Information and Communication Theoretical Foundations of the Internet of Plants, Principles, Challenges, and Future Directions

Ahmet B. Kilic; Ozgur B. Akan

REVIEW 2 major objections 4 minor 251 references

Information and Communication Theoretical Foundations of the Internet of Plants, Principles, Challenges, and Future Directions

T0 review · 2 major / 4 minor · reviewed 2026-08-04 · deepseek-v4-flash

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Pith's one-line read The paper argues that plant-to-plant signaling—chemical, electrical, fungal, and acoustic—can be systematically modeled as communication links, and that this abstraction opens the way to an Internet of Plants.

desk verdict A genuinely useful tutorial that overstates its own title: the ICT framing is an analogy for at least two of the four modalities, and the paper knows it. read the letter →

arxiv 2509.08434 v1 pith:DLHGMDY4 submitted 2025-09-10 eess.SP

Ahmet B. Kilic , Ozgur B. Akan This is my paper

classification eess.SP

keywords plantcommunicationmolecularvolatileorganiccompoundsmycorrhizalnetworkselectricalsignalingacousticInternetofPlantsinformationtheory

verification ladder T0 review T1 audit T2 compute T3 formal

The pith

A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.

The reading

Plants signal one another by several physical means: volatile chemicals above and below ground, fungal networks, electrical potentials, and sound. This paper tries to show that all four can be described with one engineering language—each signal has a transmitter, a channel, and a receiver—so that questions like noise, interference, channel memory, and coding become well-defined. If the mapping holds, biology and communications engineering can share tools: modelers can compute capacities and intersymbol interference for chemical plumes, fungal webs, and electrical pulses, and engineers can build monitoring systems that read plant stress the way radios read carriers. The paper's larger target is the Internet of Plants, a vision of plants as interconnected nodes in a network that could serve precision agriculture and ecosystem sensing. It positions itself as a tutorial plus survey, collecting the biological evidence and the available mathematical models for each modality.

What carries the argument

The organizing object is the transmitter–channel–receiver decomposition applied to each plant signaling modality. Within it, the load-bearing models are the Green-function solution of the advection–diffusion–reaction equation (the chemical channel's impulse response), the Michaelis–Menten boundary kinetics for root and hyphal exchange, the cable equation and Hodgkin–Huxley-type spike models for electrical propagation, the graph Laplacian for mycorrhizal network diffusion, and the two-state Boltzmann model for mechanosensitive-channel sound reception. These give the paper's claims their quantitative content: parameters like diffusivity, wind, degradation rate, soil moisture, and network conne

What would settle it

A controlled two-room greenhouse experiment where receiver plants are exposed only to recordings of stressed-plant ultrasound while all chemical and electrical contact is blocked; if these receivers show no more defense priming than plants in silence, the paper's acoustic-channel claim loses its empirical footing. The analogous test for mycorrhizal channels is severing the fungal connection while keeping soil chemistry identical.

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Extended reading notes

Core claim

The central claim is that inter-plant communication is not a loose metaphor but a set of physical information channels that can be abstracted into transmitter, channel, and receiver blocks. For aboveground and belowground chemical signaling, the paper assembles emission models, advection–diffusion–reaction propagation kernels, and Robin-boundary uptake models; for electrical signaling, cable equations and action-potential models; for mycorrhizal networks, Michaelis–Menten transfer kinetics and graph-Laplacian network dynamics; and for acoustic signaling, cavitation-based emission and mechanosensitive-channel reception. In every case the same communication-theoretic vocabulary applies: delay

Load-bearing premise

The load-bearing premise is that plant signaling mechanisms are genuine information-carrying channels—not just metaphors—so that treating them as transmitters, channels, and receivers yields real insight; the paper itself concedes that acoustic plant-to-plant communication lacks definitive evidence and that mycorrhizal stress-signal mechanisms remain unclear.

Editorial extensions

If this is right

Aboveground and belowground chemical channels should be treated as memory-heavy: diffusion and sorption tails make intersymbol interference unavoidable, so coding or baseline-subtraction strategies matter more than symbol timing.
Electrical signaling behaves like a fast, sparse digital link: action potentials are near-memoryless binary pulses limited by refractory periods, while variation potentials are analog and ambiguous.
Mycorrhizal networks are topology-dependent channels: graph connectivity, not just diffusivity, sets latency and capacity, so a dense fungal web acts like a low-latency backbone and a sparse one like a bottleneck.
Acoustic signaling is the least established link: the paper treats it as a high-speed but poorly controlled channel and explicitly notes the absence of definitive plant-to-plant evidence.
If the framework holds, plant stress becomes a measurable communication quantity, opening the way to networked monitoring for agriculture and ecosystems—the Internet of Plants.

