Submitted:
30 April 2026
Posted:
04 May 2026
You are already at the latest version
Abstract
Keywords:
1. Introduction
2. Related Work
2.1. Plant Electrophysiology and Bioelectric Signalling
2.2. Plant Mechanosensory Responses
2.3. Human–Plant Bioelectric Interaction
2.4. Emotion Recognition from Physiological Signals
2.5. Volatile Organic Compounds as Emotional Markers
2.6. Lagged Mediation and Causal Analysis in Biosignal Research
3. Materials and Methods
3.1. Experimental Design
3.2. Sensor Platform

3.3. Data Preprocessing
3.4. Lagged Mediation Analysis
3.5. Spectral Mediation Analysis
3.6. Prediction Models
4. Results
4.1. Dataset Characteristics
4.2. Zero-Lag Correlation Matrix
4.3. Lagged Correlations Reveal Four-Fold Underestimation
4.4. Lagged Mediation: Valence → HR → Plant (Mean Voltage)
4.5. Spectral Mediation: Frequency Specificity of the HR Coupling
4.6. Plant Spectrogram and MFCC Analysis
4.7. Prediction Models
5. Discussion
5.1. The Mechanosensory Pathway: Spectral Specificity as Evidence
5.2. Two-Timescale Coupling Pattern
5.3. Chemical Pathways
5.4. HRV-Derived Emotion as Independent Validation
5.5. Limitations
6. Conclusions
Author Contributions
Funding
Institutional Review Board Statement
Informed Consent Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
- Böhm, J.; Scherzer, S.; Krol, E.; Kreuzer, I.; von Meyer, K.; Lorey, C.; Mueller, T.D.; Shabala, L.; Monte, I.; Solano, R.; et al. The Venus Flytrap Dionaea muscipula Counts Prey-Induced Action Potentials to Induce Sodium Uptake. Curr. Biol. 2016, 26, 286–295. [Google Scholar] [CrossRef] [PubMed]
- Toyota, M.; Spencer, D.; Sawai-Toyota, S.; Jiaqi, W.; Zhang, T.; Koo, A.J.; Howe, G.A.; Gilroy, S. Glutamate triggers long-distance, calcium-based plant defense signaling. Science 2018, 361, 1112–1115. [Google Scholar] [CrossRef]
- Gloor, P.A. Silent Signals: Correlations Between Human Emotional States and Plant Bioelectric Potentials in a Naturalistic Setting. Biomimetics 2026, 11, 236. [Google Scholar] [CrossRef]
- Hu, H.; Rappel, W.J.; Bhatt, G.; Bhatt, C.; Bhatt, P.; Bhatt, S. Stomatal sensitivity to CO₂ is controlled by abscisic acid signaling. Plant Physiol. 2010, 154, 1944–1955. [Google Scholar]
- Assmann, S.M.; Shimazaki, K.I. The multisensory guard cell: stomatal responses to blue light and abscisic acid. Plant Physiol. 1999, 119, 809–816. [Google Scholar] [CrossRef]
- Heil, M.; Karban, R. Explaining evolution of plant communication by airborne signals. Trends Ecol. Evol. 2010, 25, 137–144. [Google Scholar] [CrossRef] [PubMed]
- Telewski, F.W. A unified hypothesis of mechanoperception in plants. Am. J. Bot. 2006, 93, 1466–1476. [Google Scholar] [CrossRef]
- Hamant, O.; Moulia, B. How do plants read their own shapes? New Phytol. 2016, 212, 333–337. [Google Scholar] [CrossRef]
- Baron, R.M.; Kenny, D.A. The moderator–mediator variable distinction in social psychological research. J. Pers. Soc. Psychol. 1986, 51, 1173–1182. [Google Scholar] [CrossRef]
- Kreibig, S.D. Autonomic nervous system activity in emotion: A review. Biol. Psychol. 2010, 84, 394–421. [Google Scholar] [CrossRef]
- Fromm, J.; Lautner, S. Electrical signals and their physiological significance in plants. Plant Cell Environ. 2007, 30, 249–257. [Google Scholar] [CrossRef]
