, 1962; Ekman, 1972, 1980, 1984, 1992, 1999a, 1999b; Izard, 1992, 2007; Damasio, 1994, 1999; Panksepp, 1998, 2000, 2005; Prinz, 2004).”

“As noted above, when threat levels rise, feeding is suppressed (Gray, 1987; Lima and Dill, 1990; Blanchard et al., 1990; Fanselow, 1994).”

“…1994, 1999; Leventhal and Scherer, 1987; Scherer, 2000; Ortony and Turner, 1990; Öhman, 1986, 2009; Johnson-Laird and Oatley, 1989; Ellsworth, 1994; Zajonc, 1980; Lazarus, 1981, 1991a, 1991b; Barrett, 2006a, 2006b; Barrett et al., 2007; Kagan, 2007; Prinz, 2004; Scarantino, 2009; Griffiths, 2004;…”

“Body feedback has played an important role in emotion theory for more than a century (James, 1884; Lange, 1885/1922; Schachter and Singer, 1962; Tomkins, 1962; Adelmann and Zajonc, 1989; Buck, 1980; Damasio, 1994, 1999).”

“…Scherer, 2000; Ortony and Turner, 1990; Öhman, 1986, 2009; Johnson-Laird and Oatley, 1989; Ellsworth, 1994; Zajonc, 1980; Lazarus, 1981, 1991a, 1991b; Barrett, 2006a, 2006b; Barrett et al., 2007; Kagan, 2007; Prinz, 2004; Scarantino, 2009; Griffiths, 2004; Ochsner and Gross, 2005; Lyons, 1980).”

“A third consequence of survival circuit activation is ‘‘generalized arousal’’ (Moruzzi and Magoun, 1949; Lindsley, 1951; 662 Neuron 73, February 23, 2012 ª2012 Elsevier Inc. Schober et al., 2011; Lang, 1994; Pfaff et al., 2008).”

“Cortisol crosses the blood brain barrier and binds to receptors in a variety of areas, while adrenergic hormones affect the CNS indirectly (McGaugh, 2000).”

“…transcription factors (e.g., cyclic AMP response element binding protein) involved in defense conditioning Aplysia (e.g., Hawkins et al., 2006; Kandel, 2001; Carew and Sutton, 2001; Glanzman, 2010; Mozzachiodi and Byrne, 2010) have been implicated in defense conditioning in the mammalian…”

“Nevertheless, there have from time to time been calls for linking reinforcement more directly to specific neurobiological systems.”

“…behaviorally, involves approach toward desired outcomes and avoidance of undesired outcomes (Tolman, 1932; McClelland et al., 1953; Schneirla 1959, Elliot and Church, 1997; Cofer, 1972; Cofer and Appley, 1964; Miller, 1944; Trowill et al., 1969; Bindra, 1969; Davidson, 1993; Gray, 1982; Lang et…”

“Body feedback has played an important role in emotion theory for more than a century (James, 1884; Lange, 1885/1922; Schachter and Singer, 1962; Tomkins, 1962; Adelmann and Zajonc, 1989; Buck, 1980; Damasio, 1994, 1999).”

“Research on the neural basis of feelings in humans is in its infancy (Panksepp, 1998; 2005; Damasio, 2003; Damasio et al., 2000; Ochsner et al., 2002; Barrett et al., 2007; Rudrauf et al., 2009; Critchley et al., 2004; Pollatos et al., 2007).”

“The evidence for conservation across mammals of mechanisms underlying survival functions such as defense (e.g., LeDoux, 1996, 2012; Phelps and LeDoux, 2005; Motta et al., 2009; Choi et al., 2005; Kalin et al., 2004; Amaral, 2003; Antoniadis et al., 2007), reproduction (e.g., Pfaff, 1999; Oomura et…”

“The discovery that behavior could be reinforced by electrical stimulation of brain areas (Olds and Milner, 1954), and findings that electrical reinforcement could summate with different natural reinforcers (Coons and White, 1977; Conover and Shizgal, 1994), were compatible with a generic mechanism…”

“Other activities, such as eating, drinking, and sex, are actively suppressed (Gray, 1987; Lima and Dill, 1990; Blanchard et al., 1990; Fanselow, 1994; Choi et al., 2005).”

“…1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et al., 2006; Nestler, 2004; Kelley, 2004).”

“Research on the neural basis of feelings in humans is in its infancy (Panksepp, 1998; 2005; Damasio, 2003; Damasio et al., 2000; Ochsner et al., 2002; Barrett et al., 2007; Rudrauf et al., 2009; Critchley et al., 2004; Pollatos et al., 2007).”

“…et al., 2007), reproduction (e.g., Pfaff, 1999; Oomura et al., 1988; Blaustein, 2008), thermoregulation (Nakamura and Morrison, 2007), fluid balance (Johnson, 2007; Fitzsimons, 1979), and energy/nutritional regulation (Elmquist et al., 2005; Morton et al., 2006; Saper et al., 2002) is strong.”

“The anticipatory/exploratory/search phase is guided by incentives (Bindra, 1968; Trowill et al., 1969; Balleine and Dickinson, 1998; Cardinal et al., 2002; Johnson et al., 2009; Petrovich et al., 2002; Berridge, 1999, 2007, 2004; Rolls, 1999, 2005; Glimcher, 2003).”

