https://doi.org/10.
1093/brain/awac194
Homeostatic feelings and the biology
of consciousness
In a new theory of consciousness, Antonio and Hanna Damasio argue that interoception and the generation
of homeostatic feelings are crucial to understanding how conscious states emerge.
Consciousness occurs when mind contents, such as perceptions and
thoughts, are ‘spontaneously identified as belonging to a specific or-
ganism/owner’. The identification is provided by a continuous flow
of ‘homeostatic feelings’, which translate the process of life regulation
and include salient fluctuations, e.g. hunger, pain, wellbeing, and
states closer to equilibrium, e.g. plain feelings of life/existence.
We propose that homeostatic feelings were the inaugural phe-
nomena of consciousness in biological evolution and were selected
because the spontaneous information they provided regarding the
current state of life regulation conferred extraordinary advantages
to the respective organisms. The ‘knowledge’ carried by conscious
homeostatic feelings enabled overt guidance of life regulation.
We also outline a mechanism for generating homeostatic feelings
based on a two-way interaction between (i) the nervous system,
specifically neural elements of early interoceptive systems; and (ii)
non-neural components of the organism, the body’s interior, name-
ly, viscera and circulating chemical molecules. Feelings emerge from
this interaction, as hybrid and continuous phenomena, related
simultaneously to neural/representational phenomena, and to non-
neural/bodily processes. They identify the organism in which they
inhere by continuously generating the ‘feelingness’ of traditional
philosophical definitions of consciousness.
The problem of consciousness
Neuroscience has been remarkably successful at elucidating the
physiological mechanisms behind some of the most complex func-
tions of living organisms, namely memory, emotions, language,
attention, and intelligence. When it comes to consciousness, how-
ever, the situation could not be more different. Although a variety
of solutions have been proposed, none has emerged as satisfactory.
Strangely, the failure has been accompanied by the notion that the
problem cannot be solved, at least within the realm of biology,1 and
that the solutions, if they exist, might actually be found in the do-
main of physics.2 The multiplicity of conceptions associated with
the term consciousness looms large among the difficulties faced
by these studies. It is often not clear which specific conception is
targeted by the proposed solutions. In search of mechanisms re-
sponsible for subjectivity, the prominent solutions often focus,
Received May 09, 2022. Accepted May 12, 2022. Advance access publication May 30, 2022
© The Author(s) 2022. Published by Oxford University Press on behalf of the Guarantors of Brain. All rights reserved. For permissions, please e-mail:
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instead, on mechanisms underlying other complex processes,
such as attention and integration of information.3–5 That is why
we begin by attempting to make clear what we regard as the es-
sence of consciousness.
Consciousness
Consciousness occurs when the contents of mind—what we per-
ceive and recollect and ponder—are ‘spontaneously identified as
belonging to a specific owner’: our own organism, located in its
body. This definition applies to any ‘scale’ of mind—large and over-
flowing with contents, as when we contemplate a complex prob-
lem, or small, with only a few contents, as when we wake up
from deep sleep. Consciousness makes its appearance when a
mind process—of large scale or small—becomes enriched by
‘knowledge’ that links it to a specific organism. Consciousness
should not be confused with the mere integration of images, or
with the scale of that integration, or with the attention accorded
to some images in detriment of others. ‘We become conscious
when we know, without any question being asked, that the con-
tents of our minds belong to our respective bodies.’
Sensing and minding
The phenomenon of consciousness needs to be distinguished from
the simpler ability to ‘sense’ or ‘detect’ objects and conditions in the
environment.6 Numerous living species, from bacteria to plants,
are capable of sensing/detecting and even responding to what is
sensed. It is likely that sensing, whose physiology hinges on cell
membranes, provides a deep biological foundation for more com-
plex processes of ‘minding’ and ‘consciousness proper’ that subse-
quently appeared in evolution in more complex organisms. But
sensing does not require the sensing organism to possess a nervous
system, nor does the sensing organism make internal representa-
tions of what is sensed or detected via generating ‘mapped pat-
terns’ or ‘images’. Sensing dispenses with representations.
By contrast, we see consciousness as involving and depending
on representations of the kind that constitute minds. The most ob-
vious and common kind of representation takes the form of a
‘mapped pattern’ or ‘image’ of objects, events and qualities (Fig. 1).
2232 | BRAIN 2022: 145; 2231–2235
Figure 1 Photographer Martin Liebscher plays the piano and watches himself play, perhaps the ultimate stage of consciousness! (C-print diasec,
Collection of A. and H. Damasio).
