I would like to express my profound sense of gratitude to my guide Dr. S. Kiran for his kind and able guidance. I am grateful for his help in preparation of this manuscript. I would like to express my profound sense of gratitude to Principal Dr. Netaji S. Ganesan, Director of life sciences Dr. P. S. Rao and Head of the Department Dr. G. S. JagannathaRao for extending their support. I thank all the Teaching and Technical staff of the Department of Biotechnology Engineering who have helped me on various occasions during the course of Engineering. My sincere thanks to all my Friends who have helped me on various occasions during the course of Engineering. I am deeply indebted to my Parents and Siblings who have encouraged and supported me throughout the course of Engineering.

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Acknowledgement01 Abstract 1. Introduction04
1.1. Definition 1.2. Senses 1.3. Non-human senses


2.Plant senses
2.1. Plants and vision 2.2. Plants sense of touch 2.3. Plants can taste 2.4. Plant hearing 2.5. Plant can smell


3. Stomatal action
3.1. Definition 3.2. Mechanism of stomatal action 3.3. Active potassium theory 3.4. Split root experiment


4. Some special sense of plants 5. Case studies 27 6. Conclusion 7. References 31 8. Annexure 9. Articles 49 9.1.Up, down, and all around: How plants sense and respond to environmental Stimuli.
9.2. Mechanoreceptor Cells on the Tertiary Pulvini of Mimosa pudica .




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Plants are bursting with movement. They are rich in sensation, and respond to the stimulation of the Surrounding world every moment of their active lives. They can send messages to one another about overcrowding or a threatened attack by a new pest. Within each plant there is ceaseless activity as purposive as that in an animal. Many of them share hormones that are remarkably similar to our own. Their senses are sophisticated: some can detect the lightest touch (better than the sensitivity of the human fingertips), and they all have a sense of vision. Plants are intelligent forms of life who are capable of intention, preference, and a will to survive, thrive and interact. Scientific research indicates that plants communicate with insects, animals, human beings and other plants in order to keep themselves alive and safe. Evidence also reveals that plants are telling us how to achieve health and wholeness for humanity and the earth herself. Plants may not have complex tissues and a nervous system, but they still "feel". If they sprout, develop and flower at the right time and place, that's because they are sensitive to environmental factors. The vegetationboomduring each spring shows they obey precise rhythms, which come encoded in their genes. Plants do not have only a touch sensitivity (that has been known for quite some time), but also a chemical one, not to mention their sensitivity to light and temperature variations. This way, plants can appreciate the length of the day and the air temperature, adopting aposition fitting their neighborhood. Awound, stress or a disease trigger specific defense mechanisms. Information about their state and environment circulates along signals transmitted from one cell to another, from plant to plant, or even from one plant to other beings. Their sensitivity translates via movements, growth directions and metabolism changes.

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INTRODUCTION Department of Biotechnology Feb11 -Mar11 Page 4 .PLANTS HAVE SENSE! 1.

Non-human animals also have receptors to sense the world around them. 1. other senses include temperature (thermoception). cognitive psychology (or cognitive science). and philosophy of perception. Some species of animals are able to sense the world in a way that humans cannot. touch. A broadly acceptable definition of a sense would be "A system that consists of a group of sensory cell types that responds to a specific physical phenomenon. and detect water pressure and currents. Humans are considered to have at least five additional senses that include: nociception (pain). hearing (audioception).however. photoreception. such as electroreception and detection of polarized light. Non-human animals may possess senses that are absent in humans.1.PLANTS HAVE SENSE! 1. dedicated to each sense. Humans have a comparatively weak sense of smell. with degrees of capability varying greatly between species. leading to difficulties in defining what exactly a sense is. classification. smell and taste. and six more if interoceptive senses (see other internal senses below) are also considered. This categorisation has been criticized as too restrictive. kinesthetic sense (proprioception). pain (nociception). and possibly an additional weak magnetoception (direction). whilst other species may lack one or more of the traditional five senses. proprioception and kinaesthesia (joint motion and acceleration). Human beings have a multitude of senses.000. hearing. mechanoreception.000 olfactory receptors. and touch (tactioception). balance (equilibrioception) and acce leration (kinesthesioception). Definition There is no firm agreement among neurologists as to the number of senses because of differing definitions of what constitutes a sense. thermoception (temperature differences). and that corresponds to a particular group of regions within the brain where the signals are received and interpreted. In addition to the traditionally recognized five senses of sight (ophthalmoception). The senses and their operation. taste (gustaoception). a classification attributed to Aristotle. most notably neuroscience." Disputes about the number of senses typically arise around the classification of the various cell types and their mapping to regions of the brain. The human olfactory system contains about 10. One commonly recognized categorisation for human senses is as follows: chemorecenptio. Department of Biotechnology Feb11 -Mar11 Page 5 . INTRODUCTION: Senses are the physiological capacities within organisms that provide inputs for perception. and theory are overlapping topics studied by a variety of fields. as it does not include categories for accepted senses such as the sense of time and sense of pain. One definition states that an exteroceptive sense is a faculty by which outside stimuli are perceived. equilibrioception (balance). with some species able to sense electrical and magnetic fields. The traditional five senses are sight. smell (olfacoception or olfacception). The nervous system has a specific sensory system or organ. and thermoception. What constitutes a sense is a matter of some debate. Other species may also intake and interpret senses in very different ways. sense of time.

