Plant perception (physiology)

The leaf closing after touch in Mimosa pudica depends upon electrical signals.

Vine tendril. Note how the plant reaches for and wraps around the galvanised wire provided for the purpose. This is a very tough twig and appears to have no other purpose than support for the plant. Nothing else grows from it. It must reach out softly, then wrap around and then dry and toughen. See more at thigmotropism.

Plant perception is the ability of plants to sense and respond to the environment to adjust their morphology, physiology, and phenotype accordingly.^[1] Other disciplines such as plant physiology, ecology and molecular biology are used to assess this ability. Plants react to chemicals, gravity, light, moisture, infections, temperature, oxygen and carbon dioxide concentrations, parasite infestation, disease, physical disruption, sound,^[2] and touch.

Processes

Detection

Arabidopsis thaliana can perceive magnetic fields using cryptochromes.^[3] Displacement can also be detected by plants.^[4] Poplar stems can detect reorientation and inclination (equilibrioception).^[5]

Pathway signals

Wounded tomatoes are known to produce the volatile odour methyl-jasmonate as an alarm signal.^[6] Plants in the neighbourhood can then detect the chemical and prepare for the attack by producing chemicals that defend against insects or attract predators.^[6]

Plants systematically use hormonal signalling pathways to coordinate their own development and morphology.

Neurochemicals

Plants produce several proteins found in the animal neuron systems such as acetylcholine esterase, glutamate receptors, GABA receptors, and endocannabinoid signaling components. They also use ATP, NO, and ROS like animals for signaling.^[7]

Electrophysiology

Although plant cells are not neurons, they can be electrically excitable and can display rapid electrical responses (action potentials) to environmental stimuli. These action potentials can influence processes such as actin-based cytoplasmic streaming, plant organ movements, wound responses, respiration, photosynthesis, and flowering.^{[8][9][10][11]} These electrical responses can cause the synthesis of numerous organic molecules, including ones that act as neuroactive substances in other organisms. Thus, plants accomplish behavioural responses in environmental, communicative, and ecological contexts.

Signal response

A plant's concomitant reactive behavior is mediated by phytochromes, kinins, hormones, antibiotic or other chemical release, changes of water and chemical transport, and other means. These responses are generally slow, taking at minimum a number of hours to accomplish, and can best be observed with time-lapse cinematography, but rapid movements can occur as well. Plants respond to volatile signals produced by other plants.^[12][13] Jasmonate levels also increase rapidly in response to mechanical perturbations such as tendril coiling.^[14]

Plants have many strategies to fight off pests. For example, they can produce different toxins (phytoalexins) against invaders or they can induce rapid cell death in invading cells to hinder the pests from spreading out.

Some plants are capable of rapid movement: the mimosa plant (Mimosa pudica) makes its thin leaves point down at the slightest touch and carnivorous plants such as the Venus flytrap snap shut by the touch of insects.

In plants, the mechanism responsible for adaptation is signal transduction.^{[15][16][17][18]}

Adaptive responses include:

• Active foraging for light and nutrients. They do this by changing their architecture, physiology and phenotype.^[19][20][21]
• Leaves and branches are positioned and oriented in response to light source.^[19][22]
• Ability to detect soil volume and adapt growth accordingly independently of nutrient availability.^[23][24][25]
• Adaptively defend against herbivores.

Aspects of perception

Light

The sunflower, a common heliotropic plant which perceives and reacts to sunlight by slow turning movement

Many plant-organs contain photo-sensitive compounds (phototropins, cryptochromes and phytochromes), each reacting very specifically to certain wavelengths of light. These light sensors tell the plant if it is day or night, how long the day is, how much light is available and from where the light comes. Shoots grow towards light and roots usually grow away from light. These responses are called phototropism and skototropism respectively. They are brought about by light sensitive pigments like phototropins and phytochromes and the plant hormone auxin. Many plants exhibit certain phenomena at specific times of the day; for example, certain flowers open only in the mornings. Plants keep track of the time of the day with a molecular clock. This internal clock is set to the solar clock every day using sunlight. The internal clock coupled with the ability to perceive light also allows plants to measure the time of the day and so find the season of the year. This is how many plants know when to flower. (see photoperiodism) The seeds of many plants sprout only after they are exposed to light. This response is carried out by phytochrome signalling. Plants are also able to sense the quality of light and respond appropriately. For example, in low light conditions, plants produce more photosynthetic pigments. If the light is very bright or if the levels of harmful UV increase, plants produce more of their protective pigments that act as sunscreens.^[26]

