Fungal Minds

Andrew Adamatzky, Jordi Vallverdu, Antoni Gandia, Alessandro Chiolerio,

Oscar Castro, and Gordana Dodig-Crnkovic

Abstract 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.

A. Adamatzky (B)
Unconventional Computing Lab, UWE, Bristol, UK
e-mail: andrew.adamatzky@uwe.ac.uk
J. Vallverdu · O. Castro
Autonomous University of Barcelona, Catalonia, Spain
e-mail: jordi.vallverdu@uab.cat
A. Gandia
Institute for Plant Molecular and Cell Biology, CSIC-UPV, Valencia, Spain
e-mail: anganfer@alumni.upv.es
A. Chiolerio
Istituto Italiano di Tecnologia, Center for Converging Technologies, Soft Bioinspired Robotics,
Via Morego 30, 16165 Genova, Italy
e-mail: alessandro.chiolerio@iit.it
G. Dodig-Crnkovic
Chalmers University of Technology, Gothenburg, Sweden
e-mail: dodig@chalmers.se

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 409
A. Adamatzky (ed.), Fungal Machines, Emergence, Complexity and Computation 47,
https://doi.org/10.1007/978-3-031-38336-6_26
410 A. Adamatzky et al.

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[1, 2],
it is still controversial to discuss those capacities in other lifeforms such as Fungi,
Protista and Monera, the latest corresponding to single-celled organisms without
true nucleus (particularly bacteria) [3, 4]. However, the current move in research
towards basal cognition and intelligence shows how already unicellular organisms
possess basal levels of cognition and intelligent behaviour [5–11]. Further perplexity
arises when considering consciousness in living organisms. Humans are conscious,
and some allow consciousness in animals provided 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 [12, 13].
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 “Humongous fungus”. Its size is impressive: 910 hectares, possibly
weighing as much as 35,000 tons and having an estimated age of 8,650 years [14].
Identification of Armillaria species used two methods: diploid-diploid pairings, and
restriction fragment length polyporphisms (RFLPs) of the intergenic spacer I (IGS-I)
ribosomal DNA region. A total of 112 Armillaria isolates were collected over two
years campaigns, from six conifer species [15].
Furthermore, current estimations sum up a total of 3.8 million existing species
of fungi, out of which only 120.000 are currently identified [16], 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 impor-
tant 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 dev-
astation (Puccinia graminis responsible for stem, black or cereal rust) [17]. They
have also shown particular features, including interaction with the nervous system of
parasitised superior organisms, to induce them performing actions which are instru-
mental to further fungi propagation. This is the case of Ophiocordyceps sinensis, also
known with its Tibetan name yartsa gumbu, an enthomopathogenic fungus parasitis-
ing ghost moths larvae, that is able to induce them standing vertically under the soil
surface, to facilitate spores spreading in spring times [18]. Similarly, 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 collec-
tively 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 [19].
Fungal Minds 411

By studying Fungi kingdom we can better hope to understand the origin of life
[20] 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 [21–23].

Fungi exhibit trains of action-potential like spikes, detectable by intra-and extra-
cellular recordings [24–26]. In experiments with recording of electrical potential of
oyster fungi Pleurotus djamor (Fig. 1a) we discovered a wide range of spiking activity
(Fig. 1b). Two types were predominant high-frequency (period 2.6 m) and low-freq
(period 14 m) [26]. While studying other species of fungus, Ganoderma resinaceum,
we found that most common width of an electrical potential spike is 5–8 m [27]. In
both species of fungi we observed bursts of spiking in the trains of the spike similar
to that observed in central nervous system [28, 29]. 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 respond to mechanical,
chemical and optical stimulation by changing pattern of its electrically activity and,
in many cases, modifying characteristics of their spike trains [30, 31]. There is also
evidence of electrical current participation in the interactions between mycelium and
plant roots during formation of mycorrhiza [32]. In [33] 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. As per the speed of propagation of information by means of spike-like
structures in the bioelectric signal, we found a spike velocity in slime mold (Physarum
policephalum) ca. 0.08 – 0.17 mm/s [34], in Acetobacter aceti speed between 0.37
and 0.5 mm/s [35], and we know that in superior animals a spike propagates with a
broad spectrum of speeds, ranging from the order of 1 m/s up to 100 m/s.
In [36] we recorded extracellular electrical activity of four species of fungi. We
speculated that fungal electrical activity is a manifestation of the information commu-
nicated between distant parts of the fungal colonies and the information is encoded
into trains of electrical potential spikes. We attempted to uncover key linguistic phe-
nomena 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, how-
ever 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
412 A. Adamatzky et al.

