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).”

“Further, arousal may enhance processing in the survival circuit—a hardwired, evolutionarily conserved set of defensive mechanisms activated in challenging and possibly life-threatening situations—which facilitates activity in sensory, memory, and attentional processes (LeDoux, 2012, 2022).”

“Over time, practicing in emotion regulation, either directly (ABM, reappraisal) or indirectly (by enhancing cognitive control), could re-configure our survival circuits [1], such that events or stimuli that were once considered as a threat to survival (or alternatively highly hedonic) loses their ability to trigger su…”

“Emotional systems are found in both animals and humans (LeDoux, 2012), which presents the question of whether or not emotions rely on consciousness.”

“2007; Murray 2007), suggesting that the amygdala processes rewards and threats in two different ways (LeDoux 2012).”

“There is a general consensus that the amygdala plays an important role in the acquisition, storage, and expression of cued fear-related behavioral responses, while the hippocampus plays an important role in encoding the contextual environment and the prefrontal cortex plays an important role in modulating the expressi…”

“Furthermore, differential adaptive links with different neuronal sources of input to the GW (cognitive workspace, LeDoux 2000, 2012), such as survival circuit activity (and the amygdala in particular), environmental stimuli, language, and body feedback, can be associated to different facets of…”

“Several brain areas involved in cerebral processing of these emotions are described (Panksepp, 2011; LeDoux, 2012), with sad states being regulated most prominently by the anterior cingulate cortex and the dorsomedial prefrontal cortex (Murphy et al., 2003).”

“…and cognitive processes are tightly intertwined, in terms of both their psychological function and neural substrates, and the effort moved to understand the mechanisms of this connection (Clore and Colcombe, 2003; Pessoa and Adolphs, 2011; LeDoux, 2012; Winkielman et al., 2015a; Barrett, 2017).”

“Importantly, if mnemonic information is conjured up when reactivating a brain area, each of these findings makes specific hypotheses with regards to the phenotypic consequences resulting from acute and chronic modulation of each area (Feldman Barrett and Wager, 2006; LeDoux, 2012).”

“Next, we wished to distinguish fear and anxiety from other negative emotions, consistent with theory and evidence that threat or fear-inducing stimuli are processed in different ways than other negative stimuli (e.g., LeDoux, 2012; March, Gaertner, & Olson, 2017; Öhman & Mineka, 2001).”

“Influential theories of emotion describe pleasure as an integral part of core affect (Lindquist et al., 2012) or survival functions (LeDoux, 2012), and neuroimaging evidence links pleasurable music listening with brain areas implicated in emotion and the dopaminergic reward system (Blood et al.,…”

“Note that various factors might shape the available repertoire, for example, based on innate programs, past experience, context, but also hormonal factors (LeDoux, 2012).”

“…in affective science relates to theoretical approaches that take a broad functional perspective (e.g. Adolphs, 2017b; Adolphs and Anderson, 2018; LeDoux, 2012) and those that take a psychological constructionist perspective (e.g. Barrett, 2017a, 2017b); it may not be possible to bring these…”

“The PAG is commonly thought to be involved in regulating systems of the body’s internal milieu, particularly when dealing with moments of motivated behavior when energy expenditures are higher, such as during episodes of pain, sexual behavior, distress, and feeling threatened (Fanselow, 1991; LeDoux, 2012; Keay and Ba…”

“Facial and bodily expressions therefore both activate a brain region involved in the avoidance motivation system for action (Cacioppo & Berntson, 1994; Lang, Bradley, & Cuthbert, 1997; Ledoux, 2012).”

“It is important to emphasize that emotional appraisal does not necessarily entail the conscious elaboration of stimuli [37,39,40 ], which, incidentally, would be difficult to envisage in rats.”

“…additional explanatory power is o ff ered by connecting neural structure and function to an ecologically situated, embodied and embedded cognitive architecture selected for by evolutionary pressures (Bracha et al., 2004; Clinchy et al., 2011; Dodgson & Gordon, 2009; LeDoux, 2012; Roelofs, 2017).”

