Teaching & Learning Guide for: Plant Cognition: A Primer

The study of plant cognition has emerged as a provocative and rapidly developing area at the intersection of philosophy, cognitive science, and plant biology. Long-standing assumptions that restrict cognition to organisms with nervous systems have been challenged by empirical research documenting flexible, context-sensitive, and goal-directed behaviours in plants. These findings raise foundational questions: What counts as cognition? How should it be studied in organisms without neurons? And what philosophical and ethical implications follow if cognition is not restricted to animals? The two companion articles addressed by this guide are designed to be read together. Plant Cognition–A Methodological Primer provides the conceptual and methodological scaffolding needed to navigate the debate, surveying competing theoretical approaches and identifying key epistemic challenges. Plant Cognition—An Empirical Primer complements this by reviewing the current empirical evidence and exploring its philosophical and ethical significance. Together, the articles offer an integrated overview of the field and a case study in how philosophy and empirical science can mutually inform one another. Trewavas, A. 2014. Plant Behavior and Intelligence. Oxford University Press. This book is one of the foundational texts of contemporary plant cognition research. Drawing on decades of experimental plant biology, Trewavas argues that plants exhibit genuine behavior and problem-solving capacities grounded in self-organization, signaling networks, and fitness-relevant responsiveness to environmental challenges. Rather than appealing to neural analogies, the book develops a systems-level perspective in which intelligence is understood as an emergent property of complex, distributed biological organization. Particularly valuable for teaching, the volume combines historical context (including Darwin, Pfeffer, Bose, and McClintock), detailed empirical case studies (e.g., root navigation, foraging, competition, and signaling), and explicit methodological reflections on why plant behavior has been systematically overlooked. The book serves both as a comprehensive empirical resource and as a conceptual bridge between plant science, systems biology, and philosophical debates about the nature and scope of cognition. Baluška, F., S. Mancuso, and D. Volkmann, eds. 2006. Communication in Plants: Neuronal Aspects of Plant Life. Springer. This edited volume is an influential early contribution to the development of what later became known as plant neurobiology. Bringing together work on electrical signaling, long-distance communication, mechanosensation, chemical signaling, and whole-plant integration, the book explores the idea that plants possess sophisticated internal communication systems enabling coordinated, adaptive behavior. For teaching purposes, the volume is best read as a historically and conceptually important landmark rather than as a settled account: many of its claims and metaphors—particularly those invoking neuronal or brain-like analogies—have sparked sustained debate and criticism. As such, the book is especially valuable for illustrating how empirical findings, conceptual framing, and metaphor choice interact in emerging scientific fields and for helping students critically assess both the promise and the pitfalls of extending cognitive language to plant biology. Calvo, P. 2023. Planta Sapiens: The New Science of Plant Intelligence. W. W. Norton (With N. Lawrence). This book provides an accessible, empirically informed introduction to contemporary research on plant behavior and cognition for a broad readership. Written in a popular-science style, it synthesizes findings from plant biology, comparative cognition, and philosophy of psychology to challenge entrenched assumptions about intelligence as necessarily brain-bound. Although the exposition is nontechnical, the arguments are grounded in peer-reviewed research, and each chapter is supported by extensive references that allow interested readers to trace the underlying empirical and theoretical debates in detail. For teaching purposes, the book is particularly useful as an entry point for students encountering plant cognition for the first time, helping to overcome intuitive resistance and “plant blindness” while providing clear signposts to the primary scientific and philosophical literature discussed more systematically in the accompanying academic articles. Chamovitz, D. 2020. What a Plant Knows: A Field Guide to the Senses: Updated and Expanded Edition. Scientific American/Farrar, Straus and Giroux. This book provides an accessible introduction to plant sensory biology for a broad readership, using familiar sensory categories, such as light detection, chemical sensing, and mechanosensation, as an entry point into contemporary plant science. Written in a clear popular-science style, the book translates core findings from molecular and physiological plant biology into conceptually intuitive narratives that help readers understand how plants detect and respond to environmental information. Although not framed explicitly in terms of plant cognition, the volume offers a strong empirical foundation for discussions about plant behavior, signaling, and environmental responsiveness. For teaching purposes, it is especially valuable as a bridge text: it helps students build biological literacy while preparing them to engage more critically with philosophical and cognitive interpretations of plant behavior explored in more