Fungal Biology 125 (2021) 257e259

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Fungal Biology
journal homepage: www.elsevier.com/locate/funbio

Hyphal and mycelial consciousness: the concept of the fungal mind

Nicholas P. Money*
Western Program and Department of Biology, Miami University, Oxford, OH, 45056, USA

a r t i c l e i n f o a b s t r a c t

Article history: Like other cells, fungal hyphae show exquisite sensitivity to their environment. This reactiveness is
Received 3 February 2021 demonstrated at many levels, from changes in the form of the hypha resulting from alterations in pat-
Received in revised form terns of exocytosis, to membrane excitation, and mechanisms of wound repair. Growing hyphae detect
8 February 2021
ridges on surfaces and respond to restrictions in their physical space. These are expressions of cellular
Accepted 11 February 2021
consciousness. Fungal mycelia show decision-making and alter their developmental patterns in response
Available online 23 February 2021
to interactions with other organisms. Mycelia may even be capable of spatial recognition and learning
coupled with a facility for short-term memory. Now is a fruitful time to recognize the study of fungal
Keywords:
Cellular consciousness
ethology as a distinctive discipline within mycology.
Mycelia © 2021 The Author(s). Published by Elsevier Ltd on behalf of British Mycological Society. This is an open
Decomposers access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Fungal pathogens
Wood wide net

Sensitivity or irritability are not the same things as conscious-
ness, but they are starting points in considering whether filamen-
tous fungi have minds. We will begin this inquiry with single
hyphae and move on to fungal colonies or mycelia. Hyphae are thin,
pressurized tubes of cytoplasm adapted for invasive growth and
feeding in solid materials. These microscopic threads elongate at
their tips, where the exocytosis of thousands of vesicles per minute
provides new membrane and cell wall components (Riquelme et al.,
2018). The vesicles are produced by an endomembrane system that
stretches along the hypha and they are guided by the cytoskeleton
to a vesicle supply center (VSC) located close to the tip region.
Vesicles move to the cell surface from this hub and new VSCs are
organized behind the tip to support emerging branches. The
sensitivity of the hypha is clear from the behavior of these VSCs,
which are visible as clouds of vesicles called Spitzenko €rpers in
ascomycete and basidiomycete fungi (Riquelme and Sa nchez-Leo n,
2014). Spitzenko € rpers form and dissolve as hyphae grow and stop
growing, and variations in their size and position are correlated
with changes in the rate of hyphal extension and the shape of the
tip (Reynaga-Pen ~ a et al., 1997). There is nothing artificial about this
intelligence. The continuous flow of information in the live cell
would overwhelm the most complex robot.
Sensitivity is very evident when we manipulate the fungus in
the lab. If the cell membrane is punctured with a glass micropipet,
* Corresponding author
E-mail address: moneynp@miamioh.edu.
the hypha responds by mobilizing cytoplasmic vesicles to repair the
leakage. When the hypha is sliced open with a scalpel blade, more
radical repair mechanisms are deployed. Filamentous ascomycetes
use organelles called Woronin bodies to seal-off hyphal compart-
ments to prevent the whole hypha from rupturing and different
defensive measures have evolved in other groups of fungi (Nguyen
et al., 2021). These patching systems are comparable to blood-
clotting mechanisms in animals.
Recordings of membrane potential using microelectrodes pro-
vide another illustration of hyphal sensitivity (Slayman and
Slayman 1962). The membrane potential is responsive to physical
perturbation of the cell and to changes in the surrounding chemical
environment. Seemingly spontaneous waves of membrane depo-
larization followed by repolarization occur too (Slayman et al.,
1976). These resemble the action potentials or nerve impulses in
animals, but their function in fungi is a mystery. Changes in
membrane potential may be features of a signaling mechanism that
allows hyphae to detect microscopic ridges on surfaces. This
behavior was discovered in rust fungi grown on microfabricated
silicon wafers and serves the fungus in nature as it negotiates the
surface topography of its host plants (Hoch et al., 1987).
Hyphae are reactive to more generalized physical contact be-
tween the cell surface and surrounding environment. Recent ex-
periments have shown that the growth rate and branching patterns
of hyphae are affected by confinement in microfluidic chambers
(Baranger et al., 2020). Hyphae find themselves in similarly cram-
ped situations in nature as they spread in soil and penetrate the
tissues of plants and animals. Indeed, fungi may display unnatural