“The nucleus accumbens emerged as a key focal point of this general motivational system (Graybiel, 1976; Mogenson et al., 1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009;…”

“Themodern paradigmatic example of a generic reinforcement mechanism is the role of dopamine in the striatum as a reward prediction error signal (Schultz, 1997).”

“It is therefore of great interest that recent work on the role of dopamine as a reward prediction error signal is beginning to recognize the importance of specific motivational states in modulating the effects of dopamine as a reward prediction error signal (Schultz, 2006; Glimcher, 2011).”

“Research on the neural basis of feelings in humans is in its infancy (Panksepp, 1998; 2005; Damasio, 2003; Damasio et al., 2000; Ochsner et al., 2002; Barrett et al., 2007; Rudrauf et al., 2009; Critchley et al., 2004; Pollatos et al., 2007).”

“…is deliberate, controlled and often conscious, determines stimulus meaning and predisposes actions (e.g., Arnold, 1960; Bowlby, 1969; Frijda, 1986; Lazarus, 1991a, 1991b; Leventhal and Scherer, 1987; Lazarus and Folkman, 1984; Smith and Ellsworth, 1985; Scherer, 1988; Scherer et al., 2001; Sander…”

“…and Davidson, 1994; LeDoux, 1996; Panksepp, 1998, 2000, 2005; Rolls, 1999, 2005; Damasio, 1994, 1999; Leventhal and Scherer, 1987; Scherer, 2000; Ortony and Turner, 1990; Öhman, 1986, 2009; Johnson-Laird and Oatley, 1989; Ellsworth, 1994; Zajonc, 1980; Lazarus, 1981, 1991a, 1991b; Barrett,…”

“With each subsequent decade, small increases resulted, until the last decade, when emotion titles grew exponentially—more than 2,000 hits.”

“…behavior, in spite of discussion and debate that dates back to the earliest days in modern biology and psychology (e.g., Darwin, 1872; James, 1884; Cannon, 1927, 1931; Duffy, 1934, 1941; Tomkins, 1962; Mandler, 1975; Schachter, 1975; Ekman, 1980, 1984, 1992; Izard, 2007; Frijda, 1986; Russell,…”

“…region; PMH, premammilary nucleus of the hypothalamus. the medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998; Paré et al., 2004; Maren and Quirk,…”

“This debate is centered on the question of whether emotions are ‘‘natural kinds,’’ things that exist in nature as opposed to being inventions (constructions) of the human mind (e.g., Panksepp, 2000; Griffiths, 2004; Barrett, 2006a; Izard, 2007; Scarantino, 2009).”

“…et al., 2007), reproduction (e.g., Pfaff, 1999; Oomura et al., 1988; Blaustein, 2008), thermoregulation (Nakamura and Morrison, 2007), fluid balance (Johnson, 2007; Fitzsimons, 1979), and energy/nutritional regulation (Elmquist et al., 2005; Morton et al., 2006; Saper et al., 2002) is strong.”

“…et al., 2007), reproduction (e.g., Pfaff, 1999; Oomura et al., 1988; Blaustein, 2008), thermoregulation (Nakamura and Morrison, 2007), fluid balance (Johnson, 2007; Fitzsimons, 1979), and energy/nutritional regulation (Elmquist et al., 2005; Morton et al., 2006; Saper et al., 2002) is strong.”

“Modulators also facilitate processing areas involved in retrieving forming, and storing memories (McGaugh, 2003; Roozendaal et al., 2009).”

“In such cases of stimulus-response habit formation, the neural control switches from the ventral to the dorsal striatum (Everitt and Robbins, 2005;Wickens et al., 2007; Packard and Knowlton, 2002).”

“This debate is centered on the question of whether emotions are ‘‘natural kinds,’’ things that exist in nature as opposed to being inventions (constructions) of the human mind (e.g., Panksepp, 2000; Griffiths, 2004; Barrett, 2006a; Izard, 2007; Scarantino, 2009).”

“…medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998; Paré et al., 2004; Maren and Quirk, 2004; Quirk and Mueller, 2008; Haubensak et al., 2010).”

“that recent work on the role of dopamine as a reward prediction error signal is beginning to recognize the importance of specific motivational states in modulating the effects of dopamine as a reward prediction error signal (Schultz, 2006; Glimcher, 2011).”

“It is therefore of great interest that recent work on the role of dopamine as a reward prediction error signal is beginning to recognize the importance of specific motivational states in modulating the effects of dopamine as a reward prediction error signal (Schultz, 2006; Glimcher, 2011).”

“…to generalized arousal are peripheral systems that release hormones into the circulation (e.g., cortisol released from the adrenal cortex, adrenergic hormones, epinephrine and norepinephrine, from the adrenal medulla; and others) (Axelrod and Reisine, 1984; McEwen, 2009; Sapolsky et al., 1986).”

“…through the representation of experience in a cognitive workspace involving neocortical areas, especially prefrontal and parietal cortical areas (Crick and Koch, 1990, 2004, Dehaene and Changeux, 2004, Baars, 2005; Frith and Dolan, 1996; Frith et al., 1999; Frith, 2008; Shallice, 1988; Shallice…”

“…and motivated states have profound effects on the brain, recruiting widespread areas into the service of the immediate situation, monopolizing and/or synchronizing brain resources, has been proposed previously (Gallistel, 1980; Maturana and Varela, 1987; Scherer, 2000; LeDoux, 2002, 2008).”