Hearing is all about vibration. and brightness.2. hues. Mechanorecptors turn motion into electrical nerve pulses which are located in the inner ear. 1.3. Cones distinguish colors. salty. The inability to hear is called deafness. Rods are very sensitive to light. There are at least four types of tastes that "buds" (receptors) on the tongue detect.2. There are two types of photoreceptors: rods and cones. given that each receptor conveys information to a slightly different region of the brain. damage to the optic nerve that connects each eye to the brain. The inability to see is called blindness.2. was first theorised in 1908 and its existence confirmed in 2000. These smooth muscles control airway contraction and dilation . The umami receptor detects the amino acid glutamate. the detection of these vibrations. that is the sense of the hearing. Sight: Sight orvision is the ability of the eye(s) to focus and detect images of visible light on photoreceptors in the retina of each eye that generates electrical nerve impulses for varying colors.2.2. Sound can also be detected as vibrations conducted through the body by tactition. Some argue that stereopsis. also constitutes a sense. mild chilli and olive oil is perceived as a peppery tickle at the back of the throat (which senses polyphenols in unprocessed olive oil). bitter substances such as quinine or chloroquine opened contracted airways. Another taste sense for ginger. a neurotransmitter commonly found in meat and in artificial flavourings containing monosodium glutamate. for a sensation called umami. Temporary or permanent blindness can be caused by poisons or medications. Blindness may result from damage to the eyeball. sour. There is some disagreement as to whether this constitutes one. 1. Hearing at high frequencies declines with an increase in age. given that different receptors are responsible for the perception of color and brightness. and hence there are anatomists who argue that these constitute five or more different senses. although the receptors for sweet and bitter have not been conclusively identified. Department of Biotechnology Feb11 -Mar11 Page 6 . Hearing: Hearingorauditionis the sense of sound perception. The four well-known receptors detect sweet.contrary to expectation. Lower frequencies than that can be heard are detected this way. and/or from stroke (infarcts in the brain). but are less sensitive to dim light. especially to the retina. Since sound is vibrations propagating through a medium such as air. but it is generally regarded as a cognitive (that is.Senses 1. but do not distinguish colors. Neuroanatomists generally regard it as two senses. offering new insight into asthma. A fifth receptor. two or three senses.PLANTS HAVE SENSE! 1.1. the perception of depth using both eyes. postsensory) function of the visual cortex of the brain where patterns and objects in images are recognized and interpreted based on previously learned information.Taste: Taste or gustation is one of the two main "chemical" senses. and bitter. with substantial variation between individuals. is a mechanical sense because these vibrations are mechanically conducted from the eardrum through a series of tiny bones to hair-like fibers in the inner ear which detect mechanical motion of the fibers within a range of about 20 to 20. The inability to taste is called ageusia.000 hertz. The University of Maryland School of Medicine announced in 2010 that bitter taste receptors had been discovered on smooth muscle in human lung bronchi.

This combination of excitatory signals from different receptors makes up what we perceive as the molecule's smell. the heat flux (the rate of heat flow) in these areas. Paresthesia is a sensation of tingling.2. Unlike taste. 1. Odor molecules possess a variety of features and thus excite specific receptors more or less strongly.6. 1.4. orvestibular sense is the sense which allows an organism to sense body movement. brushing. In the brain. The vestibular nerve conducts information from sensory receptors in three ampulla that sense motion of fluid in three semicircular canals caused by three-dimensional rotation of the head. The inability to smell is called anosmia.2. or numbness of the skin that may result from nerve damage and may be permanent or temporary. The organ of equilibrioception is the vestibular labyrinthine system found in both of the inner ears. pricking. or rather.PLANTS HAVE SENSE! Note: that taste is not the same as flavour. The loss or impairment of the ability to feel anything touched is called tactile anesthesia. Olfactory receptor neurons in the nose differ from most other neurons in that they die and regenerate on a regular basis. telling wind direction. A variety of pressure receptors respond to variations in pressure (firm. Department of Biotechnology Feb11 -Mar11 Page 7 . There are specialized receptors for cold (declining temperature) and to heat. sustained. olfaction is processed by the olfactory system.2. linear acceleration. The touch sense of itching caused by insect bites or allergies involves special itch-specific neurons in the skin and spinal cord. etc. and to attain and maintain postural equilibrium and balance. direction. throat.5. but also in the tongue.7. but they are known together as equilibrioception.Touch: Touchalso called tactition or mechanoreceptionis a perception resulting from activation of neural receptors.). Some neurons in the nose are specialized to detect pheromones.2. and mucosa. The thermoceptors in the skin are quite different from the homeostaticthermoceptors in the brain (hypothalamus) which provide feedback on internal body temperature. Technically this organ is responsible for two senses of angular momentum acceleration and linear acceleration (which also senses gravity). there are hundreds of olfactory receptors. flavour includes the smell of a food as well as its taste. each binding to a particular molecular feature. generally in the skin including hair follicles.Smell: Smell or olfaction is the other "chemical" sense.Temperature: Thermoception is the sense of heat and the absence of heat (cold) by the skin and including internal skin passages.Balance and acceleration: Balance. 1. The vestibular nerve also conducts information from the utricle and the saccule which contain hair-like sensory receptors that bend under the weight of otoliths (which are small crystals of calcium carbonate) that provide the inertia needed to detect head rotation. The heat receptors are sensitive to infrared radiation and can occur in specialized organs for instance in pit vipers. and the direction of gravitational force. The cold receptors play an important part in the dog's sense of smell. equilibrioception. 1. and acceleration.

Without pain. For example. the kinesthetic sense provides the parietal cortex of the brain with information on the relative positions of the parts of the body. pressure. Assuming proper proprioceptive function. The chemoreceptor trigger zone is an area of the medulla in the brain that receives inputs from blood-borne drugs or hormones. for example headache caused by vasodilation of brain arteries. and communicates with the vomiting center. The main function of pain is to warn us about dangers. such as stretch receptors that are neurologically linked to the brain. These involve numerous sensory receptors in internal organs. 1. and temperature. including touch. The three types of pain receptors are cutaneous (skin). and their impairment results in surprising and deep deficits in perception and action. 1. Stimulation of stretch sensors that sense dilation of various blood vessels may result in pain.8. at no time will the person lose awareness of where the hand actually is. Department of Biotechnology Feb11 -Mar11 Page 8 . Sensory receptors in pharynx mucosa. Proprioception and touch are related in subtle ways. or during acid reflux. y y y y y y y y Pulmonary stretch receptors are found in the lungs and control the respiratory rate. but recent studies show that pain is registered in the anterior cingulate gyrus of the brain. and thus is dangerous.Pain: Nociception (physiological pain) signals nerve-damage or damage to tissue.9. Stimulation of sensory receptors in the urinary bladder and rectum may result in sensations of fullness. Stimulation of sensory receptors in the esophagus result in sensations felt in the throat when swallowing.2. Cutaneous receptors in the skin not only respond to touch. even though it is not being detected by any of the other senses. vomiting. sense foreign objects such as food that may result in a gag reflex and corresponding gagging sensation. but also respond to vasodilation in the skin such as blushing.Kinesthetic sense: Proprioception. Other internal senses: An internal sense or interoception is "any sense that is normally stimulated from within the body". humans avoid touching a sharp needle or hot object or extending an arm beyond a safe limit because it hurts.10. similar to touch receptors in the skin. people could do many dangerous things without realizing it. It was previously believed that pain was simply the overloading of pressure receptors. Pain was once considered an entirely subjective experience. but research in the first half of the 20th century indicated that pain is a distinct phenomenon that intertwines with all of the other senses.2. Stretch receptors in the gastrointestinal tract sense gas distension that may result in colic pain. Neurologists test this sense by telling patients to close their eyes and touch their own nose with the tip of a finger. somatic (joints and bones) and visceral (body organs).2.PLANTS HAVE SENSE! 1.