Plant intelligence

Plants do not have a brain or neuronal network, but reactions within signalling pathways may provide a biochemical basis for learning and memory in addition to computation and problem solving.^[27] Controversially, the brain is used as a metaphor in plant intelligence to provide an integrated view of signalling.^[28]

Plants respond to environmental stimuli by movement and changes in morphology. They communicate while actively competing for resources. In addition, plants accurately compute their circumstances, use sophisticated cost–benefit analysis and take tightly controlled actions to mitigate and control diverse environmental stressors. Plants are also capable of discriminating positive and negative experiences and of "learning" (registering memories) from their past experiences.^[29][30][31] Plants use this information to update their behaviour in order to survive present and future challenges of their environment.

Plant physiology studies the role of signalling, communication, and behaviour to integrate data obtained at the genetic, molecular, biochemical, and cellular levels with the physiology, development, and behaviour of individual organisms, plant ecosystems, and evolution. The neurobiological view sees plants as information-processing organisms with rather complex processes of communication occurring throughout the individual plant organism. It studies how environmental information is gathered, processed, integrated and shared (sensory plant biology) to enable these adaptive and coordinated responses (plant behaviour); and how sensory perceptions and behavioural events are 'remembered' in order to allow predictions of future activities upon the basis of past experiences. Plants, it is claimed by some plant physiologists, are as sophisticated in behaviour as animals but this sophistication has been masked by the time scales of plants' response to stimuli, many orders of magnitude slower than animals'.

It has been argued that although plants are capable of adaptation, it should not be called intelligence, as plant neurobiologists are relying primarily on metaphors and analogies to argue that complex responses in plants can only be produced by intelligence.^[32]"A bacterium can monitor its environment and instigate developmental processes appropriate to the prevailing circumstances, but is that intelligence? Such simple adaptation behaviour might be bacterial intelligence but is clearly not animal intelligence."^[33] However, plant intelligence fits a definition of intelligence proposed by David Stenhouse in a book about evolution and animal intelligence where he described it as "adaptively variable behaviour during the lifetime of the individual".^[34] Critics of the concept have also argued that a plant cannot have goals once it is past the development stage of plantlet because, as a modular organism, each module seeks its own survival goals and the resultant whole organism behavior is not centrally controlled.^[33] This view, however, necessarily accommodates the possibility that a tree is a collection of individually intelligent modules cooperating with, competing with and influencing each other, thus determining organism level behavior from the base up. The development into a larger organism whose modules must deal with different environmental conditions and challenges is not universal across plant species either, as smaller organisms might be subject to the same conditions across their bodies, at least, when the below and above ground parts are considered separately. Moreover, the claim that central control of development is completely absent from plants is readily falsified by apical dominance.

Charles Darwin studied the movement of plants and in 1880 published a book The Power of Movement in Plants. In the book he concludes:

It is hardly an exaggeration to say that the tip of the radicle thus endowed [..] acts like the brain of one of the lower animals; the brain being situated within the anterior end of the body, receiving impressions from the sense-organs, and directing the several movements.

In philosophy, there are few studies of the implications of plant perception. Michael Marder put forth a phenomenology of plant life based on the physiology of plant perception.^[35] Paco Calvo Garzon offers a philosophical take on plant perception based on the cognitive sciences and the computational modeling of consciousness.^[36]

Comparison to neurobiology

A plant's sensory and response system has been compared to the neurobiological processes of animals. Plant neurobiology, an unfamiliar misnomer, concerns mostly the sensory adaptive behaviour of plants and plant electrophysiology. Indian scientist J. C. Bose is credited as the first person to research and talk about neurobiology of plants. Many plant scientists and neuroscientists, however, view this as inaccurate, because plants do not have neurons.^[32]

The ideas behind plant neurobiology were criticised in a 2007 article^[32] published in Trends in Plant Science by Amedeo Alpi and 35 other scientists, including such eminent plant biologists as Gerd Jürgens, Ben Scheres, and Chris Sommerville. The breadth of fields of plant science represented by these researchers reflects the fact that the vast majority of the plant science research community reject plant neurobiology. Their main arguments are that:^[32]

• "Plant neurobiology does not add to our understanding of plant physiology, plant cell biology or signaling".
• "There is no evidence for structures such as neurons, synapses or a brain in plants".
• The common occurrence of plasmodesmata in plants which "poses a problem for signaling from an electrophysiological point of view" since extensive electrical coupling would preclude the need for any cell-to-cell transport of a ‘neurotransmitter-like’ compounds.