“…(Graybiel, 1976; Mogenson et al., 1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et…”

“Indeed, some aspects of defense stimulus processing in primates, including humans, involves preferential rapid learning to certain classes of innately ‘‘prepared’’ stimuli (Seligman, 1971; Öhman, 1986; Mineka and Öhman, 2002).”

“…medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998; Paré et al., 2004; Maren and Quirk, 2004; Quirk and Mueller, 2008; Haubensak et al., 2010).”

“And just as an appetitive CS enhances bar pressing for food, and aversive CS suppresses food-maintained bar pressing (Estes and Skinner, 1941; Hammond, 1970; Cardinal et al., 2002; Balleine and Killcross, 2006).”

“…energy and nutrition maintenance, fluid balance, thermoregulation), rather than on the actual responses controlled by the circuit, a species-independent set of criteria emerge for defining brain systems that detect significant events and control responses that help meet the challenges and…”

“For example, dopamine contributes to the invigoration or activation of behavior during the exploratory search phase of a motivated state (Berridge 2004; Berridge and Robinson, 1998; Robbins and Everitt, 2007).”

“…for much research on the neural basis of emotional functions in the human brain—a recent review uncovered 551 studies between 1990 and 2008 that used Ekman’s basic emotions faces or variants of these to study functional activity related to emotion in the human brain (see Fusar-Poli et al., 2009).”

“…distinct populations of chemically specific neurons that underlie sleep-wake cycles and the degree of arousal, attention, and vigilance while awake (Jouvet, 1969, 1999; Steriade, 1995, 2004; Jacobs et al., 1990; Jones, 2003; Aston-Jones, 2005; Monti and Jantos, 2008; Sarter et al., 2005; Arnsten…”

“…and recognized in people around the world, conserved in our close animal ancestors, and supposedly hard-wired into brain circuits by evolution (Darwin, 1872; Tomkins, 1962; Ekman, 1972, 1980, 1984, 1992, 1999a, 1999b; Izard, 1992, 2007; Damasio, 1994, 1999; Panksepp, 1998, 2000, 2005; Prinz, 2004).”

“…wired into the brain by evolution, and that these mediate functions that contribute to survival and well-being of the organism (e.g., Cannon, 1929; MacLean, 1949, 1952; Hess, 1954; Stellar, 1954; von Holst and von SaintPaul, 1962; Flynn, 1967; Olds, 1977; Siegel and Edinger, 1981; Panksepp, 1982,…”

“Once the association is formed, the CS itself has the ability to elicit innate defense responses.”

“…1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et al., 2006; Nestler, 2004; Kelley, 2004).”

“Yet, in spite of these differences, there is a highly conserved organizational plan that is characteristic of all vertebrate brains (Nauta and Karten, 1970; Northcutt and Kaas, 1995; Swanson, 2002; Butler and Hodos, 2005; Striedter, 2005).”

“…et al., 2007), reproduction (e.g., Pfaff, 1999; Oomura et al., 1988; Blaustein, 2008), thermoregulation (Nakamura and Morrison, 2007), fluid balance (Johnson, 2007; Fitzsimons, 1979), and energy/nutritional regulation (Elmquist et al., 2005; Morton et al., 2006; Saper et al., 2002) is strong.”

“…(e.g., Darwin, 1872; James, 1884; Cannon, 1927, 1931; Duffy, 1934, 1941; Tomkins, 1962; Mandler, 1975; Schachter, 1975; Ekman, 1980, 1984, 1992; Izard, 2007; Frijda, 1986; Russell, 2003;; Ekman and Davidson, 1994; LeDoux, 1996; Panksepp, 1998, 2000, 2005; Rolls, 1999, 2005; Damasio, 1994,…”

“…1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et al., 2006; Nestler, 2004; Kelley, 2004).”

“…and recognized in people around the world, conserved in our close animal ancestors, and supposedly hard-wired into brain circuits by evolution (Darwin, 1872; Tomkins, 1962; Ekman, 1972, 1980, 1984, 1992, 1999a, 1999b; Izard, 1992, 2007; Damasio, 1994, 1999; Panksepp, 1998, 2000, 2005; Prinz, 2004).”

“While these transmitters are released in widespread areas of the brain, their effects are especially profound on neurons that are actively engaged in information processing (Aston-Jones et al., 1991; Foote et al., 1983, 1991; Aston-Jones and Bloom, 1981).”

“…PAGv, venral periaqueductal gray region; PMH, premammilary nucleus of the hypothalamus. the medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998;…”

“Other activities, such as eating, drinking, and sex, are actively suppressed (Gray, 1987; Lima and Dill, 1990; Blanchard et al., 1990; Fanselow, 1994; Choi et al., 2005).”

“…Cannon, 1929; MacLean, 1949, 1952; Hess, 1954; Stellar, 1954; von Holst and von SaintPaul, 1962; Flynn, 1967; Olds, 1977; Siegel and Edinger, 1981; Panksepp, 1982, 1998, 2005; Blanchard and Blanchard, 1972; Bolles and Fanselow, 1980; Damasio, 1994, 1999; Berridge, 1999; McNaughton, 1989; Swanson,…”

“…1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et al., 2006; Nestler, 2004; Kelley, 2004).”

“The exact target remains to be determined but CEA connects with LH both directly and indirectly (Petrovich et al., 1996; Pitkänen et al., 1997).”

