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Key points
- One key theory links emotions and consciousness to body homeostasis, regulated by primitive brain regions.
- A competing theory views the prefrontal cortex as essential to the conscious feeling of emotion.
- Either way, feelings are key to understanding consciousness—as they are inherently subjective and conscious.
“Understanding the cognitive basis of emotional experience is a subset of the problem of understanding the cognitive foundations of consciousness.”
—Joseph LeDoux1
This is Part 4 of a four-part series on What Actually Are Emotions?
Antonio Damasio’s theories of emotion
Neuroscientist Antonio Damasio’s theories of emotion, and their relation to consciousness, are among the most influential in the field.2
Damasio defines "emotions" as complex interactions between sensory inputs, the brain, and the body, resulting in motor and physiological changes called "action programs." These changes are triggered by shifts in the internal or external environment to maintain or restore homeostasis.3 External stimuli, like encountering a predator, provoke emotions, while internal stimuli, such as hunger, relate to basic survival drives. Feelings are hybrid processes that accompany emotions, integrating the brain's perception of bodily changes into a coherent mental state.4
Central to Damasio's theory is the idea that consciousness depends on interactions between homeostatic routines and multilevel interoceptive5 maps, with affect and feeling at its core.6 He argues that consciousness arises from a continuous flow of feelings related to the body's internal state—sensations like fever, nausea, thirst, and well-being. Unlike exteroception, which maps external stimuli, interoception uniquely and constantly reflects internal events,7 linking the "object of perception" to the "subject" and grounding the sense of self (ownership). This process involves brainstem regions such as the central relay nuclei, which is not fully insulated from the blood-brain barrier and so interacts directly with molecules in the bloodstream.8 These nuclei facilitate a feedback loop between physiological states and subjective experience, contributing to the mind's awareness of its own body.
In his earlier work, Damasio placed significant emphasis on the cerebral cortex's role in generating conscious emotions, proposing that cortical body maps create a sense of self and awareness.9 Over time, however, he expanded his theory to incorporate the contributions of subcortical structures, particularly the brainstem, as essential for basic emotional experiences.10 While he agrees with Panksepp, Merker, and Solms (discussed in Part 2) that “there are aspects of feelings that probably can be accounted entirely by the operations of the brainstem and in no way will require a continuation upstream” (i.e., in the cortex), Damasio does stress that consciousness is more than these basic operations. It involves the "gathering and integration" of signals originating in the brainstem and body, which are then greatly influenced by higher cortical processes, “creating conscious emotions in a much richer interactive way.”11
Joseph LeDoux’s theory of emotion
While Damasio emphasizes homeostasis and the body's internal state as the basis of emotions, neuroscientist Joseph LeDoux focuses on the role of cognitive processes, particularly in the prefrontal cortex, in constructing conscious emotional experiences.
The amygdala is not a “fear circuit”: LeDoux is known for his seminal work on the amygdala's role in fear responses, though he defines the amygdala12 as a defensive survival circuit, not a “fear circuit.” When faced with danger, defensive responses like fleeing or freezing and the conscious feeling of fear occur together not because they are both processed in the amygdala, but because they share the same starting point: a threatening stimulus (such as the sight of a snake) enters the brain through the sensory (e.g., visual) system. From there, neural pathways diverge. The amygdala triggers instinctive, nonconscious defensive responses like freezing,13 while signals to the prefrontal cortex generate the conscious feeling of fear through cognitive interpretation. While amygdala activation can influence the feeling of fear, it does not create it; fear requires the belief that one is in danger.14
Emotions are conscious experiences: For LeDoux, emotions are primarily conscious experiences, and understanding them fully requires understanding consciousness. Emotions are not basic, pre-wired, reflexive states but are instead constructed cognitively, shaped by the brain’s interpretative processes15 (In this respect, his arguments are similar to Feldman Barrett’s,16 discussed in Part 3). This takes place in cortical circuits, particularly the prefrontal cortex, which LeDoux contends is a region crucial to conscious awareness.17 In contrast to basic emotions theory (discussed in Part 2) and its emphasis on subcortical circuits, he argues that fear (and other emotions) “does not bubble up pre-formed out of ancient circuits [such as the amygdala] but is instead a conscious experience that is cognitively assembled by prefrontal circuits.”18
Fear is personal: LeDoux argues that fear is shaped by a personal "fear schema"—a mental model formed from an individual's experiences with danger.19 In his theory, fear, like all emotions, is personal and requires self-awareness; without awareness of danger, one cannot feel fear. While behavioral reactions to danger are universal, the experience of fear is shaped by how the self-aware brain labels and interprets the threat. Survival circuits in the amygdala intensify the experience, but personal and cultural memories define its nature, with memory playing a key role throughout.20
Memory and emotion interact to produce conscious experience: LeDoux and his collaborator Hakwan Lau propose that conscious experiences arise from the interaction of sensory processes, emotions, memory systems, and higher-order cognitive functions.21 Drawing on psychologist Endel Tulving's memory model, which categorizes memory into anoetic, noetic, and autonoetic types,22 LeDoux and Lau extend these to consciousness:
- Anoetic consciousness involves implicit procedural memory,(22) providing immediate, non-reflective awareness without the use of explicit knowledge or introspection.