Reading between the lines

Editorial extensions of the paper, not claims the author makes directly.

If the transmitter–channel–receiver mapping is more than analogy, then VOC blend ratios and stress-state modulation could be analyzed as a real modulation alphabet, meaning information theory could predict how many distinct stress levels a blend can encode—something the paper does not calculate.
The paper's own caveats suggest a near-term research order: build and calibrate end-to-end models for chemical and electrical channels first, while treating acoustic and mycorrhizal links as hypotheses to be confirmed by experiments.
A testable extension would be to estimate actual channel impulse responses in the field—release a controlled VOC pulse, measure arrival at a receiver root, and fit the Green function—to measure capacity and delay spread in natural soil, connecting the paper's framework to measurable quantities.
The Internet of Plants vision depends on solving the multi-signal fusion problem: plants likely combine modalities, so the next step beyond single-channel models is a joint model of how chemical and electrical signals interact in a defense response.

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Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, and a circularity audit.

Desk Editor's Note

read the letter

Colleague,

This is a well-organized tutorial, not a research paper with new information-theoretic results. What it does well: it pulls together chemical, electrical, mycorrhizal, and acoustic plant signaling into one transmitter–channel–receiver taxonomy, gives the standard equations (advection-diffusion, cable equation, Michaelis-Menten, Boltzmann gating) correctly, and surveys the empirical literature and sensing tools in each modality. A communications engineer coming into plant biology will save days. The authors are also honest about biological uncertainty—acoustic plant-to-plant communication is explicitly labeled unconfirmed, mycorrhizal transport mechanisms are described as unclear, and the open-issues lists repeatedly note the absence of end-to-end models for belowground, mycorrhizal, and acoustic links.

The soft spot is the title. “Information and Communication Theoretical Foundations” promises more than the paper delivers. The equations are transport, energetics, and gating descriptions, not information-transfer relations—no alphabets, transition matrices, or capacity estimates are derived for any modality. The ICT framing is an interpretation layered on existing biology, and for acoustic and mycorrhizal links it is currently an analogy. The stress-test note is correct on that point, but it should be said that the paper itself concedes most of it; the mismatch is between the abstract/title and the body, not between the body and reality. The novelty that remains is the synthesis across four modalities and the clear mapping of existing models onto communication blocks—that is a legitimate contribution for a tutorial.

Citation pattern: reliance on the authors’ own prior OMC work (refs 21–23) is fine; those are independently citable and relevant. No code or data, but none is needed for a review.

So: who is this for? Graduate students and researchers in molecular/biological communication who want a map of plant signaling, and plant biologists curious how their field looks through an ICT lens. It deserves a serious referee—mostly to tighten the title, push the authors to mark which modalities have demonstrated channels, and encourage them to state explicitly where the framework is heuristic rather than established.

Recommendation: send to peer review with requested revisions, but treat it as a tutorial, not as foundational theory.

Referee Report

2 major / 4 minor

Summary. This manuscript is a tutorial-with-survey article on interplant communication, covering chemical (above- and belowground VOC), mycorrhizal, electrical, and acoustic modalities. It provides biological background, reformulates each modality as transmitter–channel–receiver blocks, compiles existing mathematical models (advection–diffusion, cable equation, Michaelis–Menten kinetics, graph Laplacian, Boltzmann gating), reviews empirical studies and sensing methods, and outlines the Internet of Plants (IoP) vision. The paper is explicitly positioned as a synthesis rather than a source of new derivations or experiments.

