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Plant bioacoustics: The sound expression of stress

Daniel Robert1,*
1School of Biological Sciences, University of Bristol, 24 Tyndall Avenue, Bristol BS8 1TQ, UK

*Correspondence: d.robert@bristol.ac.uk
https://doi.org/10.1016/j.cell.2023.03.004

Plants are not exactly known to be great conversationalists. In this issue of Cell, a new study highlights that
when stressed by desiccation or cutting injury, tomato and tobacco plants can produce airborne ultrasonic
emissions. These sounds are loud enough to be heard by insects and can be analytically categorized using
trained neural networks, pointing to their potential informative value.

Like all organisms, plants are inescapably
coupled to the world of acoustics. Sound,
be it chronic or evanescent, loud or
vanishingly faint, is a pervasive form of
energy in all biomes, propagating in air,
water, living tissue and soil substrates.
From roots to shoots, plants are therefore
subjected to a flow of acoustic energy,
like all matter on Earth. In essence, as
sounds make matter vibrate, vibrating
matter generates sounds.
Whether and how plants are generally
actively involved in the world of sound
and vibrations remains a fertile research
question. Publishing in this issue of Cell,
Khait et al. succeeded at measuring and
characterizing the acoustic emissions of
non-woody tomato and tobacco plants
experiencing physiological stress.1 Plants
placed in different, well-controlled condi-
tions, including a gradually increasing state
of desiccation, or a cut through the stem,
generated ultrasonic clicks that were
remarkably intense, reaching ca. 63dBSPL
(a logarithmic measure of sound pressure
level). For reference, a considerate human
conversation in a living room would take
place at similar sound pressure levels.
Plant sound emissions have been known
about for quite a few years,2 yet their
detection had been restricted to contact
vibrometric measurements, reporting vi-
brations from lignified tissues, such as
branches and trunks from mostly coni-
fers.3–5 The origin of these crackling
sounds has been proposed to stem from
the physical process of cavitation,
whereby abrupt discontinuities in the water
column generated by the ingress of air in
the tree’s xylem vascular system cause en-
ergetic implosions, or vascular embo-
lisms.2 The exact etiology and biophysics
of mechanisms at play remain unknown,
including whether such sounds only ever
happen in situations of stress or result in
actual irreversible injuries to the plant.
The question of acoustic detection
and emission has been investigated and
debated for some time,6–8 with one
emerging consensus being that more
experimental evidence is required to
establish the role of acoustics in plant
behavior and life history. Evidence exists
of sound affecting plant behavior, such
as sonotropism6 and changes in nectar
production9 (Figure 1A), yet many ques-
tions as to the proximate how and ultimate
why of putative acoustic communication
remain open to this day.7 Remarkable
recent evidence demonstrates that
beach-evening primroses (Oenothera
drummondii) enhance nectar concentra-
tion in response to sound playbacks simu-
lating the presence of insect pollinators.9
Albeit not the answer to all questions
about plant acoustic behavior, such evi-
dence provides an insight into the distinc-
tive sensory ecologies of autotroph
organisms that could not be conceived
of before.
Similarly, the new study by Khait et al. of-
fers a fresh and solid view, demonstrating
clear variations in air-borne ultrasonic
emissions propagating from the plants,
and constitutes a most valuable addition
to previous phenomenology (Figures 1B
and 1C). Hence, airborne sound emissions
are not necessarily a matter private to the
plant; other organisms can potentially
exploit them and derive information from
them. This is what evolutionary sensory
exploitation enables; acoustic energy that
inevitably reveals a physiological state
can potentially become an informative
cue within the sensory ecological niche of
other organisms. The distinction between
cue and signal is important, as here no ev-
idence exists thus far that plants broadcast
intentional, evolutionarily adaptive acous-
tic information, a bespoke signal, as they
would do for example using volatiles to
attract the predators of their undesired
herbivore visitors.10
Interestingly, broadening the scope,
Khait et al. also obtained acoustic record-
ings from wheat, corn, young grapevine
branches, and tomato plants infected
with tomato mosaic virus (TMV).1 As
TMV directly affects leaf growth and fruit
yield, this is an important finding. Sound
emissions can now become a useful diag-
nostic feature. Further tests may seek to
establish whether ultrasonic acoustic
emissions take place at early stages of
viral infection and whether those are
distinguishable from emissions occurring
during healthy growth. Time being of the
essence, acoustics may inform presymp-
tomatic treatments that reduce systemic
and population spread of TMC and
reduce cost and impact of chemical
applications.
From a sensory ecological viewpoint,
some horizon scanning might reveal
some far and hitherto hidden features
and implications of plant acoustic emis-
sions. The fact that acoustic energy is
propagating across the surrounding air
makes it potentially available to other or-
ganisms. Cavitation crackling is not private
anymore; the bearers of ultrasound-sensi-
tive ears could potentially exploit such
sounds. Could herbivorous insect or ro-
dent species assess the health of a plant
via cavitation sounds? Could egg-laying
lepidopteran species seek to avoid a plant
under hydration or viral stress? By the
same token, would a crackling noisy plant
be less attractive to herbivores, leafminers,
or other grazers? Further recordings could
test whether acoustically active plants, not
necessarily stressed, incur fewer attacks

Cell 186, March 30, 2023 ª 2023 Elsevier Inc. 1307

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Figure 1. Plant bioacoustics in context: acoustic sensitivity and emission

(A) Plants such as the tobacco plant can respond to the mechanical perturbations generated by incident sounds. Various physiological responses, with different
dynamics (inset) can involve change in growth rate, root navigation, increased nectar production.
(B) Healthy plants without environmental or physiological stress can generate vibrations that transmit only weakly into the surrounding air and may not be
detectable by insects. Here, vibrations are mostly confined to the shell of the plant and detected by surface dwelling herbivorous insects via their subgenual
organs in their mechanically sensitive knee joints.
(C) Plants under stress, such as desiccation or cutting injury, generate air-borne ultrasonic emissions that are loud enough to be detected by insects and other
ultrasound-sensitive organisms.

from natural enemies, offering a test for the
tantalizing thought of aposematic acoustic
signaling. Judicious playbacks of cavita-
tion sounds using species-specific versus
random crackling might even show the
sufficiency of such acoustic emissions.
Finally, in our search for plant varieties
and genetic constructs that are drought
resistant, acoustic emissions could act as
bio-indicators for susceptibility to drought
stress. As such, the study by Khait and col-
leagues offers a gentle, practical, and non-
invasive method to probe the sound
expression of stress.
DECLARATION OF INTERESTS
The author declares no competing interests.
REFERENCES
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