- Noetic consciousness relies on semantic memory(22) for factual understanding.
- Autonoetic consciousness uses episodic memory(22) for self-reflective awareness, connecting present experiences with personal past and future events.
LeDoux and Lau highlight that memory not only shapes our immediate conscious perceptions but is also integral to constructing the richer tapestry of our conscious experiences. Emotions, tied intricately to these memory systems, contribute to the ownership of experiences, making conscious states uniquely personal.
Meta-representations shape consciousness: LeDoux and Lau emphasize that all conscious experiences, including emotions, are mediated by meta-representations—higher-order cognitive processes that re-represent lower-order sensory, memory, and emotional information. These meta-representations, processed in areas of the prefrontal cortex, organize and filter sensory inputs, memories, and emotional responses into coherent narratives, forming the content of our conscious experiences. Emotions play a crucial role in shaping these narratives, making them deeply personal and subjective. By integrating past emotional experiences and cognitive appraisals, emotions contribute to the rich, subjective quality of our conscious life, influencing how we perceive and make sense of the world around us.
Two radically different views
The neuroscientist Biyu He23 notes that Damasio’s and LeDoux’s theories represent two radically different views, raising a key question for future research: whether conscious feelings and emotions come mainly from basic survival functions controlled by older (subcortical) brainstem areas or from more advanced thinking processes in newer (cortical) parts of the brain.24
How can you have a feeling without feeling it?
In this four-part series, we have explored and contrasted the leading theories of emotion and have come a little closer toward understanding how emotions give rise to subjective feelings. The centrality of this ongoing scientific quest is highlighted by the following observation by Mark Solms who, referring to the “hard problem of consciousness” as framed by philosopher David Chalmers, makes the point that unlike other forms of information processing, you can’t have a feeling without feeling it:
Chalmers reasonably asked: “Why is the performance of these [cognitive] functions accompanied by experience? Why doesn’t all this information processing go on ‘in the dark,’ free of any inner feel?” […] Chalmers’s question may reasonably be asked of all cognitive functions […] but the same does not apply to affective functions. How can you have a feeling without feeling it? How can we explain the functional mechanism of affect without explaining why and how it causes us to experience something?25
References
1. LeDoux J. E. (2020). Thoughtful feelings. Current biology : CB, 30(11), R619–R623.
3. Homeostasis is the process by which the body keeps its internal environment stable and balanced. This includes maintaining consistent levels of things like temperature, hydration, and pH, through complex feedback mechanisms. A simplified analogy is the way a thermostat keeps the temperature constant in a house. Emotions are integrated with the brain’s homeostatic mechanisms.
[Click 'More' to view footnotes 4-25].
4. In Damasio's theory, feelings are the mental experiences that arise when the brain becomes aware of changes in the body’s state, which are represented in neural maps. These feelings reflect physiological needs, bodily responses, and emotional states, integrating the brain’s perception of these body-state changes into a coherent subjective experience. [Damasio, A., & Carvalho, G. B. (2013). The nature of feelings: evolutionary and neurobiological origins. Nature reviews. Neuroscience, 14(2), 143–152. https://doi.org/10.1038/nrn3403.]
5. The term interoception refers to perception of sensory signals that originate from within the body, especially in the gut and other internal organs, as opposed to exteroception, which refers to perception of stimuli from the external environment.
