The "plant neurobiology" revolution.

The 21st-century "plant neurobiology" movement is an amalgam of scholars interested in how "neural processes", broadly defined, lead to changes in plant behavior. Integral to the movement (now called plant behavioral biology) is a triad of historically marginalized subdisciplines, namely plant ethology, whole plant electrophysiology and plant comparative psychology, that set plant neurobiology apart from the mainstream. A central tenet held by these "triad disciplines" is that plants are exquisitely sensitive to environmental perturbations and that destructive experimental manipulations rapidly and profoundly affect plant function. Since destructive measurements have been the norm in plant physiology, much of our "textbook knowledge" concerning plant physiology is unrelated to normal plant function. As such, scientists in the triad disciplines favor a more natural and holistic approach toward understanding plant function. By examining the history, philosophy, sociology and psychology of the triad disciplines, this paper refutes in eight ways the criticism that plant neurobiology presents nothing new, and that the topics of plant neurobiology fall squarely under the purview of mainstream plant physiology. It is argued that although the triad disciplines and mainstream plant physiology share the common goal of understanding plant function, they are distinct in having their own intellectual histories and epistemologies.

Consciousness and cognition in plants.

Unlike animal behavior, behavior in plants is traditionally assumed to be completely determined either genetically or environmentally. Under this assumption, plants are usually considered to be noncognitive organisms. This view nonetheless clashes with a growing body of empirical research that shows that many sophisticated cognitive capabilities traditionally assumed to be exclusive to animals are exhibited by plants too. Yet, if plants can be considered cognitive, even in a minimal sense, can they also be considered conscious? Some authors defend that the quest for plant consciousness is worth pursuing, under the premise that sentience can play a role in facilitating plant's sophisticated behavior. The goal of this article is not to provide a positive argument for plant cognition and consciousness, but to invite a constructive, empirically informed debate about it. After reviewing the empirical literature concerning plant cognition, we introduce the reader to the emerging field of plant neurobiology. Research on plant electrical and chemical signaling can help shed light into the biological bases for plant sentience. To conclude, we shall present a series of approaches to scientifically investigate plant consciousness. In sum, we invite the reader to consider the idea that if consciousness boils down to some form of biological adaptation, we should not exclude a priori the possibility that plants have evolved their own phenomenal experience of the world. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Philosophy > Consciousness Neuroscience > Cognition.

Plants have neither synapses nor a nervous system.

The alleged existence of so-called synapses or equivalent structures in plants provided the basis for the concept of Plant Neurobiology (Baluska et al., 2005; Brenner et al., 2006). More recently, supporters of this controversial theory have even speculated that the phloem acts as a kind of nerve system serving long distance electrical signaling (Mediano et al., 2021; Baluska and Mancuso, 2021). In this review we have critically examined the literature cited by these authors and arrive at a completely different conclusion. Plants do not have any structures resembling animal synapses (neither chemical nor electrical). While they certainly do have complex cell contacts and signaling mechanisms, none of these structures provides a basis for neuronal-like synaptic transmission. Likewise, the phloem is undoubtedly a conduit for the propagation of electrical signaling, but the characteristics of this process are in no way comparable to the events underlying information processing in neuronal networks. This has obvious implications in regard to far-going speculations into the realms of cognition, sentience and consciousness.

An approach to the plant body: Assessing concrete and abstract aspects.

The paper aims at proposing a representation of plants as individuals. The first section selects the population of plants to which this study is addressed. The second section describes the effective architecture of plants as modular systems with fixed and mobile elements, in other words, plants and their extensions. The third section presents how plants integrate the fixed and mobile modules into functional units through three areas of particular relevance to plant growth and development: nutrition, defence and pollination. Based on the tangible elements introduced in the previous sections, the fourth section presents the main issue of the proposal which is not apparent at first glance, namely, the local-global relationship in plants' architecture that determines their individuality as organisms. Finally, in the conclusion, we issue the challenge of developing a collective presentation of plants which satisfies their complementary dimension.