Minds are made of flowing streams of sensory representations
or images constructed with the help of a nervous system.6,7 In es-
sence, the images are of two kinds. The usually dominant kind of
image ‘represents the world surrounding each organism’, a world
which includes other living organisms, varied objects and physical
structures, and their interactions. Those images are based on ex-
teroceptive sensory processes (e.g. vision, hearing, touch, smell
and taste) and map an extraordinary repertoire of the universe ex-
ternal to the organism: perceptions (actual or recalled from mem-
ory), thoughts we hold relative to some item we perceive or
retrieve from memory, the verbal and syntactical translation of
imagetic contents, as well as the conclusions we reach when we
reason about a specific issue by logically manipulating images.
The other kind of images is usually less dominant and originates
in the organism’s interior via interoception. Those images re-
present aspects of viscera and visceral states—a racing heartbeat,
the flow of air in the respiratory system, a gut colic, the contraction
or dilation of blood vessels in the thick of the skin—and include the
intensity and quality, pleasant or unpleasant, of what is perceived,
e.g. the more or less intense burning pain caused by acid indiges-
tion, or the degree of wellbeing caused by extensive relaxation of
smooth and striated muscle fibres throughout the body. Such
images manifest themselves as the class of feelings generated by
life regulation. Those feelings naturally reflect an organism’s man-
agement of existence and are known as homeostatic, to distinguish
them from the emotional feelings caused by the triggering of emo-
tions. As we point out, the physiological apparatus that supports
the making of feelings is radically different from that of exterocep-
tion. (For the sake of brevity, we leave out another perceptual
channel, proprioception, which is primarily concerned with repre-
senting musculoskeletal structures and the movements they
execute but also contributes to homeostatic feelings.)
How is consciousness made: a new hypothesis
We propose that the critical mechanism behind consciousness—
the identification of a particular mind with a particular organism—is
Essay
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naturally provided by the continuous flow of homeostatic feelings
within that organism. Prominent examples of salient homeostatic
feelings include hunger, thirst, pain, wellbeing, malaise, and desire,
as well as subtler feelings of existence that follow unimpeded
regulatory states closer to homeostatic balance. Such feelings are
generated by the ongoing life regulation as it attempts to maintain
operations in the homeostatic range. In brief, homeostatic feelings
as dramatic as ‘pain’ or as subtle as a feeling of ‘existence’, are gen-
erated continuously in the awake state and support the respective
continuity of consciousness, provided they operate above a requis-
ite threshold.7
How and why do feelings provide the critical knowledge that
constitutes consciousness? The answer is transparent: the most
distinctive aspect of the feeling process comes from the fact that
‘each homeostatic feeling is itself spontaneously and automatically
conscious’. When you feel pain or hunger or wellbeing, you are ne-
cessarily conscious of each of these particular states. Your mind is
being offered important and conscious information about how life
is progressing inside your organism and about what the organism
needs. On the basis of that qualitative and graded ‘knowledge’
you can respond accordingly, by making corrections that may
help life continue, e.g. pain signals the possibility of tissue damage
while hunger signals the need for additional energy sources; well-
being serves notice that the organism does not have a specific
need at the moment and can engage in exploration. Had feelings
not been spontaneously conscious they would not have been able
to assist living creatures with curating the life process.
We venture to propose that homeostatic feelings were the in-
augural phenomena of consciousness and that they were selected
and prevailed in biological evolution because the spontaneous
knowledge they provided concerning the current state of life con-
ferred an extraordinary advantage to the organisms that were so
endowed. The information/knowledge carried by conscious
homeostatic feelings enabled organisms to guide life regulation in
an overt manner and behave according to their principal interest:
survival. Homeostatic feelings made deliberate life regulation pos-
sible by going beyond the already existing covert guidance, which
Essay
had been occurring in organisms endowed with sensing/detecting
but not minding and consciousness.
In parallel with advancing life regulation, homeostatic feelings
also helped generate a platform for ‘self-consciousness’. Self-
consciousness emerges when the contents of ongoing perceptions,
the recall of previous perceptions, and the results of reasoning over
such mental contents become connected to a specific organism and
are coherently organized as the mental counterpart of that organ-
ism. Current experiences and a history of past experiences, relative
to a specific organism, produce a conscious being in the full sense of
the term.