specialized literature. Kallhoff, A., M. Di Paola, and M. Schörgenhumer, eds. 2018. Plant Ethics: Concepts and Applications. Routledge. This edited volume is the first systematic, interdisciplinary attempt to develop plant ethics as a distinct field of philosophical inquiry rather than as a marginal extension of environmental or animal ethics. Bringing together contributions from ethics, philosophy of technology, aesthetics, and environmental humanities, the book explores how plants can be ethically significant despite lacking sentience, agency, or consciousness in the animal sense. Particularly valuable for teaching, the volume clarifies key conceptual options—axiological, relational, and practice-based approaches—while explicitly rejecting simplistic extensionist strategies that merely transfer moral categories from animals to plants. The second half of the book grounds these frameworks in concrete applications, including agriculture, forestry, gardening, biotechnology, seed banks, and plant-inspired robotics. The collection provides essential background for understanding the ethical stakes of plant cognition research and offers students a nuanced map of how empirical findings about plant life intersect with questions of value, responsibility, and moral consideration. Shettleworth, S. 2010. Cognition, Evolution, and Behavior. Oxford University Press. This book provides a complete picture of comparative psychology, including its history, its methods, and some of the most important theoretical debates in the discipline. Shettleworth systematically integrates behavioral evidence with evolutionary theory, emphasizing adaptive function, ecological context, and phylogenetic continuity while maintaining a cautious stance toward over-interpretation. In addition, the book is organized by cognitive abilities, from sensing and learning to episodic memory and communication. Even though the book covers nonhuman animals exclusively, it will function as a guide to approach the empirical work being done in comparative psychology, including the most important experimental paradigms. At the same time, the book serves as an excellent grounding in the standards of explanation, inference, and comparison that have shaped modern comparative cognitive science, standards that are especially instructive when evaluating controversial claims about cognition in nonneural organisms. Andrews, K. How to Study Animal Minds. Cambridge University Press, 2020. Although focused on animal cognition, this concise volume offers methodological lessons that translate directly to research on plant cognition. Written in the Cambridge Elements in the Philosophy of Biology format, the book provides a compact and accessible overview of the principles and pitfalls of comparative psychology, including debates about anthropomorphism, Morgan's Canon, bias, and the evidential standards for attributing mental capacities. Of particular relevance for plant cognition research is Andrews's critique of methodological prohibitions that prematurely restrict explanatory vocabulary and her defense of cross-species functional comparison as a legitimate scientific strategy. For teaching purposes, the book functions as an ideal “pocket guide” to comparative methodology, helping students see how challenges traditionally associated with animal minds—such as inference, bias management, and the problem of other minds—re-emerge, often in sharpened form, when cognition is investigated in organisms without nervous systems. https://plantbehavior.org/about-us A long-standing international scientific society is dedicated to research on sensory plant biology, signaling, and communicative ecology. Its mission is to foster interactions among molecular biologists, physiologists, ecologists, and other specialists who study how plants detect, integrate, and act on environmental information, from cellular signaling networks to whole-plant behavior. The society maintains a website with news, conference and symposium information, and links to its flagship journal, Plant Signaling and Behavior. For students and instructors, this site offers a window into an active research community at the intersection of plant science and cognition studies, including announcements of meetings where conceptual and empirical debates about plant behavior are discussed. https://plantae.org Plantae is an online platform sponsored by the American Society of Plant Biologists (ASPB) and other scientific partners, designed to support and connect the global plant science community. The site offers news, research highlights, career resources, webinars, job postings, and discussion forums, making it a lively hub for up-to-date developments in plant biology across scales, from cellular signaling to ecology and evolution. Notably, Plantae curates educational content—such as recorded webinars and teaching tools—that can be used in classroom settings to introduce empirical work on plant behavior and responses to the environment. For instructors and students engaging with plant cognition and comparative philosophy of mind, the platform provides a scientific grounding that complements philosophical analysis, especially by highlighting current lines of research and debates within the broader plant biology community. http://plantinitiative.org The Plant Initiative is an educational and advocacy organization dedicated to promoting greater respect for plants and fostering discourse on plant intelligence, sentience, and human–plant relationships. Although not a scientific society in the traditional academic sense, the site