6. Damasio explains: "Feelings are hybrid processes, based largely on the body’s interior and carried out, physiologically, as an interaction between (i) homeostatically driven, life-regulatory routines occurring in the chemical-visceral interior, and (ii) interoceptive maps generated at multiple peripheral levels (e.g., spinal ganglia; trigeminal ganglion) and central nervous system (e.g., brainstem nuclei; insular cortices; cingulate cortices). Whereas exteroception accomplishes the mapping of external objects and actions that are strictly separate from the perceiving organism, interoception accomplishes the mapping of internal organism events. As a result, an interaction between ‘object of perception’ and ‘subject’ is maintained throughout the process and results in a unique hybrid. Moreover, the anatomical and physiological features of interoception are also unique. The critical step in the generation of consciousness, then, is the revelation, made within the mind, that the mind belongs to and is located within a specific living organism. This revelation is firstly accomplished by feeling but is complemented by (a) proprioceptive signals arising from the musculoskeletal frame, by (b) exteroceptive signals which define organism perspective in coordinate space, and by (c) recall of memorized material related to critical aspects of the organism’s history." [Damasio A. (2021). Feeling & knowing: Making minds conscious. Cognitive neuroscience, 12(2), 65–66. https://doi.org/10.1080/17588928.2020.1846027]
7. In contrast to the exteroceptive and proprioceptive (awareness of body position and movement) nervous systems, which operate via more regulated and precise, but indirect, synaptic transmission.
8. He B. J. (2023). Next frontiers in consciousness research. Neuron, 111(20), 3150–3153. https://doi.org/10.1016/j.neuron.2023.09.042.
9. Earlier versions of his theory had proposed that body maps in the brain were central to creating a sense of self and consciousness. These maps, primarily formed in the cortex, allowed for the internal representation and manipulation of sensory and emotional information, which he believed were necessary for conscious experience. [Damasio, A. (1994). Descartes' error: Emotion, reason, and the human brain. Avon Books.]
10. Damasio, A. (2010). Self comes to mind: Constructing the conscious brain. Pantheon Books.
11. Damasio, A., & Damasio, H. (2023, June 27). Presentation at the National Institutes of Health workshop: Next Frontiers in Consciousness Research. NIH, Bethesda, MD. NIH VideoCast - Neural Mechanisms of Conscious Awareness in Humans (Day 2) (These points were made by Damasio during the Q & A following the presentation, at 6:08:27 in the video recording). Panksepp (if he were still alive), Merker and Solms would probably not disagree fundamentally. The difference is one of emphasis, in terms of the relative importance placed on subcortical versus cortical processes in the generation of conscious emotion. Panksepp, Merker and Solms would argue more vehemently for the sufficiency of brainstem structures in producing basic conscious emotions.
12. The amygdala and interconnected downstream areas, such as the hypothalamus and periaqueductal gray region. All of these, including the amygdala, are subcortical regions. [LeDoux J. E. (2021). What emotions might be like in other animals. Current biology : CB, 31(13), R824–R829. https://doi.org/10.1016/j.cub.2021.05.005.]
13. In certain situations, it can be shown that these automatic responses are not necessarily coupled to a conscious feeling of fear.
14. LeDoux J. E. (2020). Thoughtful feelings. Current biology : CB, 30(11), R619–R623. https://doi.org/10.1016/j.cub.2020.04.012
16. While Feldman Barrett acknowledges the role of subcortical structures like the amygdala, her theory places more emphasis on distributed brain networks and the cortex's role in constructing emotions. But the two of them agree that conscious feelings are a cortical, cognitive construction. LeDoux’s theory proposes a dual-pathway model for processing emotional information. The 'low road' is a fast, subcortical pathway through the amygdala, triggering immediate defensive reactions. The 'high road' involves slower cortical processing, allowing for detailed and conscious appraisal of emotional stimuli. While the amygdala handles rapid responses, the conscious experience of fear arises from activating a 'fear schema' in the prefrontal cortex, which integrates memories and context. [LeDoux J. E. (2021). What emotions might be like in other animals. Current biology : CB, 31(13), R824–R829. https://doi.org/10.1016/j.cub.2021.05.005.]
17. The prefrontal cortex is particularly advanced in humans and supports processes like hierarchical reasoning, prediction, higher-order conceptualization, self-awareness, emotional regulation, attentional control, working memory, and cognitive flexibility—all essential for human conscious experience. LeDoux favors a version of Higher Order Theories of consciousness, which suggests that consciousness arises when the prefrontal cortex re-represents sensory information, transforming it into a conscious experience.