“…and recognized in people around the world, conserved in our close animal ancestors, and supposedly hard-wired into brain circuits by evolution (Darwin, 1872; Tomkins, 1962; Ekman, 1972, 1980, 1984, 1992, 1999a, 1999b; Izard, 1992, 2007; Damasio, 1994, 1999; Panksepp, 1998, 2000, 2005; Prinz, 2004).”

“It is therefore of great interest that recent work on the role of dopamine as a reward prediction error signal is beginning to recognize the importance of specific motivational states in modulating the effects of dopamine as a reward prediction error signal (Schultz, 2006; Glimcher, 2011).”

“…sleep-wake cycles and the degree of arousal, attention, and vigilance while awake (Jouvet, 1969, 1999; Steriade, 1995, 2004; Jacobs et al., 1990; Jones, 2003; Aston-Jones, 2005; Monti and Jantos, 2008; Sarter et al., 2005; Arnsten and Li, 2005; Robbins, 2005; Nieuwenhuys, 1985; Nishino, 2011).”

“…1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et al., 2006; Nestler, 2004; Kelley, 2004).”

“Indeed, different cultures and their languages express emotions differently (Kitayama and Markus, 1994; Wierzbicka, 1994; Averill, 1980).”

“…Frijda, 1986; Russell, 2003;; Ekman and Davidson, 1994; LeDoux, 1996; Panksepp, 1998, 2000, 2005; Rolls, 1999, 2005; Damasio, 1994, 1999; Leventhal and Scherer, 1987; Scherer, 2000; Ortony and Turner, 1990; Öhman, 1986, 2009; Johnson-Laird and Oatley, 1989; Ellsworth, 1994; Zajonc, 1980;…”

“…amygdala; BA, basal amygdala; CEA, central amygdala; LA, lateral amygdala; LH, lateral hypothalamus; MEA, medial amygdala; NAcc, nucleus accumbens; VMH, ventromedial hypothalamus; PAGd, dorsal periaqueductal gray region; PAGv, venral periaqueductal gray region; PMH, premammilary nucleus of the…”

“A recent study by Lin et al. (2011) showing distinct populations of cells in the ventromedial hypothalamus that contribute to mating and intruder attack is a prime example.”

“…representation of experience in a cognitive workspace involving neocortical areas, especially prefrontal and parietal cortical areas (Crick and Koch, 1990, 2004, Dehaene and Changeux, 2004, Baars, 2005; Frith and Dolan, 1996; Frith et al., 1999; Frith, 2008; Shallice, 1988; Shallice et al., 2008).”

“That certain emotions are wired into the brain is also a major tenet of evolutionary psychology (e.g., Tooby and Cos- 654 Neuron 73, February 23, 2012 ª2012 Elsevier Inc. mides, 1990; Pinker, 1997; Nesse, 1990).”

“That is, they modulate rather than initiate neural activity, regulating neuronal excitability and neurotransmission (Schildkraut and Kety, 1967; Hasselmo, 1995; Lopez and Brown, 1992).”

“…Flynn, 1967; Olds, 1977; Siegel and Edinger, 1981; Panksepp, 1982, 1998, 2005; Blanchard and Blanchard, 1972; Bolles and Fanselow, 1980; Damasio, 1994, 1999; Berridge, 1999; McNaughton, 1989; Swanson, 2000; Ferris et al., 2008; Choi et al., 2005; Motta et al., 2009; Lin et al., 2011; Öhman, 2009).”

“…medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998; Paré et al., 2004; Maren and Quirk, 2004; Quirk and Mueller, 2008; Haubensak et al., 2010).”

“…medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998; Paré et al., 2004; Maren and Quirk, 2004; Quirk and Mueller, 2008; Haubensak et al., 2010).”

“…and often conscious, determines stimulus meaning and predisposes actions (e.g., Arnold, 1960; Bowlby, 1969; Frijda, 1986; Lazarus, 1991a, 1991b; Leventhal and Scherer, 1987; Lazarus and Folkman, 1984; Smith and Ellsworth, 1985; Scherer, 1988; Scherer et al., 2001; Sander et al., 2005;…”

“…and motivated states have profound effects on the brain, recruiting widespread areas into the service of the immediate situation, monopolizing and/or synchronizing brain resources, has been proposed previously (Gallistel, 1980; Maturana and Varela, 1987; Scherer, 2000; LeDoux, 2002, 2008).”

“…wired into the brain by evolution, and that these mediate functions that contribute to survival and well-being of the organism (e.g., Cannon, 1929; MacLean, 1949, 1952; Hess, 1954; Stellar, 1954; von Holst and von SaintPaul, 1962; Flynn, 1967; Olds, 1977; Siegel and Edinger, 1981; Panksepp, 1982,…”

“Themodern paradigmatic example of a generic reinforcement mechanism is the role of dopamine in the striatum as a reward prediction error signal (Schultz, 1997).”