“For these and other reasons, I have argued that the amygdala’s role in detecting and responding to threats is more appropriately considered in terms of a non-conscious defensive survival circuit than a conscious fear circuit [4,5].”

“…recently shifted his focus away from “ emotion ” circuits and towards “ survival circuits ” , information about which feeds into the cognitive workspace along with information from explicit memory, language, environmental activity, body feedback, and central nervous system arousal (LeDoux, 2012).”

“(2018) found that direct fear learning activates a neural network that has been linked to the acquisition, storage, and expression of conditioned fear (LeDoux, 2012), as well as the aversive value of pain (Kober et al.”

“LeDoux (2012), on the other hand, placed further distance between the study of animal affective states and that of human emotional feelings, advocating the use of emotion neutral terminology such as ‘defensive survival circuits’ rather than ‘fear’, because human emotion words are too easily…”

“regulatory feedback projection forms part of the brain circuitry involved in emotion regulation [95, 97].”

“, 2017), and that fear is highly conserved throughout the animal kingdom (LeDoux, 2012; Adolphs, 2013), it is plausible that fear conditioning-based studies in rodents can reveal causative pathological mechanisms that govern the development of PTSD.”

“Even though both fear and aggression have been extensively studied [3], their underlying neural mechanisms remain largely unknown.”

“…the central nucleus of the amygdala to the lateral hypothalamus are involved in activating autonomic sympathetic nervous system responses, and projections to the ventrolateral periaqueductal gray direct the expression of behavior responses, such as defensive freezing (Davis, 1992; LeDoux, 2012).”

“The affective internal motivational experiences of animals, long debated and still often denied by behaviorists and behavioral neuroscientists (LeDoux, 2012) and ethologists (Dawkins, 2012), can finally be addressed empirically through affective neuroscience strategies (Panksepp, 1998, 2011).”

“Still, prominent investigators (usually those who do no brain research) claim we can never know whether nonspeaking animals experience anything (e.g., Dawkins, 2012); a viewpoint shared by some prominent behavioral neuroscientists (LeDoux, 2012, 2014).”

“Several studies suggest that emotional cues activate cortico-limbic appetitive and defensive systems, which, in turn, enhance attention allocation, and prepare the organism for action (Anderson, Laurent, & Yantis, 2011; Gottlieb, 2012; Hickey, Chelazzi, & Theeuwes, 2010; Lang, Bradley, & Cuthbert, 1997; Lang & Bradley…”

“Some emotion researchers now agree (not always for the same reasons) that the human focus has enabled continued reliance on long-standing theoretical assumptions predominantly derived from first-person intuitions about human experience, to the detriment of the field [1,2,4,5].”

“Extensive research with animals (e.g., Davis, 2000; Fanselow & Poulos, 2005; Kapp, Supple, & Whalen, 1994; LeDoux, 2003, 2012) has defined a set of neural structures that makeup the brain’s survival network (for an overview, and its relationship to research with human participants, see Davis &…”

“Vertebrates, including humans and rodents, share a highly conserved threat response circuitry that impacts learning and memory to facilitate the acquisition of salient cues that inform defensive behavioral responses, the functioning of which is hypothesized to be a key dimension underlying PTSD (LeDoux, 2012; Milad et…”

“Examples are the updated Jo ur na l P re -p ro of version of the LeDoux (2012) theory of fear and Berridge’s model of distinct neural circuits involved in wanting (i.e. motivation) and liking (i.e. experience) (Berridge & Robinson, 2016; Castro & Berridge, 2014).”

“One theory has proposed that emotional processing integrates information from deliberate, conscious pathways with that from automatic, unconscious pathways (LeDoux, 1996, 2012).”