Homeostatic feelings are not conscious because they borrow
consciousness from some other process, but because they naturally
produce knowledge about the current life state. As a consequence,
homeostatic feelings can actually ‘lend’ consciousness to other
processes, namely, exteroceptive sensory images. For example,
the retinas, visual pathways and visual cortices produce refined
maps of the outside world and offer us the respective images. But
in order for us to be conscious of those images, we propose that
homeostatic feelings must be present as well, so that such feelings
can identify the visual images that provoke them as belonging to
our respective organism/body. We believe that only then can we be-
come conscious of those images.
Relative to its subjective owner, consciousness emerges from
the homeostatic feelings conjured by many or by few imagetic con-
tents, on any theme. There is no need to invoke an additional mech-
anism to provide consciousness to large arrays of contents.
How do living organisms generate homeostatic
feelings?
At first glance, it might appear that by making homeostatic feelings
responsible for consciousness, we would be moving the problem of
consciousness to a subsidiary domain while leaving the mystery
unsolved. That is not the case, however, because we also propose
a possible mechanism for the generation of homeostatic feelings,
namely, a two-way interaction between (i) the nervous system;
and (ii) non-neural components of the organism.7,8 The typical
homeostatic feelings emerge from this interaction as a novel kind
of phenomenon, in the form of ‘hybrid’ representations that
Figure 2 Artist Fatima Mendonça’s dramatic envisioning of interoception as a dialogue of brain and body. (Gouache and ink on paper, Collection of
A. and H. Damasio).
BRAIN 2022: 145; 2231–2235 | 2233
describe aspects of visceral anatomy and document the qualities
and intensity of visceral events, for example, the duration and in-
tensity of pain. By continuously generating representations of the
state of the organism in which they occur, feelings refer to and
identify the living organism in which they inhere, and thus make
it known in the mind process. Feelings continuously bring con-
sciousness to the minds of ‘their’ organisms because they unmis-
takably connect those minds to the body where feelings ‘happen’.
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The physiological setting of interoception could not be more
distinct from that of exteroception and goes a long way toward
explaining the nature of feelings (Fig. 2). Exteroception produces de-
tailed maps of structures and events external to the organism, while
interoception attempts to describe structures and events inside the or-
ganism. The conditions are ideal for the nervous system to produce
mapped representations of internal structures and events and for al-
lowing those representations to interact directly and bidirectionally
with the structures themselves. For example, neural and chemical sig-
nals can travel from the gut to the brain and produce images, but the
brain can respond to the point of origin of such signals and modify the
local conditions. In the interoceptive world, object and map create a
hybrid—by which we mean that they not only link up with each other
but interact, literally commingle. There is no such equivalent in the
world of exteroception—we cannot have the map of a landscape inter-
act with the landscape itself, which means that they cannot commin-
gle in the same way that they do in the world of interoception. This
unique physiological setting suggests that the ‘feeling imagery’ pro-
duced by interoception ought to be quite distinct from exteroceptive
imagery. We doubt the physiology of consciousness can be properly
understood without considering this particular point.
The distinction becomes even more marked, and physiologically
supported, because the neural elements charged with operating in-
teroceptive interactions are, in of themselves, quite different from
those that assist exteroception. On the neural side we find that in-
teroceptive axons (i) are not insulated by myelin; (ii) make frequent
non-synaptic contacts; and that (iii) interoceptive neurons are not
systematically protected by a blood–brain barrier. On the body side,
we see that the components of the body’s interior, namely viscera
and circulating chemical molecules, can gain direct access to many
interoceptive neural elements thanks to the absence of protective
barriers such as myelin and the blood–brain barrier (Table 1).
2234 | BRAIN 2022: 145; 2231–2235
Table 1 Contrasting the anatomical and physiological characteristics of interoception with those that support exteroception,
movement and cognition
Interoception
Predominant nerve type of axons Poorly myelinated or unmyelinated (Aδ/C)
Mode of signal transmission Both nonsynaptic and synaptic
Time scale Both slow (s/min/h) and fast
Neural processes suported Interoception/visceroception, affect
(moods, emotions, feelings)
Blood–brain barrier Absent or with major gaps
Main neurotransmitters and Monoamine (dopamine, noradrenaline, serotonin),
neuromodulators acetylcholine, neuropeptides
Impaired consciousness
Disorders of consciousness, a common occurrence in clinical neur-
ology, provide an opportunity to interpret the impairment of con-
sciousness in our perspective.