“By focusing on the evolved function of a circuit (defense, reproduction, energy and nutrition maintenance, fluid balance, thermoregulation), rather than on the actual responses controlled by the circuit, a species-independent set of criteria emerge for defining brain systems that detect significant…”

“…1952; Hess, 1954; Stellar, 1954; von Holst and von SaintPaul, 1962; Flynn, 1967; Olds, 1977; Siegel and Edinger, 1981; Panksepp, 1982, 1998, 2005; Blanchard and Blanchard, 1972; Bolles and Fanselow, 1980; Damasio, 1994, 1999; Berridge, 1999; McNaughton, 1989; Swanson, 2000; Ferris et al., 2008;…”

“…1996; Panksepp, 1998, 2000, 2005; Rolls, 1999, 2005; Damasio, 1994, 1999; Leventhal and Scherer, 1987; Scherer, 2000; Ortony and Turner, 1990; Öhman, 1986, 2009; Johnson-Laird and Oatley, 1989; Ellsworth, 1994; Zajonc, 1980; Lazarus, 1981, 1991a, 1991b; Barrett, 2006a, 2006b; Barrett et…”

“…focal point of this general motivational system (Graybiel, 1976; Mogenson et al., 1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007,…”

“…conscious, determines stimulus meaning and predisposes actions (e.g., Arnold, 1960; Bowlby, 1969; Frijda, 1986; Lazarus, 1991a, 1991b; Leventhal and Scherer, 1987; Lazarus and Folkman, 1984; Smith and Ellsworth, 1985; Scherer, 1988; Scherer et al., 2001; Sander et al., 2005; Jarymowicz, 2009).”

“Body feedback has played an important role in emotion theory for more than a century (James, 1884; Lange, 1885/1922; Schachter and Singer, 1962; Tomkins, 1962; Adelmann and Zajonc, 1989; Buck, 1980; Damasio, 1994, 1999).”

“…premammilary nucleus of the hypothalamus. the medial CEA (CEm) to the ventrolateral PAG (PAGvl) (Johansen et al., 2011; LeDoux, 2000; Maren, 2001; Fanselow and Poulos, 2005; Davis et al., 1997; Rosenkranz and Grace, 2002; Cousens and Otto, 1998; Paré et al., 2004; Maren and Quirk, 2004; Quirk…”

“…system consisting of multiple subsystems that work in concert to sustain life both on a moment to moment to basis and over long time scales (Bernard, 1878–1879; Cannon, 1929; Lashley, 1938; Morgan, 1943; Stellar, 1954; Selye, 1955; McEwen, 2009; Damasio, 1994, 1999; Pfaff, 1999; Schulkin, 2003).”

“…(defense, reproduction, energy and nutrition maintenance, fluid balance, thermoregulation), rather than on the actual responses controlled by the circuit, a species-independent set of criteria emerge for defining brain systems that detect significant events and control responses that help meet…”

“…sleep-wake cycles and the degree of arousal, attention, and vigilance while awake (Jouvet, 1969, 1999; Steriade, 1995, 2004; Jacobs et al., 1990; Jones, 2003; Aston-Jones, 2005; Monti and Jantos, 2008; Sarter et al., 2005; Arnsten and Li, 2005; Robbins, 2005; Nieuwenhuys, 1985; Nishino, 2011).”

“Dorsolateral prefrontal cortex, a key component of the cognitive workspace, is lacking in most other mammals, and is less developed in nonhuman primates than in humans (Reep, 1984; Braak, 1980; Preuss, 1995; Wise, 2008).”

“For example, there are more than 30 words in English for gradations of fear (fear, panic, anxiety, worry, trepidation, consternation, etc.) (Marks, 1987).”

“…to unconditioned threats, especially odors that signal predators or potentially dangerous conspecifics (Dielenberg et al., 2001; Canteras, 2002; Petrovich et al., 2001; Markham et al., 2004; Blanchard et al., 2003; Motta et al., 2009; Choi et al., 2005; Vyas et al., 2007; Pagani and Rosen,…”

“And just as an appetitive CS enhances bar pressing for food, and aversive CS suppresses food-maintained bar pressing (Estes and Skinner, 1941; Hammond, 1970; Cardinal et al, 2002; Balleine and Killcross, 2006).”

“…stimulus processing is nevertheless subject to epigenetic modulation by various factors inside and outside the organism during development, and throughout life (Bendesky and Bargmann, 2011; Monsey et al., 2011; McEwen et al, 2012; Brown and Hariri, 2006; Casey et al., 2011; Zhang et al., 2004).”

“…for food by a hungry obtain food is facilitated by a conditioned incentive that signals food, is facilitated less by one that signals water and is inhibited by one that signals shock (Corbit and Balleine, 2005; Hammond, 1970), indicating that motivation is tied to specific survival functions.”

“The accumbens, once thought to be mainly involved in processing appetitive stimuli, is now know to contribute to the processing of aversive incentives as well (Salamone, 1994; Schoenbaum and Setlow, 2003; Roitman et al., 2005; Reynolds and Berridge, 2008).”

“Thus, a rat can recognize a predator (cat, fox) by specific chemical constituents of predator odors (Wallace and Rosen, 2000; Vyas et al., 2007; Dielenberg et al., 2001; Markham et al., 2004; Blanchard et al., 2003) and does not have to recognize the predator as a complex perceptual pattern.”

“The detection of a threat by defense circuits of the amygdala leads to the activation of central neuromodulatory and peripheral hormonal systems (see Gray, 1993; LeDoux, 1992, 1995; Davis, 1992; Rodrigues et al., 2009).”