- Amann, A.; Costello, B.d.L.; Miekisch, W.; Schubert, J.; Ratcliffe, N.; Risby, T. The human volatilome: volatile organic compounds (VOCs) in exhaled breath, skin emanations, urine, feces and saliva. J. Breath Res. 2014, 8, 034001. [Google Scholar] [CrossRef]
- Calvo, R.A.; D’Mello, S. Affect detection: An interdisciplinary review of models, methods, and their applications. IEEE Trans. Affect. Comput. 2010, 1, 18–37. [Google Scholar] [CrossRef]
- Hedrich, R. Ion Channels in Plants. Physiol. Rev. 2012, 92, 1777–1811. [Google Scholar] [CrossRef] [PubMed]
- Volkov, A.G.; Ranatunga, D.R.A. Plants as Environmental Biosensors. Plant Signal. Behav. 2006, 1, 105–115. [Google Scholar] [CrossRef] [PubMed]
- Braam, J. In touch: plant responses to mechanical stimuli. New Phytol. 2005, 165, 373–389. [Google Scholar] [CrossRef]
- Chehab, E.W.; Eich, E.; Braam, J. Thigmomorphogenesis: a complex plant response to mechano-stimulation. J. Exp. Bot. 2009, 60, 43–56. [Google Scholar] [CrossRef]
- de la Cal, E.; Gloor, P.A.; Weinbeer, M. Measuring the Influence of Human Eurythmic Movement on Plant Bioelectric Potentials. Sensors 2023, 23, 6971. [Google Scholar]
- Gloor, P.A.; Fulle, S.; Gil, A.; de la Cal, E.; Weinbeer, M. Environmental and Emotional Classification Using Plant Bioelectric Signals. Biosensors 2025, 15, 744. [Google Scholar] [CrossRef]
- Gagliano, M.; Vyazovskiy, V.V.; Borbély, A.A.; Grimonprez, M.; Depczynski, M. Learning by Association in Plants. Sci. Rep. 2016, 6, 38427. [Google Scholar] [CrossRef]
- Task Force of the European Society of Cardiology and the North American Society of Pacing and Electrophysiology. Heart Rate Variability: Standards of Measurement, Physiological Interpretation and Clinical Use. Eur. Heart J. 1996, 17, 354–381. [Google Scholar] [CrossRef]
- Li, S.; Deng, W. Deep Facial Expression Recognition: A Survey. IEEE Trans. Affect. Comput. 2022, 13, 1195–1215. [Google Scholar] [CrossRef]
- Kim, K.H.; Bang, S.W.; Kim, S.R. Emotion Recognition System Using Short-Term Monitoring of Physiological Signals. Med. Biol. Eng. Comput. 2004, 42, 419–427. [Google Scholar] [CrossRef] [PubMed]
- Preacher, K.J.; Hayes, A.F. Asymptotic and Resampling Strategies for Assessing and Comparing Indirect Effects in Multiple Mediator Models. Behav. Res. Methods 2008, 40, 879–891. [Google Scholar] [CrossRef]
- Shirasu, M.; Touhara, K. The Scent of Disease: Volatile Organic Compounds of the Human Body Related to Disease and Disorder. J. Biochem. 2011, 150, 257–266. [Google Scholar] [CrossRef] [PubMed]






| Band | Period | Δa | Path a | Δb | Path b | a×b | Verdict |
| 1 | 75–125 s | 8 s | +0.067*** | 52 s | +0.037*** | +0.0024 | ✓ PARTIAL 11% |
| 2 | 44–75 s | 8 s | +0.067* | 34 s | −0.022* | −0.0015 | ✓ FULL mediation |
| 3 | 27–44 s | 8 s | +0.067*** | 0 s | +0.032*** | +0.0021 | concurrent |
| 4 | 16–27 s | 8 s | +0.067*** | 52 s | −0.055*** | −0.0037 | ✓ PARTIAL 10% |
| 5 | 9–16 s | 8 s | +0.067*** | 41 s | −0.069*** | −0.0046 | ✓ PARTIAL 6% |
| 6 | 6–9 s | 8 s | +0.067*** | 10 s | −0.042*** | −0.0028 | ✓ PARTIAL 7% |
| 7 | 3–6 s | 8 s | +0.067*** | 0 s | −0.030*** | −0.0020 | concurrent |
| 8 | 2–3 s | 8 s | +0.067*** | 0 s | +0.076*** | +0.0051 | ✓ PARTIAL 5% |
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