“Before proceeding, it is important to note that FC studies, especially those conducted in rodents, relate directly to implicit Pavlovian memories and defensive responses [33].”

“Likewise, emotional responses in children with hydranencephaly could be triggered unconsciously via subcortical networks, such that these responses do not necessarily reflect emotional experiences (LeDoux (2012); LeDoux and Brown (2017)).”

“Emotion processing in the brain is an intricate activity, asynchronous across multiple regions [17].”

“vol. 182, no. 6 the american naturalist december 2013 Effects of the Emotion System on Adaptive Behavior”

“In biological theories relevance is assessed in terms of survival, but it is also thought to be a necessary condition to evoke an emotion (LeDoux, 2012; Panksepp, 1998).”

“However, it is not clear so far whether the categorical structure and the representation of the emotion labels is the same at earlier processing levels, especially at a non-conscious level [42, 43].”

“Anderson and Phelps 2000; LeDoux 1996, 2012.”

“4 The idea that basic emotions have dedicated neural circuits (LeDoux, 1996) has been amended; LeDoux (2012) only claims that survival circuits have dedicated, innate neural substrates, and these survival circuits mediate fundamental phenomena related to emotion. destinations, consumer goods, and…”

“Hedonic experience requires the contemporary activation of neuronal circuits situated in mesocorticolimbic areas (Damasio, 2010; Panksepp, 2011; LeDoux, 2012) which have undergone an extraordinary evolution in time, precisely because affective reactions guarantee a significant objective gain for…”

“…responses, especially those linked to our biological fitness (e.g., those occurring when in danger or when hungry or thirsty), have been sculpted in ‘survival circuits’ throughout evolution (LeDoux, 1996, 2012) profoundly influenced the interpretation of behavioral reactions to valenced stimuli.”

“First, Joseph LeDoux (2012a, 2012b), as noted earlier, has solidified the position that the scientific book on emotional feelings in other animals is epistemologically closed, thereby mandating a continued use of behavioronly models of emotions in animals, as represented by his ongoing fine…”

“Defence behaviours essential to survival can be innate, but they can also be learnt, and during evolution are conserved across species, including humans (Takahashi, 1992a,b; Blanchard et al. 2001a,b; Gross & Canteras, 2012; LeDoux, 2012).”

“…actions through association and reinforcement; and (3) the generalized arousal in which a feedback loop is established to facilitate the continuing activation of survival circuits, to enhance attention to external stimuli, and to stimulate memory retrieval and formation (LeDoux, 2012).”

“These studies have shown that the amygdala, insula, and anterior cingulate cortex (ACC) are activated during the processing of negative emotions, with hyperactivation in individuals with high levels of anxiety [6, 7].”

“Although evidence for a similar role in humans is only beginning to emerge (e.g. Noriuchi et al., 2008; Bartels & Zeki, 2004; Laurent & Ablow, 2012), it has been strongly argued that the brain circuits subserving survival functions are conserved across mammals (Ledoux, 2012).”

“Thus there is not likely to be a unitary state with a single neural representation, but rather multiple forms depending on the underlying survival circuit activated (LeDoux 2012; Moyer 1976).”

“Besides the BG, the amygdala is another important brain structure that is strongly involved in the processing of emotional stimuli (LeDoux 2012).”

“In sum, there is compelling evidence of complex bi-directional controls over emotion responding and the complexity of these processes may make them particularly challenging to disentangle in humans (LeDoux, 2012).”

“Finally, in the circa-strike stage the threat is most imminent and the organism engages in active behavioural strategies accompanied with increased physiological activation (Kim et al., 2013; LeDoux, 2012).”

“Hence, there have been great experimental challenges in defining the underlying neuronal circuits, and even the general organization of the circuits is still debated, such as whether the subjective feeling underlying an emotion is a cause (upstream) or consequence (downstream) of the external manifestations (James, 18…”

“Infiltrating immune cells causes vascular oxidative stress via NOX2 activity [25,34].”