“…et al., 1980; Balleine and Killcross, 1994; Killcross and Robbins, 1993; Everitt et al., 1999; Cardinal et al., 2002; Ikemoto and Panksepp, 1999; Parkinson et al., 1999; Koob, 2009; Sesack and Grace, 2010; Berridge, 2007, 2009; Berridge and Robinson, 1998; Hyman et al., 2006; Nestler, 2004;…”

“That is, they modulate rather than initiate neural activity, regulating neuronal excitability and neurotransmission (Schildkraut and Kety, 1967; Hasselmo, 1995; Lopez and Brown, 1992).”

“Fearful and aggressive faces of conspecifics are also a potent innate defense trigger in humans and other primates (Adolphs, 2008; Davis et al., 2011).”

“…sleep-wake cycles and the degree of arousal, attention, and vigilance while awake (Jouvet, 1969, 1999; Steriade, 1995, 2004; Jacobs et al., 1990; Jones, 2003; Aston-Jones, 2005; Monti and Jantos, 2008; Sarter et al., 2005; Arnsten and Li, 2005; Robbins, 2005; Nieuwenhuys, 1985; Nishino, 2011).”

“Similarly, the fact that both appetitive and aversive stimuli activate the amgydala in fMRI studies (e.g., Canli et al., 2002; Hamann et al., 2002; Lane et al., 1999) does not mean that these stimuli are processed the same by the amygdala.”

“Thus, a rat can recognize a predator (cat, fox) by specific chemical constituents of predator odors (Wallace and Rosen, 2000; Vyas et al., 2007; Dielenberg et al., 2001; Markham et al., 2004; Blanchard et al., 2003) and does not have to recognize the predator as a complex perceptual pattern.”

“Similarly, the fact that both appetitive and aversive stimuli activate the amgydala in fMRI studies (e.g., Canli et al., 2002; Hamann et al., 2002; Lane et al., 1999) does not mean that these stimuli are processed the same by the amygdala.”

“…processing, for which analogies in behaviorally relevant peripheral odor-encoding and central representation occur using similar organizational principles in anatomically distinct (nonhomologous) structures in Drosophila and rodents (see Bargmann, 2006; Sosulski et al., 2011; Wang et al., 2011).”

“Dorsolateral prefrontal cortex, a key component of the cognitive workspace, is lacking in most other mammals, and is less developed in nonhuman primates than in humans (Reep, 1984; Braak, 1980; Preuss, 1995; Wise, 2008).”

“…and lesion results from humans (e.g., Phelps, 2006; Damasio, 1994, 1999; LaBar and Cabeza, 2006; Whalen and Phelps, 2009; Büchel and Dolan, 2000; Mobbs et al., 2009; Schiller and Delgado, 2010) show that the amygdala plays a key role in defense conditioning, and thus suggest that, at least to…”

“For example, Glickman and Schiff (1967) proposed that reinforcement is a facilitation of activity in neural systems that mediate species-specific consummatory acts.”

“In the field of emotion, these are described as unconditioned and conditioned fear stimuli.”

“…across mammals of mechanisms underlying survival functions such as defense (e.g., LeDoux, 1996, 2012; Phelps and LeDoux, 2005; Motta et al., 2009; Choi et al., 2005; Kalin et al., 2004; Amaral, 2003; Antoniadis et al., 2007), reproduction (e.g., Pfaff, 1999; Oomura et al., 1988; Blaustein,…”

“Within the accumbens information processing segregated along motivational lines—aversive and appetitive stimuli are processed separately at the cellular and molecular level (Roitman et al., 2005, 2008).”

“…the years (e.g., Duffy, 1941; Lindsley, 1951; Schachter and Singer, 1962; Schachter, 1975; Schildkraut and Kety, 1967; Mandler, 1975; Lang, 1994; Robbins, 1997) and is also important in contemporary dimensional theories of emotion (Russell, 1980, 2003; Russell and Barrett, 1999) and some neural…”

“…words that refer to feelings as a means of classifying and studying human emotions has been questioned by a number of authors over the years (e.g., Duffy, 1934, 1941; Mandler, 1975; Russell, 1991, 2003; Barrett, 2006a, 2006b; Kagan, 2007; Griffiths, 1997; Rorty, 1980; Dixon, 2001; Zachar, 2006).”

“Research on the neural basis of feelings in humans is in its infancy (Panksepp, 1998; 2005; Damasio, 2003; Damasio et al., 2000; Ochsner et al., 2002; Barrett et al., 2007; Rudrauf et al., 2009; Critchley et al., 2004; Pollatos et al., 2007).”

“For many, the core of emotion is the subjective experience (Barrett, Mesquita, Ochsner, & Gross, 2007; LeDoux, 2012).”

“…Darwin's logic, modern affective neuroscience also posits brain mechanisms of emotional reactions to mediate evolved ''survival functions'' (LeDoux, 2012), with emotional ''core features that can form the basis for studies of emotion across phylogeny'' (p. 198) (Anderson and Adolphs,…”

“2017), has wreaked havoc with the scientific accumulation of knowledge about emotion (also see Barrett, 2012; LeDoux, 2012).”

“However, different subdivisions of the amygdala and hypothalamus are thought to control learned vs innate responses, in a parallel manner (Figure 1A; reviewed in [Davis, 1992; Fanselow, 1994; LeDoux, 1995, 2000; Gallagher and Chiba, 1996; Fanselow and LeDoux, 1999; Canteras, 2002; Rosen, 2004; Swanson, 2005; Gross and…”

“D ow nloaded from https://academ ic.oup.com /scan/article-abstract/12/1/1/2823712 by guest on 04 M ay 2020 2017), has wreaked havoc with the scientific accumulation of knowledge about emotion (also see Barrett, 2012; LeDoux, 2012).”