1. Reptiles like snakes and monitor lizards make extensive use of it as a smelling organ by transferring scent molecules to the vomeronasal organ with the tips of the forked tongue. 1. Magnetotactic bacteria build miniature magnets inside themselves and use them to determine their orientation relative to the Earth's magnetic field. Non-human senses: 1. trails. Although there is no dispute that this sense exists in many avians (it is essential to the navigational abilities of migratory birds). and sexual state.3. a phenomenon which is known as human echolocation. including chemicals in the water.1. and can taste anything they touch. the mechanisms and capabilities vary widely. The organ is vestigial in humans. 1.2.3. including bats and cetaceans. One study has found that cattle make use of magnetoception. 1. In mammals it is often associated with a special behavior called flehmen characterized by uplifting of the lips. Department of Biotechnology Feb11 -Mar11 Page 9 . their own footsteps).3. as they tend to align themselves in a north-south direction.3. because associated neurons have not been found that give any sensory input in humans.1. Echolocation: Certain animals. 1. They most often use this to navigate through poor lighting conditions or to identify and track prey. Taste: Flies and butterflies have taste organs on their feet.1. Insects have olfactory receptors on their antennae. although the mechanism is similar. It has also been observed in insects such as bees. it is not a well-understood phenomenon. allowing them to taste anything they land on. including many of the senses listed above for humans. However.3.PLANTS HAVE SENSE! 1. They follow the nostril that first detected the smell.3. reptiles.1. Smell: Most non-human mammals have a much keener sense of smell than humans. Direction: Magnetoception (or magnetoreception) is the ability to detect the direction one is facing based on the Earth's magnetic field.4.1. mammals) have a vomeronasal organ that is connected with the mouth cavity. Sharks combine their keen sense of smell with timing to determine the direction of a smell. a task that has proven difficult in practice. Vomeronasal organ: Many animals (salamanders.3. Blind people report they are able to navigate by interpreting reflected sounds (esp. have the ability to determine orientation to other objects through interpretation of reflected sound (like sonar).5.1. There is currently an uncertainty whether this is simply an extremely developed post-sensory interpretation of auditory perceptions or it actually constitutes a separate sense.1. Catfish have taste organs across their entire bodies. In mammals it is mainly used to detect pheromones to mark their territory. Analogous to human senses: Other living organisms have receptors to sense the world around them. Directional awareness is most commonly observed in birds.3. Resolution of the issue will require brain scans of animals while they actually perform echolocation.

the platypus has the most acute sense of electroception. a system consisting of three appendages of vertebrae transferring changes in shape of the gas bladder to the middle ear.3. Fish like the weather fish and other loaches are also known to respond to low pressure areas but they lack a swim bladder.1. This is however not electroception as it is a post-sensory cognitive action. 1. 1. mostly consisting of vortices. This sense plays a role in the navigational abilities of several animal species and has been postulated as a method for animals to develop regional maps.3. pythons and some boas have organs that allow them to detect infrared light. Among these mammals. the same mechanoreceptors for vestibular sense and hearing. hunting. Several species of fish. y y Department of Biotechnology Feb11 -Mar11 Page 10 . some generate their own weak electric fields. Some fish passively sense changing nearby electric fields. An electrically charged balloon. y Magnetoreception (or magnetoception) is the ability to detect a magnetic field to perceive direction. a reflective membrane that optimizes the image.3. Pressure detection uses the organ of Weber. Vision: Cats have the ability to see in low light due to muscles surrounding their irises to contract and expand pupils as well as the tapetumlucidum. and some use these electric field generating and sensing capacities for social communication.2. altitude or location. however in general humans (and probably other mammals) can detect electric fields only indirectly by detecting the effect they have on hairs.PLANTS HAVE SENSE! 1. Body modification enthusiasts have experimented with magnetic implants to attempt to replicate this sense. It is used primarily for navigation. will exert a force on human arm hairs. including the following: y Electroreception (or electroception) is the ability to detect electric fields. It has been found that birds and some other animals are tetrachromats and have the ability to see in the ultraviolet down to 300 nanometers. Bees and dragonflies] are also able to see in the ultraviolet. Balance: Ctenophora have a balance receptor (a statocyst) that works very differently from the mammalian's semi-circular canals. Not analogous to human senses: In addition. It can be used to regulate the buoyancy of the fish. Pitvipers. such that these snakes are able to sense the body heat of their prey. and sense the pattern of field potentials over their body surface. The lateral line is also sensitive to low frequency vibrations. which can be felt through tactition and identified as coming from a static charge (and not from wind or the like). The common vampire bat may also have an infrared sensor on its nose. The receptors of the electrical sense are modified hair cells of the lateral line system. Current detectionThelateral line in fish and aquatic forms of amphibians is a detection system of water currents. The mechanisms by which electroceptive fish construct a spatial representation from very small differences in field potentials involve comparisons of spike latencies from different parts of the fish's body. sharks and rays have the capacity to sense changes in electric fields in their immediate vicinity. The mechanoreceptors are hair cells.7. and schooling. for instance.1. The only order of mammals that is known to demonstrate electroception is the monotreme order. some animals have senses that humans do not.6.

water. Most sighted humans can in fact learn to roughly detect large areas of polarization by an effect called Haidinger's brush. light. that respond to vibration. or other specific chemicals.PLANTS HAVE SENSE! Polarized light direction/detection is used by bees to orient themselves. Cuttlefish can also perceive the polarization of light. however this is considered an entoptic phenomenon rather than a separate sense. Department of Biotechnology Feb11 -Mar11 Page 11 . providing information on force and vibrations. especially on cloudy days. scents. Slit sensillae of spiders detect mechanical strain in the exoskeleton. Some plants sense the location of other plants and attack and eat part of them. y y Some plants have sensory organs. for example the Venus fly trap.