(a)

61,000 62,000 63,000 64,000

8 21,500 22,000 44,000 45,000
3
0.5 2.5
2
2.0
6 1
0 1.5
A E B

4
Electrical potential, mV

2
D
C
4 1 0
2
2
3
6 58,000 59,000
48,000 50,000
20,000 30,000 40,000 50,000 60,000 70,000
Time, sec

(b)

Fig. 1 Recording electrical activity of fungi. a Setup with an array of differential electrodes pairs.
b A variety of patterns of spike trains

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 electroencephalo-
gram as the patient transits from the conscious to the unconscious state [37–40].
In words, a rich spiking activity is a manifestation of consciousness, and reduced
activity of unconsciousness.
In [41] we demonstrated that the electrical activity of the fungus Pleurotus ostrea-
tus 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
Fungal Minds 413

Anaesthetic is introduced Lid is open and anaesthetic is vented out

4.0

3.5

3.0
Potential, mV

2.5

2.0

1.5

1.0

0.5
1 105 2 105 3 105 4 105
Time, sec

Fig. 2 Reduction of spiking activity of Pleurotus ostreatus under influence of chloroform

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 elec-
trical activity restores to a level similar to that before anaesthesia.
To summarise, in experimental laboratory studies of electrical activity of fungi
we demonstrated that fungi produce neuron-like bursts of spikes which are affected
by general anaesthetics. 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 [42] which
sees 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 [5–11, 43]. 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? [44]. 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 expla-
nations 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
414 A. Adamatzky et al.

this process, forcing us to consider the emergence of mind as the coupling between
single individual units and collective behaviour.
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 founda-
tion 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 car-
bon, phosphorus, nitrogen, water, defence compounds, or allelochemicals. Thanks
to this network, plants regulate better their survival, growth, and defence strategies.
Such symbiotic relationship provides fungi carbohydrates, which are used metabol-
ically 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 [45] (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 filamentous nets, they
leave impressions, and mycelia sense and respond to these movements. 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 [46]. Cheating,
besides, decreases when high genetic relatedness exists, a key point for sustaining
multicellular cooperation in fungi [47]. 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 [48] but also for protecting the body from external men-
aces, like somatic parasitism [49, 50]. 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 actions.

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
Fungal Minds 415

starting with humans and investigate its evolutionary origins in other living organ-
isms. Comparing humans with simpler living organisms it might be useful to make
the distinction between primary and higher order consciousness. With minimal mod-
ifications 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 intelli-
gence has been a matter of study for hundreds of years, and still there is an ongoing
controversy on its definition and functional extent [51]. There is a strong resis-
tance and reluctance to acknowledge intelligence and cognitive capacities (including
degree of consciousness) in other living beings.
Fungi can store information and recall it [52]. 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, synchronous 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 [53]. 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 intelligence) are able to perform tasks that we usually would ascribe
to conscious systems, and that consciousness is ubiquitous [54].

5 Biosensing for Data Integrating: The Mechanistic Path

to Fungi Consciousness

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 [55] that we are
416 A. Adamatzky et al.

walking there. This mechanoception process is shared by several living systems,

including plants [56].
Consider for example calcium signal transduction [57]: it has been proven that
in fungi this signalling pathway has an essential role in the survival of fungi [58],
as well as mediate stress responses, or promote virulence [59]. Mechanosensitive
channels can also be important for mating, as we see in Neurosopora Crassa [60].
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 [61]. Furthermore, sensing
capabilities so far described include also nutrient sensing (glucose, nitrogen) and
general chemophysical sensing (pH, temperature, light, gravity, electric field) [55].
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
process, not just a passive one) and to decide output actions. Due to the multimodal
nature of data, this process implies an integration and meaning hierarchies. The
properties of fungi in data integration are shown in Table 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 [91]. 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 stud-
ies: alarmones [92]. These are regulatory molecules used to communicate exclusively
among single cells (not as host). Following [92], pages 13–14, we notice that a static
organism, like a filamentous fungi can live very long and move by hyphal growth,
Fungal Minds 417