Significance. If the claims are appropriately calibrated, this is a useful interdisciplinary entry point for the communications community. Its strengths are the breadth of the survey, the clear taxonomy of modalities, and the honest enumeration of open issues and missing end-to-end models. The mathematical formulas are standard and generally correctly presented, and the biological descriptions align with cited reviews. There is no machine-checked proof or reproducible code, which is not expected for a tutorial. The main risk is overstatement: the paper calls the result 'Information and Communication Theoretical Foundations' while the central abstraction is largely analogical, and the paper itself concedes that two of the four modalities lack confirmed plant-to-plant communication evidence.

major comments (2)

[Section III, esp. III-B and III-E; Sections II-B and II-D] The central claim—that all four modalities can be abstracted into transmitters, channels, and receivers—is load-bearing but not equally supported. The text itself states that 'definitive evidence of plant-to-plant acoustic communication is lacking' (Section II-D) and that mycorrhizal stress-signal mechanisms 'remain unclear' (Section II-B). Yet Sections III-B and III-E assign full ICT block models to these modalities. Equations (7), (18), and (24) are transport/energetics descriptions, not demonstrated information-transfer relations; no alphabets, channel transition probabilities, or mutual-information/capacity estimates are provided. The paper's own open-issues lists concede the absence of end-to-end models for belowground, mycorrhizal, and acoustic links. The title's 'Foundations' therefore overstates what is established. I recommend reframing the contribution as a perspective/roadmap
[Section III, general] The paper repeatedly invokes information-theoretic concepts (modulation, ISI, capacity, noise, interference) but never computes an information-theoretic quantity. For example, Section III-A.5 describes noise as AWGN-like or colored, and III-B.5 describes severe ISI, but no signal-to-noise ratio, error probability, or capacity estimate is derived for any plant channel. For a tutorial, qualitative mapping is acceptable, but the claim of 'information and communication theoretical foundations' requires at least one quantitative demonstration that the framework is operational. A single worked example—e.g., computing channel memory or a capacity lower bound for a VOC pulse under Eq. (7) with realistic parameters—would substantially strengthen the paper. As written, the title and abstract promise more than the manuscript delivers.

minor comments (4)

[Equation (21)] The resonance-frequency formula is typeset ambiguously: 'fm = m 2 vl L' should be fm = m v_l / (2L). Please correct the notation.
[Reference [73]] The author name 'B. A. Kilic' in reference [73] appears inconsistent with the first author's name elsewhere; please verify.
[Section IV-E / Figure 5] Figure 5 labels interplant acoustic communication as 'remains undetected,' while the text cites one study suggesting this possibility. Please align the wording: e.g., 'no definitive evidence' rather than 'undetected,' to avoid contradicting Section II-D.
[Section III-A.2] The Green's function in Eq. (7) is for instantaneous point release; the text correctly notes continuous emissions matter. A sentence connecting this formula to the later continuous-emission discussion would improve readability for ICT readers.

Circularity Check

0 steps flagged · score 0.0 of 10

No circular derivation: the paper is a tutorial that assembles external models; the few self-citations are not load-bearing.

full rationale

The paper makes no fitted-parameter 'predictions' and derives no new numerical result. Its contribution is a reformulation of existing biological signaling studies into transmitter/channel/receiver blocks. The equations presented are standard transport/electrophysiology models cited to external sources (e.g., advection-diffusion Green's function Eq. (7) [74], Robin boundary condition Eq. (8) [89,90], cable equation Eq. (20) [136], Boltzmann gating Eq. (24) [171]). The authors' own works [21], [23] are used to supply OMC/end-to-end modeling vocabulary, but no conclusion in the paper depends on the truth of a specific numerical prediction from those works; they are literature models, not self-confirming fits. The paper explicitly identifies missing end-to-end models (Section III-C: 'there is no end-to-end model that explains the induction and emission from the plant root...') and missing evidence (Section II-D: 'definitive evidence of plant-to-plant acoustic communication is lacking'). Such admissions are correctness/empirical-support limitations, not circularity. The central claim is a framing claim, so it cannot reduce by construction to its own inputs.

Assumptions & free parameters 6 free parameters · 4 assumptions · 0 invented entities

The paper introduces no new physics or biology; it borrows models from prior literature. The central claim depends on the transferability of these models and the interpretative leap that biological signaling can be quantified as communication channels.

free parameters (6)