Consciousness can be radically lost in cases of coma caused by
focal damage of the nervous system.7,9,10 The critical site for
lesion-induced coma is the posterior sector of the brainstem region,
where structures such as the nucleus tractus solitarius, the para-
brachial nucleus, and the peri-acqueductal grey collect signals
from the body’s interior. The first integration of feeling-related sig-
nals corresponding to large sectors of the body is accomplished in
this region. The consequence of its damage is a direct comprom-
ise of integrated body representations and a severing of their
links to the exteroceptive mental contents; the latter are largely
assembled in the posterior cerebral cortices and integrated in
cortical association areas. While the cerebral cortex is the base
of operations for creating the predominant imagetic contents of
minds, the subcortical regions are needed to support the feelings
that identify those imagetic contents as belonging to a specific
body. Because of the disparity between the way the CNS handles
interoception and exteroception, small brainstem lesions can
have a devastating effect on consciousness, whereas even exten-
sive lesions of the cerebral cortex may not. Once the integration
of interoceptive information, whose functional culmination in
the CNS occurs at brainstem level, is no longer possible, the world
of exteroception—whose wide base is in the temporal, parietal
and occipital cortices—can no longer be grounded on the body
to which it belongs.
On the matter of impaired consciousness, we must briefly
comment on general anaesthesia, which is by far the most fre-
quent cause of lost consciousness and is both benign and con-
trolled. The anaesthetics that are commonly used in surgery
suspend consciousness rapidly, drastically, and temporarily.
Their physiological point of action is the plain ‘sensing and de-
tecting’ process, located well below the functional level of mind-
ing at which feelings are generated. As noted above, all living
organisms including bacteria, share a basic ability to sense/de-
tect and respond. This ability should not be mistaken for the
more complex phenomena of feeling and consciousness, which
require nervous systems and representations. It is probable,
nonetheless, that sensing/detecting/responding, which depend
on the operation of cell membranes, are the foundational level
of a functional hierarchical chain that eventually enables feeling
and consciousness once nervous systems are present. General
anaesthetics seem to pre-empt the first link in the chain and thus
preclude interoception and exteroception simultaneously. Once
feelings are no longer possible consciousness is also suspended.
Curiously, anaesthetics also interrupt regular life activities in
Essay
Exteroception, movement, cognition
Well myelinated (Aα/Aβ)
Predominantly synaptic
Very fast (μs–ms)
Fine perception, learning, reasoning, calculation,
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language, movement
Continuous
Glutamate, GABA
plants.7,11 At first glance, this is astonishing, since plants do not
have nervous systems or feelings or consciousness. However, be-
cause general anaesthetics can preclude sensing/detecting pro-
cesses, the animation of plants and the complex transactions
executed by their roots and leaves, also come to a halt. Plants become
just as immobile as anaesthetized patients in the operating room,
their animation temporally suspended.
Conclusion
In conclusion, consciousness can be found in many complex living
organisms though not in all. It cannot be found in inanimate ob-
jects, regardless of how complex they may be. Consciousness is
present in living organisms capable of constructing sensory repre-
sentations of components and states of their own bodies, but not in
organisms limited to sensing/detecting. Because nervous systems
are a requirement for the building of mapped imagetic representa-
tions, it follows that only organisms with nervous systems are like-
ly to be conscious. Still, we caution against concluding that because
nervous systems are necessary for consciousness they would be
solely responsible for consciousness, as is commonly assumed.
We believe that consciousness ‘requires a partnership of nervous
systems with the bodies they serve’. Most declarations of failure
relative to the problem of consciousness, claim that ‘neuroscience’
failed to solve the problem but there is no reason why neuroscience
should have solved it alone, because consciousness does not de-
pend on the nervous system alone.
Antonio Damasio and Hanna Damasio
Brain and Creativity Institute, Dornsife College of Letters, Arts and
Sciences, University of Southern California, Los Angeles, USA
Correspondence to: Antonio Damasio
E-mail: damasio@usc.edu
Antonio Damasio is University Professor and David Dornsife Professor of
Neurology, Psychology and Philosophy at the University of Southern
California, where he directs the Brain and Creativity Institute. His most recent
book is Feeling & Knowing: Making Minds Conscious, published by Pantheon.
Hanna Damasio is University Professor and Dana Dornsife Professor of
Neurology and Psychology at the University of Southern California, where she
directs the Dornsife Cognitive Neuroimaging Center.
Competing interests
The authors report no competing interests.
Essay BRAIN 2022: 145; 2231–2235 | 2235

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