PLANT SENSES Department of Biotechnology Feb11 -Mar11 Page 12 .PLANTS HAVE SENSE! 2.

plants orientate in space and time. Riboflavin may have connections with human life. The disturbance of the soil brought to the surface many seeds which were stimulated to germination. riboflavin (but not carotene) is inactivated. It was originally known as vitamin B2 and is essential for health. which was proposed as a light receptor as a result of some simple experiments. when we go to quickly slap a fly. the action is in slow motion. there is a relative excess of dark red. This explains the poppy fields of the battlegrounds of the First World War. can no lon be ger sustained. That is because we are also living with a limited Perception of life. the churned fields were cloaked in blood-red poppies. The inhibiting process stops and the stems grow longer. the eye can become reddened. Such interesting coincidences between human sight and the photoreceptors of the plant kingdom remind us of the universality of the senses. 2. The notion that perception is confined to humans. PLANT SENSES: It is not easy to describe the senses that plants have. the plant fails to respond to light. The photoreceptors of the plants are sensitive not only to the amount of light they receive. Within weeks of terrifying ground attacks. People deficient in riboflavin find bright light unbearable. but also to its quality. and a sense of itching or soreness develops. How plants follow light? The sense of light detection exists even in seeds.Under a dense plant cover. plants possess an excellent internal communication system from roots to leaf tip. Cataracts can develop. Although there has been much research into this potentially revealing area. a process called " the syndrome of shadow avoiding". but the light red stimulates seed germination and chlorophyll synthesis.1. enabling plants to sense the presence of their green neighbours. Phytochrome.1. Department of Biotechnology Feb11 -Mar11 Page 13 . and this photophobia can be so severe that they cannot endure even normal daylight. or even to the animal kingdom. it is hard to tell the relationship between a light sensitive pigment and the way a plant responds. inhibiting stem length growth. others to blue or ultraviolet.1.PLANTS HAVE SENSE! 2. A plant seems to have an even slower outlook on its environment. the balance moves towards the inactive form of the phytochrome. They also distinguish if the light contains more light red or dark red.Plants and vision: There are light-sensitive chemicals in plants which do not take part in photosynthesis. For example. for instance. but give the plant its sense of vision. for the fly. This is where molecular biology could hold the key to unravelling the mechanisms hidden in the green plant's sense of sight. An insect with compound eye structure has a different look on life and time.Some photoreceptors pigments are more sensitive to red. can sense the relationship between red and far-red light. In that state. It is already difficult enough to imagine why a snake can see in the dark by means of infrared sight. Both types of light are present in the daylight. In this case. which strongly supports the idea that riboflavin acts as a sensory compound. Depending on light. If plant tissues are treated with potassium iodide. Another example is phytochrome. 2. It is also known that plants can receive signals from their environment and that they can also signal back. Another is riboflavin. A lack of riboflavin in the hum diet leads to an problems with the eye. Many plant seeds require sunlight to germinate.

in 1875. 2. over 300 protocarnivorous plant species in several genera show some but not all these characteristics. a simple rod of plant tissue. so it is natural that a plant should be able to detect light well enough to ensure that it derives maximum benefit from the sun.2. They try to resist wind by strengthening tissues that are being swayed. However.But a sense of touch is something every plant has. produce digestive enzymes. and absorb the resulting available nutrients. and around the far edge are Department of Biotechnology Feb11 -Mar11 Page 14 . The extra energy expended stif ening tissue can cost f farmers dear. the facts stand and more research will doubtless unearth the real answer to plants and their reactions to light. yields drop by 30 to 40% compared with unshaken plants. One experiment showed that when maize plants are shaken for 30 seconds each day. True carnivory is thought to have evolved independently six times in five different orders of flowering plants. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients. which can cause damage to foliage. An old farming tale suggests that fields ploughed at night grow fewer weeds. Charles Darwin wrote Insectivorous Plants. 2. The plant-growth hormones. as µCoughlin¶ describes: "Ordinary plants need a sense of touch to respond to the buffeting of the wind. a shoot will always grow towards the light. and these are now represented by more than a dozen genera. the growth will continue in a straight line. Plant¶s sense of touch: 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.PLANTS HAVE SENSE! It was the sunlight which made the poppies grow. Light is the green plant's primary source of energy. This does not make immediate sense to me . Carnivorous plants: Carnivorous plants are plants that derive some or most of their nutrients (but not energy) from trapping and consuming animals or protozoans. Some of the mechanisms of sense are simple.even if the seeds were exposed at night. These include about 630 species that attract and trap prey. stimulate cells to grow. If light shines onto the left of the shoot. Additionally. On the glandular inner surface of each lobe are three fine spiny hairs. for it has jaws which snap shut in one-third of a second. Imagine an upward-growing shoot. Some recent research suggests that levels of weed growth can be cut by 80 per cent by ploughing at night.1. however. The increase in size of the tissues on the darker (right) side will therefore tend to bend the shoot toward the light. the first wellknown treatise on carnivorous plants. Plant growth hormones in stems are more concentrated in dark tissues than in those exposed to light. Each trap is composed of a leaf divided into two oval portions. typically insects and other arthropods. The effect of light on growth hormone seems to 'drive it away'. In this way. they would surely be stimulated by sunlight the next day.2. Venus flytrap: (Dionaeamuscipula) The Venus Fly Trap seems to be the closest to an animal. then this lighter side will continue to grow rather slower than before. such as acidic bogs and rock outcroppings. and teeth which hold the prey. which open like a butterfly's wings. Once illumination is evenly detected on all sides. especially nitrogen. auxins.

The electrical impulse was recorded immediately after the sensitive hairs were touched. He had two main reasons for drawing thisconclusion. This scepticism resulted in a lack of further interest in the study of plant movement and the subject lay dormant for a century. thespeed of the impulse was too slow.PLANTS HAVE SENSE! interlocking teeth like those of a man-trap. A slight movement against one of the hairs triggers the trap into action. US scientists started to study how impulses are transmitted by cells. The cell is covered with a thin skin. and soon found an electrical signal in 1873. it was believed that they collapsed mechanically inwards and allowed the leaves to close. who concluded that there was no true nervous activity in the fly-trap. the cell membrane. Just because we measure a peak of electrical activity does not necessarily mean that there is a nervous reaction. the two halves spring towards each other so fast that they prey cannot escape. Do plants have nervous system ? Sir John Burdon-Sanderson (1828 . If the chemical change were produced by the stimulation of one of the trigger hairs on a fly-trap. and similarities between plants and animals started to emerge.sodium chloride. and before any movement was detected. and closure of the trap. (1832 . But in the 1960s.1905) hitched the leaves up to the apparatus that he was using to study how nerves send signals to muscles. Surely. von Sachs argued if there were nerve impulses. This objection was raised. And second. was only 20 cm (8 in) per second. there are no nerve cells in the fly-trap. we would expect it to be detectable as an electrical response.97) an eminent German botanist. and one of the fundamental mechanisms of any living cell is a way of controlling the level of salt . which is a fatty layer insulating the inside of the cell Department of Biotechnology Feb11 -Mar11 Page 15 . they must have nerves along which the impulses travel. Keeping the sodium ions from saltwater atbay is done by restricting their movement in and out of a living cell. This is very similar to the way an animal responds to a stimulus. electrical impulse. The earliest research into the mechanism behind the movement of the leaves of sensitive plant showed that contact produced a collapse in the cells that ordinarily hold the leaves erect. First. at the time by Julius von Sachs. while the rate in the Venus fly-trap. Impulses can travel along animal nerves at thousands ofcentimetres per second. Life on earth first developed in the salt water of the seas. we could merely be measuring the results of a chemicalchange inside the cells. As water passed rapidly out of these supporting cells. Here was a series of three events: stimulation of the hair.

