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Fungal organisms can perceive the outer world in a way similar to what animals sense. Does that mean that they have full awareness of their environment and themselves? Is a fungus a conscious entity? In laboratory experiments we found that fungi produce patterns of electrical activity, similar to neurons. There are low and high frequency oscillations and convoys of spike trains. The neural-like electrical activity is yet another manifestation of the fungal intelligence. We discuss fungal cognitive capabilities and intelligence in evolutionary perspective, and question whether fungi are conscious and what does fungal consciousness mean, considering their exhibiting of complex behaviours, a wide spectrum of sensory abilities, learning, memory and decision making. We overview experimental evidences of consciousness found in fungi. Our conclusions allow us to give a positive answer to the important research questions of fungal cognition, intelligence and forms of consciousness.
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Fungal States of Minds
Andrew Adamatzkya, Jordi Vallverdub, Antoni Gandiac, Alessandro Chiolerioa,d, Oscar
Castrob, Gordana Dodig-Crnkovice
aUnconventional Computing Lab, UWE, Bristol, UK
bAutonomous University of Barcelona, Catalonia, ES
cInstitute for Plant Molecular and Cell Biology, CSIC-UPV, Valencia, ES
dIstituto Italiano di Tecnologia, Center for Converging Technologies, Soft Bioinspired Robotics, Via
Morego 30, 16165 Genova, IT
eChalmers University of Technology, Gothenburg, SE
Fungal organisms can perceive the outer world in a way similar to what animals sense.
Does that mean that they have full awareness of their environment and themselves? Is a
fungus a conscious entity? In laboratory experiments we found that fungi produce patterns
of electrical activity, similar to neurons. There are low and high frequency oscillations and
convoys of spike trains. The neural-like electrical activity is yet another manifestation of the
fungal intelligence. In this paper we discuss fungal cognitive capabilities and intelligence in
evolutionary perspective, and question whether fungi are conscious and what does fungal
consciousness mean, considering their exhibiting of complex behaviours, a wide spectrum
of sensory abilities, learning, memory and decision making. We overview experimental evi-
dences of consciousness found in fungi. Our conclusions allow us to give a positive answer to
the important research questions of fungal cognition, intelligence and forms of consciousness.
Keywords: fungi, electrophysiology, consciousness, bio-computation, intelligence, memory,
1. The Basic Nature of Fungi
Cognition and intelligence in nature is a topic of debate from multiple points of view.
While nowadays it is acceptable to speak about the cognition and intelligence within the
biological kingdom Animalia, and even up to certain degree in Plantæ[80, 23], it is still con-
troversial to discuss those capacities in other lifeforms such as Fungi, Protista and Monera,
the latest corresponding to single-celled organisms without true nucleus (particularly bacte-
ria) [85, 57]. However, the current move in research towards basal cognition and intelligence
Email addresses: (Andrew Adamatzky),
(Jordi Vallverdu), (Antoni Gandia),
(Alessandro Chiolerio), (Oscar Castro), (Gordana
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shows how already unicellular organisms possess basal levels of cognition and intelligent be-
haviour [43, 44, 45, 51, 52, 16, 11]. Further perplexity arises when considering consciousness
in living organisms. Humans are conscious, and some allow consciousness in animals pro-
vided by a nervous systems. But other living creatures are typically considered not having
consciousness at all. Our focus is on fungi, remarkable organisms with surprising cognitive
capacities and behaviours which can be characterised as intelligent, and this article will
argue that they possess a level of basal consciousness.
Fungi dominated the Earth 600 million years before the arrival of plants [84, 20]. Even
today the largest known living organism in the world is a contiguous colony of Armillaria
ostoyae, found in the Oregon Malheur National Forest, and colloquially known as the “Hu-
mongous fungus”. Its size is impressive: 910 hectares, possibly weighing as much as 35,000
tons and having an estimated age of 8,650 years [71]. Furthermore, current estimations sum
up a total of 3.8 million existing species of fungi, out of which only 120.000 are currently
identified [35], representing a promising biotechnological tool-set from which human kind
has slightly scratched the surface.