serves as a cross-disciplinary hub connecting philosophical reflection, humanities perspectives, and public engagement with scientific insights about plant life. Users can find free online events, podcasts, recorded conversations, and newsletters that explore themes such as plant communication, plant blindness, ecological partnership, and ethical consideration of vegetal life. The organization also hosts and promotes collaborative networks and offers resources aimed at integrating cultural, ethical, and scientific approaches to understanding plants and their role in human and ecological contexts. For students and instructors, this site provides material that bridges empirical findings with broader questions about human–plant relations and ethical frameworks, making it a valuable complement to scientific and philosophical literature. https://cswr.hds.harvard.edu/research-programming/thinking-plants-fungi The Thinking with Plants & Fungi initiative is an interdisciplinary research and programming effort hosted by the Centee for the Study of World Religions at Harvard Divinity School. It convenes conferences, reading groups, and discussions that explore how inquiry into plant and fungal life can prompt new ways of thinking about mind, ecology, interdependence, and human–more-than-human relationships. Events draw participants from the sciences, humanities, arts, and ecological practice, reflecting an expansive engagement with vegetal life beyond narrow disciplinary boundaries. The initiative's online presence includes information about past and upcoming conferences, reading group resources, and bibliographic suggestions that connect philosophy, ecology, cultural studies, and scientific perspectives on plant and fungal capacities. This resource is especially useful for students and instructors interested in contextualizing plant cognition within broader cultural and ethical conversations about life, agency, and the more-than-human world. https://sites.arizona.edu/plants/resources/organizations This online directory is part of the University of Arizona's Literary and Cultural Plant Studies Network, an interdisciplinary research initiative focused on how plants are conceptualized, represented, and engaged across the humanities and sciences. The organizations page gathers links to a diverse set of external communities, collections, and groups related to plant studies, including botany-oriented archives (e.g., JSTOR Global Plants), speculative or artistic platforms (e.g., Botanical Speculations), interdisciplinary research hubs (e.g., Plant Humanities Initiative at Dumbarton Oaks), and community spaces such as the Plantae Global Plant Science Community. For teaching purposes, this aggregated list functions like a curated index of plant-focused organizations and resource collections across disciplinary boundaries. It offers students an accessible way to explore the breadth of plant studies—from formal scientific communities to cultural, artistic, and philosophical networks—each of which can enrich discussions of plant cognition, agency, and value from multiple perspectives. https://www.um.es/mintlab The MINT Lab (Minimal Intelligence Laboratory) at the University of Murcia is a research group dedicated to understanding intelligence in nonneural organisms and minimal systems, with a particular emphasis on plant cognition and comparative studies that challenge the traditional boundaries of mind. The lab's website showcases its research projects, publications, seminars, and collaborative activities, providing direct access to ongoing work at the crossroads of philosophy, cognitive science, and biology. For students and instructors, the MINT Lab site functions both as a source of primary research outputs (papers, talks, and datasets) and as a platform for academic engagement, including information on events, workshops, and potential collaboration opportunities. Its focus on minimal and distributed forms of intelligence makes the lab's materials especially relevant for modules that examine cognition beyond neural systems and explore methodological and conceptual frameworks for studying “plant minds.” This session introduces the core conceptual dispute surrounding plant cognition by asking what it would even mean for plants to count as cognitive systems. Rather than assuming a settled definition of cognition, the readings expose students to competing strategies for demarcating the cognitive—from pluralistic and deflationary approaches to more restrictive, representation-based accounts—and to the methodological stakes of extending cognitive vocabulary beyond animals. The session sets the stage for the course by framing plant cognition as a live philosophical controversy shaped as much by disagreements about cognitive science itself as by empirical findings about plant behavior. Reading: Allen, C. 2017. “On (Not) Defining Cognition.” Synthese 194, no. 11: 4233–4249. https://doi.org/10.1007/s11229-017-1454-4. Adams, F. R. 2018. “Cognition Wars.” Studies in History and Philosophy of Science Part A 68: 20–30. https://doi.org/10.1016/j.shpsa.2017.11.007. Segundo-Ortin, M., and P. Calvo. 