Table 1 Different cognitive tasks performed by fungi

Fungal Abilities Evidence
Decision-making and spatial recognition Fungi use an elaborate growth and space-searching strategy
comprising two algorithmic subsets: long-range directional
memory of individual hyphae and inducement of branching by
physical obstruction [62]. The human pathogenic fungus
Candida albicans is able to reorient thigmotropically its
hyphae to find entry points into hosts [63]
Short-term memory 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 [64]. Mycelium of
Phanerochaete velutina remembered the location in which a
bait was placed in a previous test, growing towards the same
direction in a new empty tray [52]. Saturating light stimulus
habituates Phycomyces sporangiophores to a light stimulus
but not to an avoidance stimulus [65]
Long-distance communication Vacuole-mediated long-distance transporting systems support
mycelial foraging and long-distance communication in
saprophytes and mycorrhizal fungi [66, 67]
Photo-tropism Fungi show a rich spectrum of responses to light. They sense
near-ultraviolet, blue, green, red and far-red light using up to
11 photoreceptors [68–71]
Gravi-tropism Gravitropism, as well as thigmotropism, is the strongest
tropims of fungi. This tropism is well studied and
documented [72, 73]
Chemo-tropism and chemical sensitivity 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 direction [74, 75]. Fungal
colonies communicate through volatile compounds [76–78]
Sensing touch and weight Thigmo-based responses, include thigmo differentiation,
thigmonasty, thigmotropism [31, 79–82]
Self versus non-self recognition 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 incompatible [50]
Fighting behaviour Several species of fungi are capable for capturing and
consuming nematodes [83, 84]. Antagonistic interactions
between wood decay basidiomycetes show combative
hierarchies with different species possessing different
combinations of attack and defence traits [85]
Trade behaviour The nutrient exchange mutualism between arbuscular
mycorrhizal fungi (AMFs) and their host plants qualifies as a
biological market [86–89]
Manipulating 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 [18, 19, 90]
418 A. Adamatzky et al.

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
supporting 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, creativity, judgement, are characteristics of human conscious-
ness. Are fungi able of performing these? It might be useful to make the distinction
between “primary” and “higher order” consciousness.
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 completely 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 individuals (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 that fungi sacrifice their bodies for the sake of their propa-
gation. 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 intel-
ligence 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 mor-
phology and degree of connection, we may imagine how radically different compu-
tational schemes are embedded into a fungal consciousness. For example, rather than
3-dimensional visual perception, holographic perception might be possible, consid-
ering 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.
Fungal Minds 419