Fungi have represented for humans ever since both an ally and a foe, in the first case
serving to produce fermented food and beverages (just to cite the most important one,
Saccharomyces cerevisiae, fundamental for bread, beer and wine) and in the second case,
able to attack the same raw stocks and generate famine and devastation (Puccinia graminis
responsible for stem, black or cereal rust) [29]. They have also shown particular features,
including interaction with the nervous system of parasitised superior organisms, to induce
them performing actions which are instrumental to further fungi propagation. This is the
case of Ophiocordyceps sinensis, also known with its Tibetan name yartsa gumbu, an entho-
mopathogenic fungus parasitising ghost moths larvae, that is able to induce them standing
vertically under the soil surface, to facilitate spores spreading in spring times [86]. Simi-
larly, Ophiocordyceps unilateralis a complex of species also known as “zombie ant fungus”
surrounds muscle fibres inside the ant’s body, and fungal cells form a network used to
collectively control the host behaviour, keeping the brain operative and guiding the ants to
the highest points of the forest canopy, the perfect place to sporulate [55].
By studying Fungi kingdom we can better hope to understand the origin of life[60] and
evolution of cognition, intelligence and consciousness as they gradually emerge from basal
forms and up. But, beyond all the extremely important biochemical mechanisms that make
them possible, a fundamental aspect in their organisation emerges: consciousness.
2. Neuron-like spiking of fungi
Spikes of electrical potential are an essential characteristic of neural activity [47, 14, 68].
Fungi exhibit trains of action-potential like spikes, detectable by intra- and extra-cellular
recordings [75, 63, 2]. In experiments with recording of electrical potential of oyster fungi
Pleurotus djamor (Fig. 1(a)) we discovered a wide range of spiking activity (Fig. 1(b)). Two
types were predominant high-frequency (period 2.6 min) and low-freq (period 14 min) [2].
While studying other species of fungus, Ganoderma resinaceum, we found that most com-
mon width of an electrical potential spike is 5-8 min [4]. In both species of fungi we observed
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Electrical potential, mV
Time, sec
20,000 30,000 40,000 50,000 60,000 70,000
21,500 22,000
58,000 59,000
48,000 50,000
61,000 62,000 63,000 64,000
44,000 45,000
Figure 1: Recording electrical activity of fungi. (a) Setup with an array of differential electrodes pairs. (b) A
variety of patterns of spike trains.
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bursts of spiking in the trains of the spike similar to that observed in central nervous sys-
tem [25, 42]. Whilst the similarly could be just phenomenological this indicates a possibility
that mycelium networks transform information via interaction of spikes and trains of spikes
in manner homologous to neurons. First evidence has been obtained that indeed fungi re-
spond to mechanical, chemical and optical stimulation by changing pattern of its electrically
activity and, in many cases, modifying characteristics of their spike trains [7, 8]. There is
also evidence of electrical current participation in the interactions between mycelium and
plant roots during formation of mycorrhiza [17]. In [30] we compared complexity measures
of the fungal spiking train and sample text in European languages and found that the ’fungal
language’ exceeds the European languages in morphological complexity.
In [3] we recorded extracellular electrical activity of four species of fungi. We speculated
that fungal electrical activity is a manifestation of the information communicated between
distant parts of the fungal colonies and the information is encoded into trains of electrical
potential spikes. We attempted to uncover key linguistic phenomena of the proposed fungal
language. We found that distributions of lengths of spike trains, measured in a number
of spikes, follow the distribution of word lengths in human languages. The size of fungal
lexicon can be up to 50 words, however the core lexicon of most frequently used words
does not exceed 15-20 words. Species Schizophyllum commune and Omphalotus nidiformis
have largest lexicon while species Cordyceps militaris and Flammulina velutipes have less
extensive one. Depending on the threshold of spikes grouping into words, average word
length varies from 3.3 (O. nidiformis) to 8.9 (C. militaris). A fungal word length averaged
over four species and two methods of spike grouping is 5.97 which is of the same range as
an average word length in some human languages, e.g. 4.8 in English and 6 in Russian.
General anaesthetics in mammals causes reduction of neural fluctuation intensity, shift
of electrical activity to a lower frequency spectrum, depression of firing rates, which are also
reflected in a decrease in the spectral entropy of the electroencephalogram as the patient
transits from the conscious to the unconscious state [56, 36, 38, 76]. In words, a rich spiking
activity is a manifestation of consciousness, and reduced activity of unconsciousness.