2019. “Are Plants Cognitive? A Reply to Adams.” Studies in History and Philosophy of Science Part A 73: 64–71. https://doi.org/10.1016/j.shpsa.2018.12.001. Figdor, C. 2024. “Why Plant Cognition Is Not (Yet) Out of the Woods. In Philosophy of Plant Cognition: Interdisciplinary Perspectives, edited by G. Ferretti, P. Schulte, and M. Wild, 19–36. Routledge. https://doi.org/10.4324/9781003393375. Lee, J. 2023. “What Is Cognitive About ‘Plant Cognition’?” Biology & Philosophy 38, no. 3: 18. https://doi.org/10.1007/s10539-023-09907-z. This session examines the role of philosophy as a methodological partner in plant science rather than as an external or merely interpretive enterprise. The readings highlight how assumptions about cognition, explanation, and experimental and in biological research. are to philosophy as a for identifying conceptual as of or for how evidential standards are set when studying systems. The session how philosophical can to more and conceptually scientific in the study of plant behavior and cognition. Reading: Calvo, P. Philosophy of Plant A Synthese no. Andrews, and 2014. and the Biology & Philosophy no. This session examines how plants detect and respond to information across and The readings explore the diverse sensory which plants information beyond familiar chemical and The session also examines how the emphasis on shaped frameworks of what counts as behavior. forms of and plant that across and from those of the session students to how plants offer an to the of behavior in biological systems. to these diverse forms of plants as active with behavioral rather than organisms. these materials students to assumptions about plant and to plants as organisms engaged in ongoing behavioral Reading: D. A., W. and G. Biologists Not on What Animal Behavior no. Trewavas, A. 2014. and In Plant Behavior and Intelligence. Oxford University Press. S. Biology no. J. Plant to New no. M., and R. 2014. to by no. M., 2019. to by no. This session introduces students to empirical research that plants exhibit forms of behavioral organization that are using the language of The focus is on how plants information about resource competition, and environmental to in ways that to and Rather than plant responses as or the readings students to plants as systems of evaluating and The session key questions about how such as and can be in nonneural the empirical for broader debates about plant cognition. Reading: Lee, M. Segundo-Ortin, and P. Calvo. 2023. in Plants: A Plants no. M., D. M. and K. 2017. in Plants no. Plants Biology no. A. and Plants Biology no. A. in In edited by M. Springer. M., S. M. and 2019. of by Plant Signaling & Behavior no. This session the discussion of plant cognitive by them within ecological and contexts. are to the idea of communication and memory as distributed extending to ecological on the this session examines including and interactions with and as communication the session students to cognition as an emergent property of and questions about where sensing communication and how biological systems and information Reading: R. and no. R. 2023. by Plants and no. S. W. 2018. and In and in edited by F. M. and G. This session critically examines how debates about plant cognition intersect with broader about evidential and scientific are to the idea that in emerging fields are not empirical also methodological and shaped by bias, and experimental The readings highlight how claims about learning and communication in plants have both and making plant cognition a case study for understanding the in science more The session students to on how scientific to and methodological can be when forms of cognition. Reading: F. 2019. of Science and the Philosophy no. 11: M. D. and J. D. 2023. and to on in & no. M., A. A. M. and M. by in Scientific no. A., J. C. and P. Calvo. 2024. Study of in Plants: of no. This session examines the methodological challenges in approaches from comparative psychology and research to the study of plant cognition while into ecology, and of behavior. It on how methodological when not from animal cognition to plants also from traditional physiological and molecular plant biology questions to experimental aimed at cognitive and information is on with of physiological and key of experimental and as case studies, the session links methodological to the highlighting how grounded experimental and Reading: C. and M. to and A Primer for Biology and & Biology no. C. and A. M. in Plants: in This session explores how debates about plant cognition beyond and into ethical and The readings examine and in what sense, plants moral and how such consideration be without into or are to a of ethical moral and challenge entrenched assumptions about the moral being restricted to animals. The session on how empirical findings about plant behavior and cognition should should inform ethical and on what scientific and toward plant life like of Reading: Calvo. 2017. “What Is It to be a of R. 2018. the of Plants In Plant Ethics: Concepts and edited by A. Kallhoff, M. Di Paola, and M. Schörgenhumer, Routledge. C. and of Studies no. Segundo-Ortin, M., and P. Calvo. 2023. and Animal no. Calvo, and M. 2023. the of Animal no. S. 2014. of Plant of and no. cognition and behavior be of nervous systems and or are the What methodological standards be attributing cognition to behavioral for cognitive or are internal How the plant case the philosophical problem of other claims about plant cognition ethical even in the of a behavioral studied in comparative psychology (e.g., or an that this that a comparative psychology as a guide and the from The is to the for to plants. The should a of why the cognitive and the is for a detailed of the experimental including and and an explicit discussion of potential of bias, and challenges. should also on how theoretical of cognition (e.g., would the same experimental The direct engagement with both empirical and philosophical supported by a by and by and by the by and are supported by the for Plant by The of The have to