References

1. Trewavas, A.: The foundations of plant intelligence. Interface Focus. 7, 6 (2017)

2. Calvo, P., Gagliano, M., Souza, G.M., Trewavas, A.: Plants are intelligent, here’s how. Ann.
Bot. 125(1), 11–28 (2020)
3. Westerhoff, H.V., Brooks, A.N., Simeonidis, E., García-Contreras, R., He, F., Boogerd, F.C.,
Jackson, V.J., Goncharuk, V., Kolodkin, A.: Macromolecular networks and intelligence in
microorganisms. Front. Microbiol. 5, 379 (2014)
4. Money, N.P.: Hyphal and mycelial consciousness: the concept of the fungal mind. Fungal Biol.
125(4), 257–259 (2021)
5. Levin, M.: The computational boundary of a ‘self’: developmental bioelectricity drives multi-
cellularity and scale-free cognition. Front. Psychol. 10, 2688 (2019)
6. Levin, M.: Life, death, and self: fundamental questions of primitive cognition viewed through
the lens of body plasticity and synthetic organisms. Biochem. Biophys. Res. Commun. (2020)
(in press)
7. Levin, M., Keijzer, F., Lyon, P., Arendt, D.: Uncovering cognitive similarities and differences,
conservation and innovation. Phil. Trans. R. Soc. B. 376, 20200458 (2021)
8. Lyon, P.: The biogenic approach to cognition. Cogn. Process. 7, 11–29 (2005)
9. Lyon, P.: The cognitive cell: Bacterial behaviour reconsidered. Front. Microbiol. 6, 264 (2015)
10. Ben-Jacob, E.: Bacterial wisdom, gödels theorem and creative genomic webs. Phys. A. 248,
57–76 (1998)
11. Baluska, F., Mancuso, S.: Microorganism and filamentous fungi drive evolution of plant
synapses. Front. Cell. Infect. Microbiol. 3, 44 (2013)
12. Wang, D.Y.-C., Kumar, S., Hedges, S.B.: Divergence time estimates for the early history of
animal phyla and the origin of plants, animals and fungi. Proc. R. Soc. Lond. Ser. B Biol. Sci.
266(1415), 163–171 (1999)
13. Brundrett, M.C.: Coevolution of roots and mycorrhizas of land plants (2002)
14. Schmitt, C.L., Tatum, M.L.: The malheur national forest: Location of the world’s largest living
organism (the Humongous Fungus). United States Department of Agriculture, Forest Service,
Pacific Northwest (2008)
15. Ferguson, B.A., Dreisbach, T.A., Parks, C.G., Filip, G.M., Schmitt, C.L.: Coarse-scale popula-
tion structure of pathogenic Armillaria species in a mixed-conifer forest in the Blue Mountains
of northeast Oregon. Can. J. For. Res. 33, 612–623 (2003)
16. Hawksworth, D.L., Lücking, R.: Fungal diversity revisited: 2.2 to 3.8 million species. Micro-
biol. Spectr. 5(4) (2017)
17. de Mattos-Shipley, K.M.J., Ford, K.L., Alberti, F., Banks, A.M., Bailey, A.M., Foster, G.D.:
The good, the bad and the tasty: the many roles of mushrooms. Stud. Mycol. 85, 125–157
(2016)
18. Yao, Y.-J., Wang, X.-L.: Host insect species of Ophiocordyceps sinensis: a review
19. Hazen, M.L., Loreto, R.G., Mangold, C.A., Chen, D.Z., Fredericksen, M.A., Zhang, Y., Hughes,
D.P.: Three-dimensional visualization and a deep-learning model reveal complex fungal para-
site networks in behaviorally manipulated ants
20. Moore, D.: Fungal Biology in the Origin and Emergence of Life. Cambridge University Press
(2013)
21. Lewicki, M.S.: A review of methods for spike sorting: the detection and classification of neural
action potentials. Netw. Comput. Neural Syst. 9(4), R53 (1998)
22. Baslow, M.H.: The languages of neurons: an analysis of coding mechanisms by which neurons
communicate, learn and store information. Entropy. 11(4), 782–797 (2009)
23. Pruszynski, J.A., Zylberberg, J.: The language of the brain: real-world neural population codes.
Curr. Opin. Neurobiol. 58, 30–36 (2019)
24. Slayman, C.L., Long, W.S., Gradmann, D.: “Action potentials” in Neurospora crassa, a
mycelial fungus. Biochimica et Biophysica Acta (BBA)—Biomembranes. 426(4), 732–744
(1976)
420 A. Adamatzky et al.