In [5] we demonstrated that the electrical activity of the fungus Pleurotus ostreatus
is a reliable indicator of the fungi anaesthesia. When exposed to a chloroform vapour
the mycelium reduces frequency and amplitude of its spiking and, in most cases, cease to
produce any electrical activity exceeding the noise level (Fig. 2). When the chloroform
vapour is eliminated from the mycelium enclosure the mycelium electrical activity restores
to a level similar to that before anaesthesia.
To summarise, in experimental laboratory studies of electrical activity of fungi we demon-
strated that fungi produce neuron-like bursts of spikes which are affected by general anaes-
thetics. These phenomena indicate that fungi can posses the same degree of consciousness
as creatures with central nervous system do.
3. Fungal cognition
Once accepting the unity of such a big fungal biological structure as a single living
entity, we need to face a second challenge, that is, the anthropocentric bias [79] which sees
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Anaesthetic is introduced Lid is open and anaesthetic is vented out
Potential, mV
Time, sec
Figure 2: Reduction of spiking activity of Pleurotus ostreatus under influence of chloroform.
consciousness as an exclusively human capacity. This latter is the main cause for the lack
of interest in cognition and intelligence in minimal living systems. At this very moment we
can affirm that intelligence is a property extended across all living taxa, and that we can
even talk about minimal consciousness, starting at microbial level [54, 43, 44, 45, 51, 52, 16,
11]. The functional requests that make possible the existence of such huge fungal colony
are beyond the simple or automated addition of neighbouring cells, but require a level of
cooperation and informational communication that make necessary to ask for a mechanism
that makes possible all these processes, could it be a form of consciousness? [53]. From a
phylogenetic perspective fungi provide the mechanisms for the existence of plant synapses
[11], a fundamental aspect for enabling plants complex information processing.
Our departure point is naturalistic and follows a simple idea: the biological explanations
which can be identified using a functionalist approach supervene on chemical mechanisms;
consequently, any approach to the emergence of informational minds must rely on such
embodied factors. On the other hand, social interactions modify this process, forcing us to
consider the emergence of mind as the coupling between single individual units and collective
Our approach to the study of fungal minds is not a panpsychist one (attributing sentience
to matter), but is based on an informational processing model in which we identify the fungal
self-awareness mechanisms which provide an empirical foundation for the study of fungal
minds. Two questions are orienting our study: are fungi sentient? and...if so, could we talk
about fungal collective consciousness?
From mycorrhizal relations, we know that fungi interact with plants roots and allow the
existence of mycorrhizal networks, used by plants to share or transport carbon, phosphorus,
nitrogen, water, defense compounds, or allelochemicals. Thanks to this network, plants
regulate better their survival, growth, and defence strategies. Such symbiotic relationship
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provides fungi carbohydrates, which are used metabolically to generate energy or to expand
their hyphal networks, generating therefore the collective mycelium. And such mycelium can
be considered the superstructure from which the collective fungal consciousness emerges. As
magisterially described by Stamets [77] (page 4):
The mycelium is an exposed sentient membrane, aware and responsive to changes
in its environment. As hikers, deer, or insects walk across these sensitive filamen-
tous nets, they leave impressions, and mycelia sense and respond to these move-
ments. A complex and resourceful structure for sharing information, mycelium
can adapt and evolve through the ever-changing forces of nature These sensitive
mycelial membranes act as a collective fungal consciousness.
From an evolutionary point of view, mycelia are a clear example of cooperation, but also are
used as a cheating mechanism in relation to host plants [22]. Cheating, besides, decreases
when high genetic relatedness exists, a key point for sustaining multicellular cooperation in
fungi [15]. We’ve considered possible cheating actions of fungi towards their mycorrhizal
hosts, but how can they form collective living forms without cheating among themselves?
The answer is related to the concept of allorecognition. The ability to distinguish self from
non-self is beneficial not only for self-preservation purposes [69] but also for protecting the
body from external menaces, like somatic parasitism [28] [66]. We’ve seen how fungi are able
to distinguish between themselves and others, and how several mechanisms allow them to
work in colonies, to establish symbiotic or parasitic relationships with other living systems.
Their biochemistry allows them to adapt their actions to the informational variations of
the surrounding conditions, and requires a cognitive system able to adaptively manage such
4. Consciousness as self-cognition
When observing cognitive and intelligent behaviour (adequate decision making, learning,
problem solving) of fungi we may ask whether some kind of consciousness enables their goal-
directed behaviour, where consciousness is the ability to make sense of the present situation.