The sequence becomes complex: 1. the trap remains open. if slight.This stimulus is translated into an action potential only if it is strong enough. If an impulse passes along a nerve in an animal. This means that a tiny fly will not ordinarily be trapped. a plant treated with care and affection gives out a different vibration compared to a plant subjected to torture. If the action potential is insufficient.when the stimulus itself is detected. 4. the same mechanism has been detected in them. the trap stays open. But what happens with medium strength stimuli? If a stimulus is below the threshold. an advancing wave of gates opens to allow sodium ions into the nerve and potassium ions out. 2. The receptor potential results from the hair-cells in the fly-trap being touched. Movement of a hair-cell. So a small stimulus producing a modest receptor. The timing of the responses. 3. who began to conduct experiments on plants in the year 1900. which produces only the slightest touch. the trap will snap shut. which closes the trap. He found that every plant and every part of a plant appeared to have a sensitive nervous system and responded to shock by a spasm just as an animal muscle does. One of the first to research the concept was the Indian scientist Sir Jagdish Chandra Bose. and these gates are used by the cell to regulate the passage of ions through the membrane. potential. the trap remains open. shows that they are based on mechanisms much like those in animals. There are several stages of information processing in the Venus fly-trap. may be ignored. he said: "Do not these records tell us of some property of Department of Biotechnology Feb11 -Mar11 Page 16 .PLANTS HAVE SENSE! from the outside environment. like animal nerves. The most interesting discovery of all was that plant cells. the hair-cell will still generate an action potential. Because of the ions inside the cell there is a negative electrical charge (about one-tenth of a volt). if it is strong enough. if sufficient. may be insufficient to cause the action potential to be triggered. According to him. Now that we can study plant cells. If a further stimulation within half a minute is enough to trigger a potential. In conclusion. an action potential will fire and an electrical charge will pass across the open trap. but a slowly moving insect. The central layer of fat in the cell membrane acts as an electrical insulator. the trap will close. will be trapped if it causes gentle movements of a hair cell. A later stimulation of a hair-cell may be too weak to generate a response. it is ignored. together with their nature. It is as if the cell is remembering the stimuli and adding them up. can manifest a receptor potential before the action potential itself. A strong: stimulus will create a receptor potential sufficient to generate an action potential. As a rule there needs to be a second stimulation within half a minute for the two to be associated. it causes a receptor potential to fire. in which case. but if several more small stimuli are received. Ions can pass this barrier only if tiny apertures open to allow them through. The action potential is the electrical signal that induces a response in a living cell. if it is sufficiently strong. the receptor potential is the signal created before that . which suggests that there is a kind of memory within the plant-cell. If this is small.

The message had been transported by ethylene. The pine invaded by insects digging galleries in its bark responds through an increased release of resin. plants also emit SOS signals.000 plant secondary metabolites. One chemist made an experiment: he investigated how willows react when attacked by a caterpillar. Another potential message may be the acetylsalicylic acid (or aspirin). The phenomenon was first signaledin the bean attacked by mites. which are in fact the larvae of a beetle.The presence of the Spodoptera caterpillars on the corn leaves triggers the release of a chemical cocktail that attracts the parasitoid wasps of these larvae. he picked up leaves from the two plots to feed other caterpillars. the larches produce smaller and less nutritious leaves. with toxic or rejecting actions against the plant-eating insects are known. When in danger. killing the attackedcells. peptides and terpens. when attacked by the caterpillars of the moth Heliothisvirescens. tannins. over 10. stored mostly in leaves. extracted from the willow bark ("salix" means "willow" in Latin). up to 50 % of the compounds synthesized by a tree can be used as defense products. like alkaloids. After 14 days. Plants can taste(Plantsability to sense chemicals): After "tasting" the chemical betraying the presence of an aggressor. their most vulnerable organ. but a uniform and continuous march of law?" 2. emitting chemicals that attract minute nematode worms that kill the root worms. The beech attacked by the aphid Phyllaphisfagi synthesizes chemicals that inhibit the digestion of the insects. a gas normally produced during the reaping of the fruits but also released by harmed or irritated tissues. predators of the first. stimulating prolonged defense reactions. When informed about an occurring attack. Poplars react similarly to willows to insect attack. Department of Biotechnology Feb11 -Mar11 Page 17 . At the moment. lignin and other defensive chemicals. Zeirapheradiniana.3. a fact that diminishes the number of predators for 4-5 years. Some corn varieties defend themselves against the root worm (Diabroticavirgifera). The plant emitsan array of chemicals that attract other mites. The emitted chemical signals often spread far away from the attacked point. The tobacco chemically attracts the waspCardiochillesnigriceps. When defoliated bythe caterpillars of moth.In other trees.PLANTS HAVE SENSE! matter common and persistent? That there is no abrupt break. The most sensitive plants create a strong barrier between the aggressor (virus. But what was more puzzling is that leavesfrom bothplots were equally "unpleasant" for the caterpillars.He found that the larvae that ate small amounts of leaves were growing very slowly. a plant is able to unleash a chemical war against it. bacterium or fungus) and the backward intact cells. so he assigned them to two lots: one which he invaded with caterpillars and another as a witness. but it remains inert to the presence of the caterpillars of Helicoverpazea (that attacks the corn). leaving it "untouched". What had happened? Both willow groups filled their leaves with a chemical that proved to be repulsive for the insects. Just one molecule from the saliva of the caterpillar is enough to cause the SOS signal. this time in the lab. The oak opposes to the growth of the mistletoes by producing increased amounts of toxic tannins. the reception of the message induces the synthesis of the tannins.