25. Olsson, S., Hansson, B.S.: Action potential-like activity found in fungal mycelia is sensitive to
stimulation. Naturwissenschaften 82(1), 30–31 (1995)
26. Adamatzky, A.: On spiking behaviour of oyster fungi pleurotus djamor. Sci. Rep. 8(1), 1–7
(2018)
27. Adamatzky, A., Gandia, A.: On electrical spiking of ganoderma resinaceum. Biophys. Rev.
Lett. 1–9 (2021)
28. Cocatre-Zilgien, J.H., Delcomyn, F.: Identification of bursts in spike trains. J. Neurosci. Meth-
ods. 41(1), 19–30 (1992)
29. Legendy, C.R., Salcman, M.: Bursts and recurrences of bursts in the spike trains of sponta-
neously active striate cortex neurons. J. Neurophysiol. 53(4), 926–939 (1985)
30. Adamatzky, A., Gandia, A., Chiolerio, A.: Fungal sensing skin. Fungal Biol. Biotechnol. 8(1),
1–6 (2021)
31. Adamatzky, A., Nikolaidou, A., Gandia, A., Chiolerio, A., Dehshibi, M.M.: Reactive fungal
wearable. Biosyst. 199, 104304 (2021)
32. Berbara, R.L.L., Morris, B.M., Fonseca, H.M.A.C., Reid, B., Gow, N.A.R., Daft, M.J.: Electri-
cal currents associated with arbuscular mycorrhizal interactions. New Phytol. 129(3), 433–438
(1995)
33. Dehshibi, M.M., Adamatzky, A.: Electrical activity of fungi: spikes detection and complexity
analysis. Biosyst. 203, 104373 (2021)
34. Adamatzky, A., Schubert, T.: Slime mold microfluidic logical gates. Mater. Today. 17(2), 86–91
(2014)
35. Chiolerio, A., Adamatzky, A.: Acetobacter biofilm: electronic characterization and reactive
transduction of pressure. ACS Biomater. Sci. Eng. 7, 1651–1662 (2021)
36. Adamatzky, A.: Language of fungi derived from electrical spiking activity (2021).
arXiv:2112.09907
37. Miu, P., Puil, E.: Isoflurane-induced impairment of synaptic transmission in hippocampal neu-
rons. Exp. Brain Res. 75(2), 354–360 (1989)
38. Hentschke, H., Schwarz, C., Antkowiak, B.: Neocortex is the major target of sedative con-
centrations of volatile anaesthetics: strong depression of firing rates and increase of GABAA
receptor-mediated inhibition. Eur. J. Neurosci. 21(1), 93–102 (2005)
39. Hutt, A., Lefebvre, J., Hight, D., Sleigh, J.: Suppression of underlying neuronal fluctuations
mediates EEG slowing during general anaesthesia. Neuroimage. 179, 414–428 (2018)
40. Sleigh, J.W., Steyn-Ross, D.A., Steyn-Ross, M.L., Grant, C., Ludbrook, G.: Cortical entropy
changes with general anaesthesia: theory and experiment. Physiol. Meas. 25(4), 921 (2004)
41. Adamatzky, A., Gandia, A.: Fungi anaesthesia. Sci. Rep. 12(1), 1–8 (2022)
42. Trewavas, A.: Plant Behaviour and Intelligence. OUP Oxford (2014)
43. Margulis, L., Asikainen, C.A., Krumbein, W.E.: Chimeras and Consciousness: Evolution of
the Sensory Self. MIT Cambridge (2011)
44. Margulis, L.: The conscious cell. Ann. N. Y. Acad. Sci. 929(1), 55–70 (2001)
45. Stamets, P.: Mycelium Running: how Mushrooms can Help Save the World. Random House
Digital, Inc. (2005)
46. Callow, J.A.: Advances in Botanical Research, vol. 22. Elsevier (1999)
47. Bastiaans, E., Debets, A.J.M., Aanen, D.K.: Experimental evolution reveals that high related-
ness protects multicellular cooperation from cheaters. Nat. Commun. 7(1), 1–10 (2016)
48. Scheckhuber, C.Q., Hamann, A., Brust, D., Osiewacz, H.D.: Cellular homeostasis in fungi:
impact on the aging process. In: Aging Research in Yeast, pp. 233–250. Springer (2011)
49. Czaran, T., Hoekstra, R.F., Aanen, D.K.: Selection against somatic parasitism can maintain
allorecognition in fungi. Fungal Genet. Biol. 73, 128–137 (2014)
50. Paoletti, M., Saupe, S.J., Clavé, C.: Genesis of a fungal non-self recognition repertoire. PLoS
one. 2(3), e283 (2007)
51. Cvrcková, F., Lipavská, H., Žárský, V.: Plant intelligence why, why not or where?. Plant Signal.
Behav. 4(5), 394–399 (2009)
52. Fukasawa, Y., Savoury, M., Boddy, L.: Ecological memory and relocation decisions in fungal
mycelial networks: responses to quantity and location of new resources. ISME J. 14(2), 380–
388 (2020)
Fungal Minds 421