One can search for the consciousness and its markers starting with humans and investigate
its evolutionary origins in other living organisms. Comparing humans with simpler living
organisms it might be useful to make the distinction between primary and higher order con-
sciousness. With minimal modifications we can adapt the notion of “primary consciousness
in humans to describe “primary consciousness“ (that corresponds to “basal cognition“ in
other organisms including fungi and even unicellular organisms.
Over the centuries, consciousness has been a puzzling phenomenon despite all the efforts
of the scientists who tried to unveil its mysteries. Plant cognition and intelligence has
been a matter of study for hundreds of years, and still there is an ongoing controversy
on its definition and functional extent [27]. There is a strong resistance and reluctance to
acknowledge intelligence and cognitive capacities (including degree of consciousness) in other
living beings.
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Fungi can store information and recall it [32]. Fungal memories are procedural, what is
associated with anoetic consciousness. Fungi show self-nonself recognition patterns. Fungi
can navigate and solve mazes looking for a bait. Fungi perceive their environment guiding
themselves to sources of light or higher oxygen concentrations. Self-nonself recognition, syn-
chronous perception of light, nutrients, gravity, or gas, moisture or other chemical gradients,
only involves response to internal or external stimuli, thus these cannot be quoted to support
the affirmation that fungi are conscious or self-aware. Such behaviours can be also encoded
in a computer, which will respond accordingly to the instructed parameters.
Fungus has a role in the managing of such aneuronal consciousness, as has been observed
on tree colonies[34]. The fundamental part of our debate is to show that living systems
without nervous systems (which we usually regard as necessary for self-awareness or intel-
ligence) are able to perform tasks that we usually would ascribe to conscious systems, and
that consciousness is ubiquitous[81].
5. Biosensing for Data Integrating: the Mechanistic Path to Fungi Conscious-
The first aim of consciousness, from a pragmatic point of view is that of collecting and
combining information at some specific level of detail in order to take a decision for action.
Up to now the main focus on consciousness studies has been focused on high level cognitive
processes that follow a top-down hierarchical structure. Using this model, fungi should be
automatically discarded as suitable living systems which could show consciousness. Instead
of it our approach to the notion of consciousness will follow a bottom up approach, from
basic data to its ulterior processing and the possible conscious decisions. We will present
this case from a context situation: Mycelium typically is just under the humus (the soil
cover, i.e. a mix of leaves, needles from pines, fallen branches etc). Thus when we walk in
the forest mycelium “knows“ by mechanical stimulation and stretch-activated receptors [21]
that we are walking there. This mechanoception process is shared by several living systems,
including plants [58].
Consider for example calcium signal transduction [49]: it has been proven that in fungi
this signalling pathway has an essential role in the survival of fungi [88], as well as mediate
stress responses, or promote virulence [61]. Mechanosensitive channels can also be important
for mating, as we see in Neurosopora Crassa [46]. The existence of such mechanosensitive
receptors in fungi make possible to extend some cognitive properties we have already clearly
defined in plants or mammals to fungi. Consider for example the purinergic signalling [1].
Furthermore, sensing capabilities so far described include also nutrient sensing (glucose,
nitrogen) and general chemophysical sensing (pH, temperature, light, gravity, electric field)
[21]. Are those mechanisms a sufficient basis for the grounding of the following questions:
What does mycelium feel about this? Can the mycelium trace our movement? Can the
mycelium predict that we are approaching fruiting bodies (mushrooms)?
5.1. Integration data in fungi systems
All cognitive systems display mechanisms for using captured information (an active pro-
cess, not just a passive one) and to decide output actions. Due to the multimodal nature of
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Fungal abilities Evidence
and spatial recog-
Fungi use an elaborate growth and space-searching strategy compris-
ing two algorithmic subsets: long-range directional memory of individ-
ual hyphae and inducement of branching by physical obstruction [33].
The human pathogenic fungus Candida albicans is able to reorient thig-
motropically its hyphae to find entry points into hosts [78].
Short-term mem-
ory and learning
Fungi exposed to a milder temperature (priming) stress perform better
when exposed to a potentially damaging second heat (triggering) stress.