“In turn, CeA has direct connections to periaqueductal gray (PAG), a fundamental area that organizes the threat response (LeDoux, 2012).”

“…fear extinction see Milad and Quirk (2012), for fear acquisition and extinction across development see Shechner et al., 2015; for disparities between affective and cognitive neuroscience in general see Panksepp et al., 2017; for fear vs. threat responses see LeDoux, 2012; LeDoux and Pine, 2016).”

“…given the central role of the amygdala in aversive learning, threat detection, and the mobilization of defensive responses to external stimuli (LeDoux 2003, 2012), although it is well established that the amygdala responds not only to threatening cues but also to a wide variety of stimuli…”

“…life events (Bolger, 1990); how significant others aid in coping with life-threatening events, such as cancer onset (Bolger, Foster, Vinokur, & Ng, 1996); or how contextual threat cues are encoded into memory, leading to behavior change, in animal models (LeDoux, 2012; Phillips & LeDoux, 1992).”

“…achieve a consensus definition that is accepted across fields from neuroscience to psychology to philosophy have repeatedly failed, to the extent that at least one prominent emotion researcher has suggested that we excise the word “emotion” altogether from our scientific vocabulary (LeDoux, 2012).”

“Nonetheless, the belief that emotions can be localized somewhere in the brain is very strong (see [11 ,12 ] for discussions), and efforts at topographical localization have given way to the hypothesis that emotions can be localized to specific brain networks (e.”

“71, 72 Intuition can also identify patients with life-threatening conditions that would be underappreciated by traditional clinical parameters alone.”

“This third theme goes hand in hand with current developments in the neurobiology of consciousness, and it may bring back to the scientific study of emotions a topic that, ever since Behaviorism, has been excluded (despite the fact that many modern neurobiological views on emotion now mention it [12,13,15,24,25]).”

“In humans, the highly developed ability to envisage, simulate and predict future scenarios allows us to modify our current behavior to prepare, escape, or even avoid possible future dangers (Suddendorf andCorballis, 2007).”

“The distributionist view argues that no single anatomical structure uniquely specializes for individual emotion categories [40, 41].”

“As a prototypical network subject to 5-HT-induced modulation, the circuitry of experience-dependent associative and emotional learning has been implicated in social cognition and emotion, including the associated phenomena of contextual fear responses (Figure 6; LeDoux, 2012).”

“efensive reactions to threat stimuli are a basic survival mechanism in all vertebrates 1 .”

“Decades of animal and human neuroscience research supports the presence of neural circuits, conserved across species, that respond to environmental threats and orchestrate defensive responses (LeDoux, 2003, 2012; Phelps & LeDoux, 2005).”

“Specific neural circuits, identified in animal models (LeDoux, 2003, 2012) and conserved across species (Milad & Quirk, 2012; Phelps & LeDoux, 2005), respond to potential threats, orchestrate defensive responses, and support aversive learning – encompassing amygdala, hippocampus, and dorsal and…”

“Additionally, the amygdala is involved in the learning and consolidation of emotional memories and has an essential role in conditioned responses (Davis, 1992; LeDoux, 2012).”

“This fear circuit is involved in evaluation and detection of threat cues, eliciting and regulating emotional responses, and fear learning and extinction (LeDoux, 2012).”

“The importance of distinguishing between instinctive behaviours generated by the brainstem and felt emotions generated by the cortex has been previously discussed extensively (LeDoux, 2012; LeDoux, 2013; LeDoux, 2014a; LeDoux, 2014b; Rolls, 2014a; Rolls, 2014b; Scott, 2014).”

“Most fundamental are the desires associated with basic behaviors required for survival and reproduction, such as hunger, thirst, and sex (Freud, 2001; Hull, 1943; LeDoux, 2012; Maslow, 1943).”

“LeDoux has described a theory of the neural basis of emotion (LeDoux 1992, 1995, 1996, 2012) that is probably conceptually similar to that of Rolls (Rolls 1999, 2000a, 2005, 2014a, 2018) except that he focuses mostly on the role of the amygdala in emotion (and not on other brain regions such as the…”

“The tools of modern neuroscience have provided the basis for a broader understanding and precise dissection of brain networks that process survival-related stimuli [7] and have clarified the impact of biologically relevant stimuli on brain networks related to motivation, pleasure, and learning [8].”

“…evidence for hypereactivity in core nodes of the salience network (e.g. amygdala and insula), which presumably reflects increased sensitivity to conditioned fear-cues and elevated interoceptive monitoring of arousal and aversive states (Etkin et al., 2011; LeDoux, 2012; Pessoa and Adolphs, 2010).”

“In other articles, ingredients also include basic behavioral adaptations like freezing, fleeing, and fighting, although there is no necessary one-to-one correspondence between a behavior and an emotion category (e.g., Barrett, 2012; LeDoux, 2012).”

“…rats is just one of many lines of evidence, including research on humans, demonstrating a relationship between emotions and feelings comparable to that between nociception and pain (see Rose 2002; Berridge and Winkielman 2003; Damasio 2005; Rose 2007; LeDoux 2012 for a more detailed explanation).”