It blankets and kills its host. but also in order to time their blooming. What about the hearing of the plants? Many plant lovers are convinced the sensitivity of theirindoor plants to music is real. Researches made so far did not confirm or infirm the auditory skills of the plants. As dodder seedlings have only a limited amount of food intheir seeds. blooming at the same time can also be a defense mechanism. cranberries. A 2006 research showed dodder can sense chemicalsreleased by host plants and then head for that direction. the dodders could distinguish between a favorite host plant. selecting only the preferred ones! The dodder (Cuscuta) attacks tomatoes. When Jaffe added chemicals to the pea plants inhibiting the biosynthesis of this hormone. Department of Biotechnology Feb11 -Mar11 Page 18 . as the consumers will not have enough time to eat too many flowers. Some farmers believe that sounds louder than the human voice stimulate the germination and rapid growth of some vegetables. Scientists thought the dodder simply grew at random.4. as it would happen if trees bloomed one after another. alfalfa and even flowers.PLANTS HAVE SENSE! However. 2.Mordecai Jaffe (Wake Forest University) used an instrument that made a loud "warble" and got a doubling in the growth of dwarf pea plants. When they tried to trick the dodder seedlings with artificial tomato plants. he was unable to reproduce the original effects. theymus find a host plant quickly or else they die. Scientists found in a lab experiment that 80 % of the dodders grew towards a tomato plant. such as the tomato. it begins growing in a coil around the host plant. carrots. When placed in the middle of wheat and tomato plants. trees were found to communicate not only for defense.Jaffe suspects that the plant hormone gibberellic acid. onions. injecting needles into stems and leaves to suck out water and nutrients. it turned out didn't take the bait: 77 % grew towards the real plants. citrus trees. the dodders swayed aboveground in circular movements in the direction of the tomato plants.5. with the discovery of a plant toattack being just a chance encounter. Plant can smell: Wounded tomatoes are known to produce the volatile odourmethyl-jasmonate as an alarm-signal Plants in the neighbourhoodcan then smell the danger and prepare for the attack by producing chemicals that defend insects or attract predators. When the shoot finds a victim. Scientists suspect the dodder is equipped with sense receptors that can smell their hosts. and a poor one such as wheat. which is instrumental in shoot elongation and seed germination. In fact. Plant hearing: Mechanical perturbation can also be detected by plants. Parasite plants wereeven found to sniff their host-plants. Moreover. It caneven identify favorite plant hosts over less desirable ones. isinvolved in the "hearing" response. 2.

PLANTS HAVE SENSE! 3. STOMATAL ACTION Department of Biotechnology Feb11 -Mar11 Page 19 .

It is bordered by a pair of specialized parenchyma cells known as guard cells which are responsible for regulating the size of the opening.PLANTS HAVE SENSE! 3. When the guard cells are turgid.2.1. Mechanism of stomatal action: Stomatal Movement in Monocot Plants Stomatal Movement in Dicot Plants Department of Biotechnology Feb11 -Mar11 Page 20 . Definition of Stomata: µStomata¶ is a pore through which the evaporation of water is regulated by plants and also the passage of air and carbon dioxide through the network of cells within the leaf. stoma closes. the stoma opens and when the guard cells lose water.It can open and close as the plant dictates. 3.The mechanism namely the opening and closing of stomata depends upon the turgor pressure in the guard cells.

Active Potassium Theory (Levitt 1974): It was observed by Fujino (1967) that opening of stomata occurs due to the influx of K+ ions into the guard cells. ATP helps in entry of K+ ions into the guard cells. The source of K+ ions are the neighboring subsidiary and epidermal cells. Department of Biotechnology Feb11 -Mar11 Page 21 . thereby increasing the concentration from 50mM to 300mM in guard cells.PLANTS HAVE SENSE! 3.3. The increase in K+ ion concentration increases the osmotic concentration of guard cells thus leading to stomatal opening.

During periods of drought. Each stoma is controlled by two guard cells which can open and close the aperture between them. In the 1980s it wasdiscovered that the stomata start to close down the moment the roots detect dry soil. and their fourth sense is of substances produced by organisms on the leaf surface. The evaporation of water from a plant is regulated by pores (stomata) which can open and close as the plant dictates. They also control the passage of air and carbon dioxide through the network of cells within the leaf. It is believed that subsidiary cells have an active reabsorption mechanism of K+ ions. The guard cells do not simply control the pores. Third is their ability to respond to physical stimuli that affect the leaf. Theplants are anticipating a threat beforeIt arises. One of the simplest methods of demonstrating this effect is the split-root experiment. chloride and malate ions in stomatal opening (PEPcase = Phosphoenol pyruvate carboxylase) y y y Uptake of chloride (Cl-) ions Transport of H+ ions released from organic acid (malic acid) By negative charges of organic acids when they lose H+ ions Thus all these factors lead to the opening of stomata. these correspond to four of the classic senses: sight. they react to the intensity and the quality of light. If Department of Biotechnology Feb11 -Mar11 Page 22 .4. The uptake of K+ ions is balanced by: Role of potassium. Split root experiment: Although greenplants haveno nervous system. so that the pore opens. The stomatal closure is considered to be brought about by a passive or highly catalysed excretion of K+ ions and Cl.PLANTS HAVE SENSE! Levitt (1974) observed that proton (H+) uptake by guard cells. but when the guard cells enlarge. potassium chloride migrates into the cells and starch breaks down. which can be independently supplied with water. and long before there is any change in the water reaching the leaves. and which leads to the closure of the stomata before the plant experiences water loss. but sense what is going on around the plant. chloroplasts takes place with the help of ATP. First. the water flow up a plant diminishes and the stomata close down to minimise water loss.but we now know thatthey can also be controlled fromas faraway as the tips of the roots. the stomata can detect the chemical nature of the atmosphere. like vibrations and movement caused by wind. The mechanism seems to be some form of chemical signal which the plants can send to the leaves. responding to levels of carbon dioxide and other gases. the guard cells lose volume and the pore closes. Second. they can transmit messages through the length of the plant body.ions from the guard cells to the epidermal tissue in general and the subsidiary cells in particular. smell. A plant is induced to share its root system between two pots. As Terry Mansfield of the University of Lancaster has pointed out. It has long been believed that it was the closure of these pores in the leaf which reduced the water flow. touch and taste. 3. The stomata closure is due to excretion of K+ ions from guard cells surrounding epidermal and subsidiary cells. The chemistry is complex. This leads to increase in value of pH in guard cells. When the processes reverse. and potassium chloride is released from the cell as starch reforms. Rise in pH converts starch into organic acid like malic acid. It has long been thoughtthat the stomata open and close solely in response to what they sense.