The priming state in filamentous fungi dissipates over time: memory of
the initial priming stress event for a period of time of at least 24 h [10].
Mycelium of Phanerochaete velutina remembered the location in which
a bait was placed in a previous test, growing towards the same di-
rection in a new empty tray [32]. Saturating light stimulus habituates
Phycomyces sporangiophores to a light stimulus but not to an avoidance
stimulus [64].
Vacuole-mediated long-distance transporting systems support mycelial
foraging and long-distance communication in saprophytes and mycor-
rhizal fungi [82, 87].
Fungi show a rich spectrum of responses to light. They sense near-
ultraviolet, blue, green, red and far-red light using up to 11 photorecep-
tors [24, 48, 65, 89].
Gravi-tropism Gravitropism, as well as thigmotropism, is the strongest tropims of
fungi. This tropism is well studied and documented [59, 26]
and chemical
Fungi are able to detect sources of nutrients and grow towards them
(foraging), in a similar fashion, a fungus would react against a harmful
chemical trace (e.g. toxic metals) by growing towards the opposite direc-
tion [18, 31]. Fungal colonies communicate through volatile compounds
[40, 12, 39].
Sensing touch and
Thigmo-based responses, include thigmo differentiation, thigmonasty,
thigmotropism [67, 19, 9, 8, 6].
Self vs. non-self
Fungi possesses incompatibility loci and a genetic difference at any one
of them is sufficient to trigger destruction of the mixed cell: in most
fungal species, pairs of isolates taken at random are generally incom-
patible [66].
Fighting be-
Several species of fungi are capable for capturing and consuming ne-
matodes [13, 50]. Antagonistic interactions between wood decay basid-
iomycetes show combative hierarchies with different species possessing
different combinations of attack and defence traits [37].
Trade behaviour
The nutrient exchange mutualism between arbuscular mycorrhizal fungi
(AMFs) and their host plants qualifies as a biological market [62, 41,
74, 73].
other organisms
Fungi evolved elaborate tactics, techniques and molecular mechanisms
to control other organisms, from attracting and paralysing nematodes
to programming insects behaviour and death [86, 72, 55]
Table 1: Different cognitive tasks performed by fungi.
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data, this process implies an integration and meaning hierarchies. The properties of fungi
in data integration are shown in Tab. 1.
5.2. The Self: from the genetic view to the perceptual and processing ones
It is necessary to say that the search for published scientific papers on the topics about
“fungal cognition” offers us zero results. The only connections between fungi and cognition
are related most of times to the cognitive impact for humans who are in contact with
fungi. But, of course, fungi do perform cognitive tasks, being self perception one of the
most important.The most basic notion of self identification is related to the delimitation of
the structural elements that belong or not to one system. In this sense, fungi have shown
that their hyphae are able to discriminate self/non-self and that use this skill to decide
to fuse themselves with other genetically compatible hyphae, thanks to anastomosis [70].
The mycelial networks of mycorrhizal fungi can also recognise correctly the roots of their
hosts from those of other surrounding non-hosts. Even different mycorrhizas can coexist
but never fuse together. There is a second mechanism implied in self recognition that is
omitted in most of current studies: alarmones [83]. These are regulatory molecules used
to communicate exclusively among single cells (not as host). Following [83], pages 13-14,
we notice that a static organism, like a filamentous fungi can live very long and move by
hyphal growth, although it is physically constrained and must withstand the onslaught of all
potential genetic parasites they will encounter in their long life. The skill of self-identification
and colonial identity is therefore fundamental for several purposes (such as mating control).
And some fungi even use retro-parasites for their own development.
6. Discussion
We presented experimental laboratory and philosophical studies of the fungal states of
mind. We considered several aspects of fungal cognition and provided arguments support-
ing existence of fungal consciousness. We raised many more questions than we provided
answers. The new field of fungal consciousness is opening in front of us. Let us discuss
directions of future studies. How could we make sure (in wording) that the fungal behaviour
is not mechanical (automatic) responses but that it holds intention? Abstraction, creativ-
ity, judgement, are characteristics of human consciousness. Are fungi able of performing
these? It might be useful to make the distinction between “primary“ and “higher order“
Do fungi have holistic states of mind? Do they combine/modify such states? How many
fungal states of mind could be described? Do fungi create specific relational contexts? Or
are they, on the contrary, not capable of having holistic states of mind, ajust following com-
pletely prefixed patterns? At which extent can we include fungi affects into such cognitive
processing? Fungal chemotaxis could be part of such proto-emotional states.