18. A detailed footnote on this point is provided here for those readers who are interested in the cortical-subcortical debate that has been a sub-theme of this four-part series on the neuroscience of emotion:
LeDoux is critical of the claim of researchers like Jaak Panksepp (discussed in Part 2), that the amygdala is the source of conscious fear – and that subcortical circuits are the source of conscious emotional feelings generally. Such claims suggest that emotions are pre-programmed, innate states of mind hard-wired into evolutionarily ancient circuits that are shared by all animals. LeDoux is also critical of the claim by Panksepp and others that animals have human-like feelings, based on their behaving the way we humans do – for example a rat freezing in the presence of danger. LeDoux (and others) argue that findings from animal studies really only reveal how brain circuits control behavioral physiological responses (not subjective experience).
However, LeDoux qualifies his views: “I do not claim that nonhuman animals are mindless robots that lack conscious experiences. My point, instead, is that while it is likely the case that non-human animals have some form of consciousness, it is difficult, maybe impossible, to scientifically study animal consciousness.” Attempting to reduce the divide between his own views and those of Panksepp, LeDoux notes “Just as I am often misunderstood as someone who denies animal consciousness, I think Panksepp’s views about animal emotions have also been misunderstood. Because he was so adamant about animal emotion, it was assumed he meant that emotional experiences in animals are similar to the kinds of emotional experiences that humans refer to with everyday vernacular words, such as fear or sadness. But, in fact, he maintained that emotions in animals are qualitatively different from canonical human emotional experiences.”
LeDoux reinterprets Panksepp’s view of emotion in animals to mean something similar to his own current view: “rudimentary anoetic states of ‘unknowing consciousness’” – i.e., a basic, primitive, non-reflective form of consciousness involving immediate, procedural knowledge and sensory experiences. LeDoux describes these anoetic states as “primitive states of unknowing (content-lacking) consciousness,” “experience that exists as a mental shadow on the border between consciousness and unconsciousness,” or “fringe of consciousness.” [LeDoux J. E. (2021). What emotions might be like in other animals. Current biology : CB, 31(13), R824–R829. https://doi.org/10.1016/j.cub.2021.05.005]. LeDoux is unable to confirm whether Panksepp would have concurred with this reinterpretation, since Panksepp died in 2017.
19. Schemas, according to LeDoux, are mental models or memory structures that include knowledge about the self, the situation, and typical responses to similar situations. They are bundles of conceptually interrelated memories.
20. In LeDoux’s formulation, the self is itself a dynamic, evolving construct of the brain, particularly involving the prefrontal cortex, which integrates various types of information and memories to create a coherent sense of identity. The brain constructs a ‘self-schema’—the collection of memories you have accumulated about yourself. [LeDoux J. E. (2020). Thoughtful feelings. Current biology : CB, 30(11), R619–R623. https://doi.org/10.1016/j.cub.2020.04.012] See also this previous post of mine describing LeDoux’s theory.
21. LeDoux, J. E., & Lau, H. (2020). Seeing consciousness through the lens of memory. Current biology : CB, 30(18), R1018–R1022. https://doi.org/10.1016/j.cub.2020.08.008.
22. TYPES OF MEMORY:
-- Procedural Memory: This is how you remember to do things, like riding a bike. In Tulving's terms, this is anoetic memory, a basic awareness of habits and skills without recalling past experiences.
-- Semantic Memory: This involves facts and general knowledge, like knowing that Paris is the capital of France. Tulving calls this noetic memory, which refers to knowledge you can recall without linking it to any personal event.
-- Episodic Memory: These are personal experiences, like remembering your last birthday. In Tulving's terminology, this is autonoetic memory, which allows you to mentally "travel back in time" to relive specific moments.
23. He B. J. (2023). Next frontiers in consciousness research. Neuron, 111(20), 3150–3153. https://doi.org/10.1016/j.neuron.2023.09.042.
24. This mirrors the tensions between the theories we discussed in Part 2, which emphasized basic hardwired emotions in ancient subcortical circuits and which suggested that these are sufficient for conscious experience of emotion, versus the theories we focused on in Part 3, which emphasized cognitive, cortical construction of emotions and individual and cultural influences. As we have seen, some theorists have attempted an integrative approach, to varying degrees.
25. Solms, M. (2021). The hidden spring: A journey to the source of consciousness (p. 302). Profile Books.