“The central nucleus plays a role in the modulation of autonomic and endocrine responses for a host of visceral functions (LeDoux, 2000, 2012).”

“Evidence that there are distinct circuits for behavioral adaptations does not falsify the CAT because we do not define emotions as actions (Barrett, 2012; for a similar view, see LeDoux, 2012).”

“In contrast, the amygdala may be crucial for learning associations between aversive events (signaled in part by PAG) and other sensory cues (LeDoux, 2012).”

“Internal states of arousal, motivation, and emotion exert a major influence on how the brain processes sensory information to control behavior (Berridge, 2004; Bargmann, 2012; LeDoux, 2012; Anderson and Adolphs, 2014; Anderson, 2016).”

“The term emotion has been notoriously difficult to define (Hamann, 2012; LeDoux, 2012) and it has even been asked how meaningful a categorical distinction between cognition and emotion (or “cognitive” and “affective” brain areas) is (Pessoa, 2008).”

“SOCIAL APPROACH Several models of emotion and social cognition (e.g., Phillips et al., 2003a,b; Lieberman, 2007; LeDoux, 2012) include core processes subserving rapid or automatic (sometimes even unconscious) processing of information in terms of safety versus danger, which are intrinsically linked with behavioral ten…”

“Thus, by gaining control over US delivery, subjects successfully inhibit the expression of deeply instated behavioral reactions that normally prevail in situations of expected threat (Fanselow and Lester, 1988; Blanchard et al., 2005; LeDoux, 2012).”

“Such states, for example, can be referred in terms of neural activity in a defensive survival circuit (rather than in a fear circuit) [4].”

“Prevailing models have localized fear learning largely to nuclei in the BLA (Phelps et al, 2004) although recent studies have asserted a possible role for the CMA (LeDoux, 2012).”

“Thus, in spite of groundbreaking advances in understanding the circuitry underlying basic behavioral adaptations for safety and reproduction (including ‘ threat ’ behaviors; [11]), there is no comprehensive model of the neurophysiological basis of emotional experience in humans.”

“Three general factors determining defensive behavioral patterns in response to threats have been proposed4: (1) environmental context (e.”

“The importance of distinguishing between instinctive behaviours generated by the brainstem and felt emotions generated by the cortex has been previously discussed extensively (LeDoux, 2012; LeDoux, 2013; LeDoux, 2014a; LeDoux, 2014b; Rolls, 2014a; Rolls, 2014b; Scott, 2014).”

“and there are important homologues of these mechanisms in invertebrates too [4].”

“Food-deprivation creates a type of “global organismal state change” (Kim et al., 2013; LeDoux, 2012) that is multi-dimensional: it involves multiple physiological and behavioral changes.”

“Defensive behaviors are immediate responses (LeDoux, 2012) triggered by activity in survival circuits that detects threats (LeDoux, 2014) and leading to the conscious emotion of fear that follows immediately.”

“mal neuroscience (LeDoux, 2012), but I think there is a simpler solution: do the same thing we as scientists do when we study”

“This assumption would likewise hold for anxiety disorders where the vmPFC has been shown to be essential for the top-down control (suppression) of a threat response (Cain and Ledoux, 2008; Sullivan, 2003; Ledoux, 2012), which inference is also supported by Rauschecker et al.”

“Carvalho, 2013; Panksepp, 2010), levels of arousal, actions (Bernroider and Panksepp, 2011; Gardiner, 2015), hedonics (pleasure and pain (Buck, 1985; Damasio and Carvalho, 2013; LeDoux, 2012; Panksepp, 2010), drives (Alcaro and Panksepp, 2011), cognitions (including per-”

“Emotions can be considered as being composed of two components: one being tightly related to the production of behavioural responses necessary for survival or maintenance of well-being [27], the other being the subjective feeling associated with an emotional stimulus or an emotional situation [15].”

“Other models of emotion that are not explicitly identified as psychological constructionist views but that share some of this tradition’s assumptions have also emerged (e.g., Boiger & Mesquita, in press; Coan, 2010; LeDoux, 2012).”

“As such, these areas that contribute to the emotional brain processes, ‘‘reflect functions and circuits related to survival’’ that integrate ‘‘motivation, reinforcement, and arousal’’ (LeDoux, 2012).”

“As alluded to above, one influential hypothesis has suggested that unconscious processing of emotional stimuli may arise through a subcortical pathway to the AMY, which involves the superior colliculus and the pulvinar (LeDoux, 2012).”

“Extensive work over the last two decades has focused on determining the brain substrates underlying fear and its behavioral sequelae (LeDoux, 2012; Seymour & Dolan, 2008), and similar efforts have been directed towards examining the central substrates of hunger, food intake and energy balance…”

“The restructuring of sexual behaviour —and other survival behaviours 124 —into these three phases is critical in order to achieve a meaningful species comparison; although introspection and subjective feel­ ings are major topics in human research, it is impossible to know how animals perceive the…”

“This system consists of a set of nuclei located in the hypothalamic medial zone: the anterior hypothalamic nucleus (AH), the dorsomedial portion of the ventromedial hypothalamus (VMHdm), and the dorsal premammillary nucleus (PMD).”

“These structures are critical in the detection of, as well as in the physiological and behavioral response to, aversive events and potential threats (Sehlmeyer et al., 2009; Schlund et al., 2010; LeDoux, 2012).”