A deeper consciousness state allows us to understand and accept sacrificing our “self“
for higher purposes (martyrs, heroes). Nevertheless we must recognise that human individ-
uals (and animals as well) are genetically different from each other, with the only exception
of twins, while the same could not apply to fungi. This in part supports the observation
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that fungi sacrifice their bodies for the sake of their propagation. Another thought is about
mortality: though it is, particularly in the western civilisation, uncommon for people to live
everyday life with a constant thought of being mortal, and pursue choices that quite often
go in the opposite direction, as if the individuals are immortal, it is unquestionably difficult
for humans to figure out how the world would appear for immortal beings, like certain fungi.
An immortal, or even extremely old consciousness, would be able to develop perhaps an
intelligence out of reach for us, pursuing objectives that might seem unreasonable, for our
limited perception. Perception of space and time, causality, are all aspects that we consider
our unquestionable bottom line. But given the peculiarity of fungi morphology and degree
of connection, we may imagine how radically different computational schemes are embedded
into a fungal consciousness. For example, rather than 3-dimensional visual perception, holo-
graphic perception might be possible, considering the quasi-flat distribution of mycelia and
its mechanoceptive reconstruction of moving objects (animals) at the upper boundary layer.
Non-causal consciousness might arise from this specific perception framework, eventually
hindering the time perception. All of these remain open questions for further investigations.
7. Acknowledgement
AA and NP were supported by the funding from the European Union’s Horizon 2020
research and innovation programme FET OPEN “Challenging current thinking” under grant
agreement No 858132.
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Fungal construction materials—substrates colonised by mycelium—are getting increased recognition as viable ecologically friendly alternatives to conventional building materials. A functionality of the constructions made from fungal materials would be enriched if blocks with living mycelium, known for their ability to respond to chemical, optical and tactile stimuli, were inserted. We investigated how large blocks of substrates colonised with mycelium of Ganoderma resinaceum responded to stimulation with heavy weights. We analysed details of the electrical responses to the stimulation with weights and show that ON and OFF stimuli can be discriminated by the living mycelium composites and that a habituation to the stimulation occurs. Novelty of the results cast in the reporting on changes in electrical spiking activity of mycelium bound composites in response to a heavy loads.
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Central to the study of cognition is being able to specify the Subject that is making decisions and owning memories and preferences. However, all real cognitive agents are made of parts (such as brains made of cells). The integration of many active subunits into a coherent Self appearing at a larger scale of organization is one of the fundamental questions of evolutionary cognitive science. Typical biological model systems, whether basal or advanced, have a static anatomical structure which obscures important aspects of the mind-body relationship. Recent advances in bioengineering now make it possible to assemble, disassemble, and recombine biological structures at the cell, organ, and whole organism levels. Regenerative biology and controlled chimerism reveal that studies of cognition in intact, “standard”, evolved animal bodies are just a narrow slice of a much bigger and as-yet largely unexplored reality: the incredible plasticity of dynamic morphogenesis of biological forms that house and support diverse types of cognition. The ability to produce living organisms in novel configurations makes clear that traditional concepts, such as body, organism, genetic lineage, death, and memory are not as well-defined as commonly thought, and need considerable revision to account for the possible spectrum of living entities. Here, I review fascinating examples of experimental biology illustrating that the boundaries demarcating somatic and cognitive Selves are fluid, providing an opportunity to sharpen inquiries about how evolution exploits physical forces for multi-scale cognition. Developmental (pre-neural) bioelectricity contributes a novel perspective on how the dynamic control of growth and form of the body evolved into sophisticated cognitive capabilities. Most importantly, the development of functional biobots – synthetic living machines with behavioral capacity – provides a roadmap for greatly expanding our understanding of the origin and capacities of cognition in all of its possible material implementations, especially those that emerge de novo, with no lengthy evolutionary history of matching behavioral programs to bodyplan. Viewing fundamental questions through the lens of new, constructed living forms will have diverse impacts, not only in basic evolutionary biology and cognitive science, but also in regenerative medicine of the brain and in artificial intelligence.