The control of the stomata by the roots may be by means of abscisic acid. thesignalling system is severed and the stomataopen. Analysisof rootsfrom split-root plant experiments substantiates the possibility. for the stomata immediately open. The crucial role of the roots can be demonstrated by watering the dry soil. a hormone which can cause stomata toclose if present in very small amounts. One part of abscisic acid in abillion parts of water is enough to make them shut. As soon asthe roots are detached. This occurs even if there is a plentiful supply of water to the second pot. for there is always much more abscisic acid in the dry roots than in the moist ones. the stomata over the whole plant tend to close. Department of Biotechnology Feb11 -Mar11 Page 23 .PLANTS HAVE SENSE! the soil in one of the pots dries out. Confirmation is obtained by cutting off the rootsto the dry pot.

SOME SPECIAL SENSE OF PLANTS Department of Biotechnology Feb11 -Mar11 Page 24 .PLANTS HAVE SENSE! 4.

For example. Department of Biotechnology Feb11 -Mar11 Page 25 . then the tree may well do that for itself. a great cascadeof enzymes leadsto be set in train. Early in the twentieth century. Over the following years. Trees have the ability to configure their outline during their lifetime. A sense of gravity: Trees manage to grow in well-spaced patterns. London. research at Imperial College. When an electrical storm is approaching cells within the grass leaves begin to mobilise their metabolic processes. Before a dried-up plant can fully benefit from rainfall. This belief is not new. but they can react to disease by chemical responses which parallel some of those seen in animals. and when rainfall is due. experiments were carried out where high-tension cables were stretched across a field of growing crops.PLANTS HAVE SENSE! 4. no branches were lost from the tree. as a walk through woodland will confirm. To create centripetal force the plants were grown on a record player at 45(rpm)for 14 days. Plants have great regenerative powers. The lawn really does turn green and. who reports that a tree which was partly dislodged by a gale has altered its branches to regain its balance. located sensory cells within the plant which have the ability to sense electricity. When plants are subjected to centripetal force. It is true . The driedleaves need to return to an active state of metabolism. They employ mechanisms designed to prevent overcrowding. the sooner theprocess starts the better. Plants haveadapted to this fact. light. Some special sense of plants: 4. He observed that the tree had been left leaning down wind after the storm. We have no knowledge of how a tree does this. If you do not cut down a branch. ready for the rainfall. through the grass's extraordinary senses. 4. There is one experiment which helps us get a better understanding of how plants react to what they perceive is gravity. The physiological purpose of this is complex. they can shed their branches to maintain their equilibrium. An even more remarkable ability is reported by Bill Vinten in Suffolk. it does so before the first drops of rain begin to fall. The results showed that the plant did indeed 'green up' in response to an electrical field. A sense of electricity: Plants haveanother senseof which we have little knowledge .plants really do "green up" in thundery weather before the rain starts.2. and the way they heal themselves shows immense coordination of cellular growth. which would lead to competition for food. and water. but those that remained have grown round to restore the tree's centre of gravity. This takes time.1. Therehaslong been a beliefthat a lawnbecome'sgreener before the torrential downpour of a thunderstorm. Not only can plants communicate an attack by pests to other plants in the neighbourhood.the ability to detect an electrical field. A tree from which a branch has been cut covers the site with wound tissue and makes good the damage. During the 1990s. ready to receive the water when it comes. they react to the new direction of gravity by growing against it. and the maintenance of the outline of a tree is clearly a result of its sensory awareness and is worthy of further study.

all of the cacti dropped their needles by means of defense. it was discovered that a house cat had found its way into the greenhouse. After a period of a year of being without their protective quills.PLANTS HAVE SENSE! After 14 days on the record player. 4. however. He encouraged the plants to feel even more secure by playing soothing music in the greenhouse. Plants can feel: Once plants feel safe. This experiment concluded that plants grow against the vector sum of centripetal force and gravity. the scientist talked to numerous cacti assuring them that they were protected and that he cared about them. In one study. the cacti suddenly began re-growing their bristles and new baby sprouts were born with needles again.3. Defenseless within this nurturing environment. Within several months the cacti dropped all their spikes. a scientist wanted to determine if cacti grow needles primarily for the purpose of keeping themselves from harm. Once the cacti sensed they were once again safe. the mature and new-born cacti prospered. the plants grew at a horizontal 70° angle towards the center of the record player. After some investigation. Safely housed in a greenhouse. the scientist blocked the cat's way of entry. The offspring of these bare cacti were born without needles. they may drop their need for defense. Suspecting that the cat may be the source of the perceived threat to the cacti causing the reemergence of their means of protection. Department of Biotechnology Feb11 -Mar11 Page 26 .

CASESTUDIES Department of Biotechnology Feb11 -Mar11 Page 27 .PLANTS HAVE SENSE! 5.

Using anatomical (light.1. Casestudies: 5. scanning electron and transmission electron microscopy) and electrophysiological techniques.They also found that they can generate receptor potential following mechanical stimuli and they are in connection with excitable motor cells (through plasmodesmata). 5. These cells could be one ofunidentified mechanoreceptors of mimosa.Later these endodermal cells develops into gravity sensing cells. It lacks scr protein which is a transcription factor required for asymmetric cell divisions that are essential for endodermal differentiation in roots and shoots. Oxfordcarried out some research work to determine whethercircumutation depends on gravitropism. Gravitropism : directed growth in response to gravity. We alsoprovide evidence that these red cells are derived from stomatal subsidiary cells and notguard cells. They can generate receptor potential following mechanical stimuliand they are in connection with excitable motor cells (through plasmodesmata). The SCR gene regulates circumutationand gravitropism in Arabidopsisand morning glory: Scientists in Department of Botany. Department of Biotechnology Feb11 -Mar11 Page 28 . which is important in development of action potentials and movements of plants. Special red cells were found on the adaxial surface of tertiary pulvini of Mimosa pudicaand experiments performed to determine the origin and function of these cells.PLANTS HAVE SENSE! 5. which is important indevelopment of action potentials and movements of plants.Using anatomical (light.2.This mutant plant showed abnormality in its circumutation which indicates thatthe SCR gene regulates circumutation and gravitropism in Arabidopsis and morning glory.Mechanoreceptor Cells on the Tertiary Pulvini of Mimosa pudica: Special red cells were found on the adaxial surface of tertiary pulvini of Mimosa pudicaand experiments performed to determine the origin and function of these cells. Miami University. they have demonstrated that these red cells are real mechanoreceptor cells. They developed scr mutant (scarecrow) Arabidopsis . scanning electron and transmission electron microscopy) andelectrophysiological techniques.As result of histochemical studies they found red cells contain tannin. Circumutation : it is a oscillatory movements in which plants rotate around a central axis during their growth. As histochemical studies show red cells contain tannin. we have demonstrated that these red cells are realmechanoreceptor cells.