Barbara G. Pickard - Queen of Plant Electrophysiology.

Barbara Gillespie Pickard (1936-2019) studied plant electrophysiology and mechanosensory biology for more than 50 y. Her first papers on the roles of auxin in plant tropisms were coauthored with Kenneth V. Thimann. Later, she studied plant electrophysiology. She made it clear that plant action potentials are not a peculiar feature of so-called sensitive plants, but that all plants exhibit these fast electric signals. Barbara Gillespie Pickard proposed a neuronal model for the spreading of electric signals induced by mechanical stimuli across plant tissues. In later years, she studied the stretch-activated plasma membrane channels of plants and formulated the plasma-membrane control center model. Barbara Pickard summarized all her findings in a new model of phyllotaxis involving waves of auxin fluxes and mechano-sensory signaling.

Integrated information theory does not make plant consciousness more convincing.

It has been proposed by some plant scientists that plants are cognitive and conscious organisms, although this is a minority view. Here we present a brief summary of some of the arguments against this view, followed by a critique of an article in this same issue of Biochemical and Biophysical Research Communications by Calvo, Baluska, and Trewavas (2020) that cites Integrated Information Theory (IIT) as providing additional support for plant consciousness. The authors base their argument on the assumptions that all cells are conscious and that consciousness is confined to life. However, IIT allows for consciousness in various nonliving systems, and thus does not restrict consciousness to living organisms. Therefore, IIT cannot be used to prove plant consciousness, for which there is neither empirical evidence nor support from other, neuron-based, theories of consciousness.

American racism and the lost legacy of Sir Jagadis Chandra Bose, the father of plant neurobiology.

Sir Jagadis Chandra Bose, India's first modern biologist departed boldly from mainstream botany by claiming that plants possess "nerves" and "pulsating cells" that function respectively much like the nerve and heart cells of animals. These ideas were based on highly sensitive measurements he made of various plant functions by means of assorted ingenious instruments of his own design. Despite being the most internationally celebrated plant biologist of the early 20th century, by the end of his life, Bose had become a scientific pariah whose work was expunged from Western histories of plant biology for nearly a century. In the 21st century, Bose's contributions to biology have begun to be appreciated anew, particularly within the plant neurobiology community. The present contribution examines the motivating factors behind the anti-Bose camp in the United States in the 1920s. It is concluded that the opposition to Bose's ideas during this period had less to do with scientific dialectics than with jealousy over Bose's international acclaim and the prevailing racism of the era.

Jagdish Chandra Bose & plant neurobiology.

When Jagdish Chandra Bose, a renowned physicist, devoted himself entirely to research in the field of plant physiology post his superannuation at Presidency University, Kolkata, India (earlier known as Presidency College, Calcutta), it came as a surprise to many. The research on plant nervous system by JC Bose during this period was pioneering in nature, being recognized by recent plant biologists globally as the first in the field. His findings were so revolutionary at the time of their proclamation that these aroused disbelief and contradiction. Surprisingly, not many at that time took up such investigations and once accepted with reluctance, there was practically very little activity in the field for the next several decades. More than a hundred years later, recent advances in molecular biology, genomics, ecology and neurophysiology have led to renewed interest resulting in a flurry of activity, confirming most of Bose's observations. The present review describes this pioneering scientist's work and his immense contribution in the emergence of the discipline now designated as 'Plant Neurobiology'.

Plants Neither Possess nor Require Consciousness.

In claiming that plants have consciousness, 'plant neurobiologists' have consistently glossed over the remarkable degree of structural and functional complexity that the brain had to evolve for consciousness to emerge. Here, we outline a new hypothesis proposed by Feinberg and Mallat for the evolution of consciousness in animals. Based on a survey of the brain anatomy, functional complexity, and behaviors of a broad spectrum of animals, criteria were established for the emergence of consciousness. The only animals that satisfied these criteria were the vertebrates (including fish), arthropods (e.g., insects, crabs), and cephalopods (e.g., octopuses, squids). In light of Feinberg and Mallat's analysis, we consider the likelihood that plants, with their relative organizational simplicity and lack of neurons and brains, have consciousness to be effectively nil.