The authors call for an end to "superficial analogies and questionable extrapolations" if the concept of "plant neurobiology" is to benefit the research community.^[32]

There were several responses to the criticism clarifying that the term "plant neurobiology" is a metaphor and metaphors have proved useful on several previous occasions.^[37][38] Plant ecophysiology describes this phenomenon.

Parallels in other taxa

As described above in the case of a plant, similar mechanisms exist in a bacterial cell, a choanoflagellate, a fungal hypha, or a sponge, among the many other examples. All of these individual organisms of the respective taxa, despite being devoid of a brain or nervous system, are capable of sensing their immediate and momentary environment and responding accordingly. In the case of single-celled life, the sensory pathways are even more primitive in the sense that they take place on the surface of a single cell, as opposed to a network of many cells.

See also

References

Further reading

• Baluška F, Mancuso S (ed) (2009). Signalling in Plants. Springer Verlag.
• Baluška F (ed) (2009). Plant-Environment Interactions: From Sensory Plant Biology to Active Plant Behavior. Springer Verlag.
• Brenner E, Stahlberg R, Mancuso S, Vivanco J, Baluška F, Van Volkenburgh E (2006). Plant Neurobiology: An Integrated View of Plant Signaling. Trends Plant Sci 11: 413-419.
• Cvrckova, F et al. (2009). Plant Intelligence: Why, Why Not, or Where? Plant Signal Behav 4: 394-399.
• Garzón FC (2007). The Quest for Cognition in Plant Neurobiology. Plant Signal Behav 2: 208-211.
• Gilroy S, Masson PH (2007). Plant Tropisms. Iowa State University Press.
• Karban R (2008). Plant Behaviour and Communication. Ecol Lett 11: 727-739.
• Karban R (2015). Plant Sensing and Communication. University of Chicago Press.
• Mancuso S, Shabala S (2006). Rhythms in Plants. Springer Verlag.
• Scott P (2008). Physiology and Behaviour of Plants. John Willey & Sons Ltd.
• Trewavas A (2003). Aspects of Plant Intelligence Ann Bot. 92: 1-20
• Trewavas A (2005). Plant Intelligence. Naturwissenschaften 92: 401-413.
• Trewavas A (2009). What is Plant Behaviour? Plant Cell Environm 32: 606-616.
• Volkov AG (2006). Plant Electrophysiology. Springer Verlag.
• Volkov AG (2008). Plant Electrical Memory. Plant Signal Behav 3: 490-492.

External links

• Miller, Deborah; Whitney Hable; Jennifer Gottwald; Mary Ellard-Ivey; Taku Demura; Terri Lomax; Nick Carpita (1997). Connections: The Hard Wiring of the Plant Cell for Perception, Signaling, and Response. The Plant Cell 9(12). pp. 2105–2117. Retrieved 2006-12-25. 

• Keen, Noel T; Shigeyuki Mayama, Jan E. Leach, and Shinji Tsujumu (eds) (2001). Delivery and Perception of Pathogen Signals in Plants. APS Press. p. 268. ISBN 0-89054-259-7.  Cite uses deprecated parameter |coauthors= (help)

• Taiz, Lincoln; Eduardo Zeiger (2006). Plant Physiology, fourth edition. Sinauer Associates. p. 700 (est). ISBN 0-87893-856-7. 

• Taiz, Lincoln; Eduardo Zeiger (2002). "Plant Physiology Online". a companion to Plant Physiology, Third Edition. Sinauer Associates. Archived from the original on 7 December 2006. Retrieved 2006-12-26. 

• Dierk Scheel and Claus Wastermack (May 2002). Plant Signal Transduction. Oxford University Press. p. 346. ISBN 978-0-19-963879-6. Retrieved 2006-12-25. 
• Plant Neurobiology Society
• Plant Signaling and Behavior Scientific journal for Plant Neurobiology
• Study hints plants have sensibilities
• Plants cannot "think and remember," but there's nothing stupid about them: They're shockingly sophisticated
• How Does a Venus Flytrap Work?

Physiology types Animals Fish physiology Human physiology Insect physiology Physiology of dinosaurs Plants Plant physiology Plant perception (physiology) Physiological plant disorders Cells Cell physiology Related topics Comparative physiology Electrophysiology Evolutionary physiology Molecular physiology