53. Harley, J.L., Waid, J.S.: A method of studying active mycelia on living roots and other surfaces
in the soil. Trans. Br. Mycol. Soc. 38(2), 104–118 (1955)
54. Trewavas, A.J., Baluška, F.: The ubiquity of consciousness. EMBO Rep. 12(12), 1221–1225
(2011)
55. Mailänder-Sánchez, D., Braunsdorf, C., Schaller, M.: Fungal sensing of host environment
56. Monshausen, G.B., Haswell, E.S.: A force of nature: molecular mechanisms of mechanoper-
ception in plants. J. Exp. Bot. 64(15), 4663–4680 (2013)
57. Liu, S., Hou, Y., Liu, W., Lu, C., W., Wang, C., Sun, S.: Components of the calcium-calcineurin
signaling pathway in fungal cells and their potential as antifungal targets. Eukaryot. Cell. EC–
00271 (2015)
58. Qilin, Yu., Wang, F., Zhao, Q., Chen, J., Ding, X., Wang, H., Yang, B., Guangqing, L., Zhang,
B., Zhang, B., et al.: A novel role of the vacuolar calcium channel yvc1 in stress response,
morphogenesis and pathogenicity of candida albicans. Int. J. Med. Microbiol. 304(3–4), 339–
350 (2014)
59. Muller, E.M., Mackin, N.A., Erdman, S.E., Cunningham, K.W.: Fig1p facilitates ca2+ influx
and cell fusion during mating of saccharomyces cerevisiae. J. Biol. Chem. (2003)
60. Lew, R.R., Abbas, Z., Anderca, M.I., Free, S.J.: Phenotype of a mechanosensitive channel
mutant, mid-1, in a filamentous fungus, neurospora crassa. Eukaryot. Cell. 7(4), 647–655
(2008)
61. Abbracchio, M.P., Burnstock, G., Verkhratsky, A., Zimmermann, H.: Purinergic signalling in
the nervous system: an overview. Trends Neurosci. 32(1), 19–29 (2009)
62. Hanson, K.L., Nicolau Jr, D.V., Filipponi, L., Wang, L., Lee, A.P., Nicolau, D.V.: Fungi use
efficient algorithms for the exploration of microfluidic networks. Small. 2(10), 1212–1220
(2006)
63. Thomson, D.D., Wehmeier, S., Byfield, F.J., Janmey, P.A., Caballero-Lima, D., Crossley, A.,
Brand, A.C.: Contact-induced apical asymmetry drives the thigmotropic responses of candida
albicans hyphae. Cell. Microbiol. 17(3), 342–354 (2015)
64. Andrade-Linares, D.R., Veresoglou, S.D., Rillig, M.C.: Temperature priming and memory in
soil filamentous fungi. Fungal Ecol. 21, 10–15 (2016)
65. Ortega, J.K.E., Gamow, R.I.: Phycomyces: habituation of the light growth response. Sci.
168(3937), 1374–1375 (1970)
66. Veses, V., Richards, A., Gow, N.A.R.: Vacuoles and fungal biology. Curr. Opin. Microbiol.
11(6), 503–510 (2008)
67. Young, G.: Fungal communication gets volatile. Nat. Rev. Microbiol. 7(1), 6–6 (2009)
68. Carlile, M.J.: The photobiology of fungi. Annu. Rev. Plant Physiol. 16(1), 175–202 (1965)
69. Lipson, E.D.: Phototropism in fungi. In: Biophysics of Photoreceptors and Photomovements
in Microorganisms, pp. 311–325 (1991)
70. Page, R.M.: Phototropism in fungi. In: Photophysiology: Current Topics, p. 65, (2013)
71. Zhenzhong, Yu., Fischer, R.: Light sensing and responses in fungi. Nat. Rev. Microbiol. 17(1),
25–36 (2019)
72. Moore, D.: Perception and response to gravity in higher fungi—a critical appraisal. New Phytol.
117, 3–23 (1991)
73. Corrochano, L.M., Galland, P.: Photomorphogenesis and gravitropism in fungi. In: Growth,
Differentiation and Sexuality, pp. 233–259. Springer (2006)
74. Boddy, L., Jones, T.H.: Mycelial responses in heterogeneous environments: parallels with
macroorganisms. In: Fungi in the Environment, vol. 1, pp. 112–140 (2007)
75. Fomina, M., Ritz, K., Gadd, G.M.: Negative fungal chemotropism to toxic metals. FEMS
Microbiol. Lett. 193(12), 207–211 (2000)
76. Kües, U., Khonsuntia, W., Subba, S., Dörnte, B.: Volatiles in communication of agaricomycetes.
In: Physiology and Genetics, pp. 149–212 (2018)
77. Barriuso, J., Hogan, D.A., Keshavarz, T., Jesús, M., Martínez, J.J.: Role of quorum sensing and
chemical communication in fungal biotechnology and pathogenesis. FEMS Microbiol. Rev.
22, 627–638 (2018)
78. Khalid, S., Keller, N.P.: Chemical signals driving bacterial-fungal interactions (2021)
422 A. Adamatzky et al.