The botanical multiverse of Peter Barlow.

Dr Peter Barlow, who died in 2017, was one of the most respected botanists and biologists of the latter half of the 20th Century. His interests covered a wide range of plant biological topics, e.g. root growth and development, plant cytoskeleton, effects of gravity, plant intelligence, pattern formation, and evolution of eukaryotic cells. Here we consider Peter's numerous contributions to the: elucidation of plant patterns; understanding of root biology; role of the plant cytoskeleton in growth and development; influence of the Moon on terrestrial vegetation; Cell Body concept; and plant neurobiology. In so doing we attempt not only to provide an overview of Peter's important work in many areas of plant biology, but also to place that work in the context of recent advances in plant and biological sciences.

Cotton Fabric Coated with Conducting Polymers and its Application in Monitoring of Carnivorous Plant Response.

The paper describes the electrical plant response to mechanical stimulation monitored with the help of conducting polymers deposited on cotton fabric. Cotton fabric was coated with conducting polymers, polyaniline or polypyrrole, in situ during the oxidation of respective monomers in aqueous medium. Thus, modified fabrics were again coated with polypyrrole or polyaniline, respectively, in order to investigate any synergetic effect between both polymers with respect to conductivity and its stability during repeated dry cleaning. The coating was confirmed by infrared spectroscopy. The resulting fabrics have been used as electrodes to collect the electrical response to the stimulation of a Venus flytrap plant. This is a paradigm of the use of conducting polymers in monitoring of plant neurobiology.

The 'root-brain' hypothesis of Charles and Francis Darwin: Revival after more than 125 years.

This year celebrates the 200(th) aniversary of the birth of Charles Darwin, best known for his theory of evolution summarized in On the Origin of Species. Less well known is that, in the second half of his life, Darwin's major scientific focus turned towards plants. He wrote several books on plants, the next-to-last of which, The Power of Movement of Plants, published together with his son Francis, opened plants to a new view. Here we amplify the final sentence of this book in which the Darwins proposed that: "It is hardly an exaggeration to say that the tip of the radicle thus endowed [with sensitivity] and having the power of directing the movements of the adjoining parts, acts like the brain of one of the lower animals; the brain being seated within the anterior end of the body, receiving impressions from the sense-organs, and directing the several movements." This sentence conveys two important messages: first, that the root apex may be considered to be a 'brain-like' organ endowed with a sensitivity which controls its navigation through soil; second, that the root apex represents the anterior end of the plant body. In this article, we discuss both these statements.

Plant neurobiology as a paradigm shift not only in the plant sciences.

Plants are complex living beings, extremely sensitive to environmental factors, continuously adapting to the ever changing environment. Emerging research document that plants sense, memorize, and process experiences and use this information for their adaptive behavior and evolution. As any other living and evolving systems, plants act as knowledge accumulating systems. Neuronal informational systems are behind this concept of organisms as knowledge accumulating systems because they allow the most rapid and efficient adaptive responses to changes in environment. Therefore, it should not be surprising that neuronal computation is not limited to animal brains but is used also by bacteria and plants. The journal, Plant Signaling & Behavior, was launched as a platform for exchanging information and fostering research on plant neurobiology in order to allow our understanding of plants in their whole integrated, communicative, and behavioral complexity.I always go by official statistics because they are very carefully compounded and, even if they are false, we have no others ... approximately Jaroslav Hasek, 1911.

Historical overview on plant neurobiology.

The review tracks the history of electrical long-distance signals from the first recordings of action potentials (APs) in sensitive Dionea and Mimosa plants at the end of the 19(th) century to their re-discovery in common plants in the 1950's, from the first intracellular recordings of APs in giant algal cells to the identification of the ionic mechanisms by voltage-clamp experiments. An important aspect is the comparison of plant and animal signals and the resulting theoretical implications that accompany the field from the first assignment of the term "action potential" to plants to recent discussions of terms like plant neurobiology.