VOC production/emission kinetics
Rate constants in Eqs. (1)-(6) (nu_max, w, c, k_d, k_a, k_l, k_g, eta) are left unspecified; they would need calibration for any concrete plant system.
Atmospheric transport parameters
Diffusivity D, wind vector u, and loss rate lambda in Eq. (7) are scenario-dependent and not specified.
Soil effective diffusivity and loss rate
D_eff and k_d in Eq. (14) depend on soil structure, moisture, and microbial activity; no calibration is provided.
Michaelis-Menten constants
Saturation parameters in Eqs. (10), (16), (17), (19) (e.g., K_s, q_max, J_max, K_m, V_max) are cited from root-uptake literature but not specified for the plant communication scenarios.
Cable/membrane constants
Cable equation parameters r_i, r_m, C_m in Eq. (20) are not provided for plant tissues.
Acoustic resonator parameters
Vessel geometry and sap properties in Eqs. (21)-(22) (v_l, L, R, rho_l, eta_l) are species-specific and not given.

assumptions (4)

standard math Standard advection-diffusion-reaction and Green's function solutions describe VOC transport in air and soil.
Used in Sections III-A.2 and III-B.2; these are established physics models.
standard math Cable equation and Hodgkin-Huxley-type dynamics describe plant electrical signaling.
Used in Section III-D.1; borrowed from neurophysiology and applied to plants without validation.
domain assumption Plants emit and respond to chemical VOCs, mycorrhizal networks transfer stress signals, electrical signals propagate between plants, and acoustic emissions carry plant state information.
These biological premises are the basis of the survey; the paper cites empirical studies but does not independently verify them.
domain assumption Communication-theoretic concepts (modulation, ISI, noise, capacity) are meaningful for biological signaling.
Central to the ICT framing in Section III; the paper asserts this but provides no quantitative information-theoretic analysis.

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Cite this review

Pith. "Pith review of Information and Communication Theoretical Foundations of the Internet of Plants, Principles, Challenges, and Future Directions." pith.science (2026). https://pith.science/paper/DLHGMDY4

@misc{pith2026250908434,
  author       = {Pith},
  title        = {Pith review of: Information and Communication Theoretical Foundations of the Internet of Plants, Principles, Challenges, and Future Directions},
  year         = {2026},
  howpublished = {\url{https://pith.science/paper/DLHGMDY4}},
  note         = {Machine review of arXiv:2509.08434}
}

read the original abstract

Plants exchange information through multiple modalities, including chemical, electrical, mycorrhizal, and acoustic signaling, which collectively support survival, defense, and adaptation. While these processes are well documented in biology, their systematic analysis from an Information and Communication Technology (ICT) perspective remains limited. To address this gap, this article is presented as a tutorial with survey elements. It provides the necessary biological background, reformulates inter-plant signaling within ICT frameworks, and surveys empirical studies to guide future research and applications. First, the paper introduces the fundamental biological processes to establish a foundation for readers in communications and networking. Building on this foundation, existing models of emission, propagation, and reception are synthesized for each modality and reformulated in terms of transmitter, channel, and receiver blocks. To complement theory, empirical studies and state-of-the-art sensing approaches are critically examined. Looking forward, the paper identifies open challenges and outlines future research directions, with particular emphasis on the emerging vision of the Internet of Plants (IoP). This paradigm frames plants as interconnected nodes within ecological and technological networks, offering new opportunities for applications in precision agriculture, ecosystem monitoring, climate resilience, and bio-inspired communication systems. By integrating biological insights with ICT frameworks and projecting toward the IoP, this article provides a comprehensive tutorial on plant communication for the communications research community and establishes a foundation for interdisciplinary advances.

Figures

Figures reproduced from arXiv: 2509.08434 by the authors.

**Figure 2.** Illustration of aboveground chemical communication. The process involves VOC storage and emission in the transmitter [PITH_FULL_IMAGE:figures/full_fig_p008_2.png] view at source ↗

**Figure 3.** Modeling of belowground chemical communication: a) overview of the system, b) ICT-based modeling. The link [PITH_FULL_IMAGE:figures/full_fig_p010_3.png] view at source ↗

Figures from the paper (3 more)

**Figure 4.** Figure 4: Overview of Sensing Methods for Aboveground Chem [PITH_FULL_IMAGE:figures/full_fig_p019_4.png]

**Figure 5.** Figure 5: State of experimental studies on plant acoustic communication. (a) Detection of plant stress calls [8]. (b) Plants’ [PITH_FULL_IMAGE:figures/full_fig_p023_5.png]

**Figure 6.** Figure 6: An illustrative figure of the future vision for plant [PITH_FULL_IMAGE:figures/full_fig_p024_6.png]

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