https://doi.org/10.1016/j.funbio.2021.02.001
1878-6146/© 2021 The Author(s). Published by Elsevier Ltd on behalf of British Mycological Society. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).
N.P. Money
behavior when they grow unconfined in liquid cultures or medium
solidified with agar. These observations place a premium upon
imaging fungal pathogens within the tissues of their hosts (Amich
et al., 2020; Bain et al., 2015; Jones et al., 2016; Pfister et al., 2020).
Taken together, these illustrations of cellular sensitivity
demonstrate that the living hypha is highly attuned and responsive
to its surroundings. The same is true for all cells but has escaped the
attention of mycologists as an evocation of consciousness. Arthur
Reber (2019) has championed the cellular basis of consciousness
and has made powerful arguments against the illogical privilege
that has been accorded to human consciousness. Rather than
seeing the human mind as a unique product of evolution, Reber
regards our consciousness as one expression of irritability along a
continuum of complexity. He is one of several scientists who pro-
pose that consciousness emerges from the perception of environ-
mental conditions, discrimination between helpful and harmful
stimuli, and other universal features of cellular behavior. The
literature on this subject is vast and is rooted in research on protists
in the early 1900s (Jennings 1904; Glasgow 2018). The theory of
cellular consciousness is a work in progress, but it certainly en-
courages us to begin paying more attention to the behavioral so-
phistication of the fungi.
There is a natural tendency to consider an organism conscious if
it appears to engage in decision-making that results in a unique
behavioral outcome. Whether or not humans have free will, we
take actions that seem willful: she finished her coffee, whereas her
friend left her cup half full. On a simpler level, fungi express the
same kind of individualistic behavior all the time. Because the
precise timing and position of branch emergence from a hypha
varies, every germling assumes a unique shape (Hewitt et al., 2016).
This stochastic quality of development is evident from the shape of
clonal colonies growing on a surface and is amplified when the
hyphae proliferate in three dimensions. Although there is a high
degree of predictability in the overall time-dependent form of the
emerging fungus, it seems likely that its detailed shape is irrepro-
ducible. These behavioral differences arise from something akin to
decision-making: the location of a branch is determined by its point
of emergence on the circumference of the primary hypha and the
timing of its inception. Consciousness is a product of the interaction
between the genetic controls that underlie this developmental
process and the responsiveness of the fungus to the physical
environment. The fungus is not thinking in the sense that a brained
animal thinks, but some of the underlying cellular processes of
signal transduction are bound to be homologous.
Fungi that operate as opportunistic pathogens of humans shift
to different growth forms when they enter host tissues, with the
classic yeast-mold transition facilitating invasive growth. More
subtle changes occur in response to the low levels of free iron
within animal tissues. These have been described as anticipatory
behaviors because they offer protection from host immunological
defenses before they are presented to the fungus (Brown et al.,
2019). Pathogens also shield themselves from the host by mask-
ing molecules on their cell surfaces that are recognized by the
innate immune system. Anticipatory behaviors are not comparable
with the classical conditioning of Pavlov’s dogs because they appear
to be hard-wired genetically programmed routines that are always
deployed when the fungus enters the host. Nothing is learned by
1
In some sources the term mycelium (plural mycelium) has been stretched from
its original scientific use as a count noun (each mycelium [singular] expands by
branching/mycelia [plural] expand by branching) into a mass noun that is used,
sloppily in my opinion, as a singular or plural commodity (mycelium [sing. or pl.]
expands by branching). Language evolves, but this swerve in the vocabulary of
mycology needs to be recognized.
258
Fungal Biology 125 (2021) 257e259
the individual fungus, whereas Pavlov’s dogs began to salivate
independently at the sound of the ringing bell during reward-
response experiments. On the other hand, the behavioral pro-
cesses in the dogs and the fungi are founded on the intrinsic genetic
characteristics of their species.
If individual hyphae are conscious, what happens when an
interconnected colony, or mycelium, of thousands of these cells
forms in the soil? 1 Is a mycelium more than a sum of its parts? Can
it be regarded as an integrated conscious entity? Beautiful studies
with radioactive isotopes have demonstrated that materials are
transported between locations across a mycelium, so that one part
of the colony that accesses food can feed distant hyphae that have
exhausted the available nutrients (Fricker et al., 2008). This is an
impressive illustration of resource allocation within a single
mycelium. Source-sink dynamics may explain this process, with
materials simply flowing in response to concentration gradients,
but information may be flowing too (Fricker et al., 2017). Compe-
tition between mycelia of decomposer fungi is likely to be one
interaction that involves a good deal of signaling between clonal
colonies, which can fuse, and different genets that fight for food.
Communication is also essential for the formation and mainte-
nance of mycorrhizal symbioses between fungi and plants
(McClean et al., 2017; Pellegrin et al., 2019; Simard et al., 2012).
Beyond sensitivity and responsiveness, learning and memory
may be regarded as expressions of higher consciousness. These
processes have been explored in slime molds for many years
(Vallverdú et al., 2018) and mycologists are playing catch-up at last.
Mycelia of grassland fungi that are incubated at elevated temper-
atures grow more swiftly after they are exposed to a second, more
severe heat shock than controls that have not experienced the
initial heat shock (Andrade-Linares et al., 2016). This suggests that
compensatory heat shock mechanisms adopted during the first
period of stress are activated immediately when the second shock
occurs. The fungus can remember the first stressful experience for
up to 12 h. Other experiments have shown that a mycelium
growing within a wood block that serves as a home base, or inoc-
ulum, can memorize the relative position of a second wood block
that is used as a bait (Fukasawa et al., 2020). When the original
wood block is removed from its soil tray, disconnecting the inoc-
ulum from the bait, and placed in a fresh soil tray, the mycelium will
tend to reemerge from the side of the wood block from which it had
located the bait. This is an illustration of a simple form of spatial
navigation, memory, and, arguably, intelligence.
Studies on mycelia and mycorrhizas have encouraged the
concept of the forest as a kind of super-organism with a “wood
wide net” formulated by fungal connections between trees. This
awkward allusion to the World Wide Web has some usefulness as a
metaphor, and is an attention grabber, but it does a disservice to the
fungi. In this brief essay I have considered fungal expressions of
consciousness, including sensitivity, decision making, learning, and
memory. This rich behavioral repertoire allows fungi to adapt in
real time to changes in environmental circumstances. Our internet
shows none of this inherent flexibility. It is a network of pathways
that generates nothing on its own. Life outshines the limitations of
this drab technology in every cell. With the wealth of research
revealing the sensitivity and responsiveness of individual hyphae to
their environment, coupled with the novel studies on mycelial
learning and memory, now is a fruitful time to recognize the study
of fungal ethology as a distinctive discipline within mycology.
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