79. Perera, T.H.S., Gregory, D.W., Marshall, D., Gow, N.A.R.: Contact-sensing by hyphae of
dermatophytic and saprophytic fungi. J. Med. Vet. Mycol. 35(4), 289–293 (1997)
80. Brand, A., Gow, N.A.R.: Mechanisms of hypha orientation of fungi. Curr. Opin. Microbiol.
12(4), 350–357 (2009)
81. Almeida, M.C., Brand, A.C.: Thigmo responses: the fungal sense of touch. Microbiol. Spectr.
5(2), 5–2 (2017)
82. Adamatzky, A., Gandia, A.: Living mycelium composites discern weights via patterns of elec-
trical activity. J. Bioresour. Bioprod. 7(1), 26–32 (2022)
83. Barron, G.L., Thorn, R.G.: Destruction of nematodes by species of Pleurotus. Can. J. Bot.
65(4), 774 (1987)
84. Luo, H., Mo, M., Huang, X., Li, X., Zhang, K.: Coprinus comatus: a basidiomycete fungus
forms novel spiny structures and infects nematodes. Mycol. 96, 1218 (2004)
85. Hiscox, J., O’Leary, J., Boddy, L.: Fungus wars: basidiomycete battles in wood decay. Stud.
Mycol. 89(3), 117–124 (2018)
86. Noë, R., Kiers, E.T.: Mycorrhizal markets, firms, and co-ops. Trends Ecol. Evol. 33(10), 777–
789 (2018)
87. Leake, J., Johnson, D., Donnelly, D., Muckle, G., Boddy, L., Read, D.: Networks of power
and influence: the role of mycorrhizal mycelium in controlling plant communities and agro-
ecosystem functioning. Can. J. Bot. 82(8), 1016 (2004)
88. Simard, S.W., Perry, D.A., Jones, M.D., Myrold, D.D., Durall, D.M., Molina, R.: Net transfer
of carbon between ectomycorrhizal tree species in the field. Nat. 388(6642), 579–582 (1997)
89. Simard, S.W.: Mycorrhizal networks facilitate tree communication, learning, and memory
(2018)
90. Shang, Y., Feng, P., Wang, C.: Fungi that infect insects: altering host behavior and beyond.
PLOS Pathog. 11(8), e1005037 (2015)
91. Schenk, H.E.A., Herrmann, R.G., Jeon, K.W., Müller, N.E., Schwemmler, W.: Intertaxonic
combination versus symbiotic adaptation. In: Eukaryotism and Symbiosis. Springer Science
& Business Media (2012)
92. Villarreal, L.P.: Origin of Group Identity: Viruses, Addiction and Cooperation. Springer Science
& Business Media (2008)