PLANTS HAVE SENSE! 6. CONCLUSION Department of Biotechnology Feb11 -Mar11 Page 29 .

for example). Some of the senses in the plant world are already more highly developed than ours (the sense of touch. they can detect chemicals(taste). and they cooperate to survive. or 'left' from 'right'. plants are our cousins. they have a sense of touch (sometimes to an extraordinary degree) they can smell . They have sight. their senses and the limits of their sentience are exactly what they require. Department of Biotechnology Feb11 -Mar11 Page 30 . Plants can remember stimuli and tell one form of stimulus from another.PLANTS HAVE SENSE! 6. We owe plants respect. they can here and through gravity they can tell 'up' from 'down'. No longer should science regard a green plant as a simple organism which endures what it must. Conclusion: Plants have many of the senses possessed by humans. As it is. they would have developed it. They can communicate. as far as they need it. They are not our subjects. If plants required more intelligence. and adjusts like a chemical system. for on green plants we all rely for survival.

REFERENCES Department of Biotechnology Feb11 -Mar11 Page 31 .PLANTS HAVE SENSE! 7.

http:// www. http://en. New York.tutorvista.html ³The Secret Language of Life´.com/ 6.wikipedia. Department of Biotechnology Feb11 -Mar11 Page 32 3. 1999.PLANTS HAVE SENSE! 7. Brian J. http://en. Fromm International. 2.php 5. Ford. References: 1. http:// www.

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PLANTS HAVE SENSE! 9. ARTICLES Department of Biotechnology Feb11 -Mar11 Page 49 .

Charles Darwin. circumnutation) of the shoot of a young seedlingof Brassica. Examples of tropisms include gravitropism. The experiment was performedover a period of 10 hand 45 min. in which plants rotate around a ce ntral axis during their growth. and all around: How plants sense and respond to environmental stimuli John Z. growth in response to light (3). Almost all plant parts exhibit this movement. but the direction is independent relative tothe stimulus source). The 2003 paper also reported on a series of gravitropic mutants in the plant Department of Biotechnology Feb11 -Mar11 Page 50 . OH 45056 Although plants are essentially sessile in nature. Fig. Although it is one of his lesser known works. published a fascinating book in 1880 entitled The Power ofMovement in Plants (1). bean plants) move to a different position at night.. The initial observations by this group (6) published in a previous paper (7)suggested that there was a link between circumnutation and gravitropism because an agravitropic mutant of the Japanese morning glory (Pharbitis nil) also was defective in circumnutation. Fig.who had a strong interest in plant biology during his multifaceted career.Up. To date. Nastic movements include the dramatic leaf movements of the Venus flytrap after a touch stimulus and the less dramatic (but more ubiquitous) µµsleep movements.g. 5).e. 1). Kiss* Department of Botany. This may come as a surprise to many nonbotanists. along with his son Francis.1. directed growth in response to gravity (2).Drawing from The Power of Movement in Plants (1) that illustrates the oscillatory movements(i. Miami University. The Darwins also studied oscillatory movements (also termed circumnutation. and phototropism. 1. Oxford. Articles: 9. these organisms are very much in tune with their environment and are capable of a variety of movements. but not to Charles Darwin.PLANTS HAVE SENSE! 9. they have been most recently considered by Kitazawa and coworkers(6) in a recent issue of PNAS.. down.¶¶ in which the leaves of some plants (e. and tracings wereof the shoot tip as viewed from above. Although the Darwins (1) were among the first to address these issues. The Darwins studied in great detail the two broad categories of plant movements: the tropisms (directed growth in response to external stimuli) and nastic movements (movements in response to stimuli. it is unclear whether these oscillatory movements are an endogenous nastic move ment or whether they are coupled to and depend on gravitropism. but vines show an exaggerated circumnutation (4.

which are the statocytes. and a transgenic Arabidopsis line with the mutant scrgene.] Thus. Additionally. that move in response to gravity (8). ( ) The stem consists of an epidermal layer (Ep). However. These cells contain amyloplasts (arrows) that function as statoliths and move in response to the direction ofthe gravity (toward bottom). the gravity-perceiving statocytes are located in the central columella cells of the root cap. Hatakedaet al. thus clearly linking these two processes. 2. They also found that mutation of PnSCR causes abnormal shoot circumnutation by studying gravitropism in wild-type morning glory. called statoliths. sgr2 and zig_sgr4.. whereas in stems. 2). another Arabidopsis mutant (scrfor scarecrow) lacking an endodermis that was agravitropic in stems had an even more severe lack of circumnutation. Fig. the results of some experiments (11) on the space shuttle Columbia would no doubt have heartened Charles Darwin. (a) Within the root cap at the apex. these statocytes are found in a layer of cells adjacent to the vascular tissue and are termed endodermal cells (Fig. or gravity-perceivingcells. The endodermal cells (En. Both columella and endodermal cells have dense amyloplasts (i. other Arabidopsis mutants that have the endodermis but with abnormal statolith movement (e. a morning glory scrmutant. The results of these studies and the various control experiments clearly demonstrate that the SCR gene regulates both circumnutation and gravitropism in both Arabidopsis and morning glory.Diagram of a root (a) and stem (b) of Arabidopsis to illustrate the regions of gravit perception.g. with small statoliths) of Arabidopsis that had reduced gravitropism also exhibited a decreased circumnutation. Sunflower seedlings were Department of Biotechnology Feb11 -Mar11 Page 51 . nil Scr (PnSCR) using the scrmutant of Arabidopsis. In addition. some y cells develop into the gravity-sensing columella cells (yellow).. Those mutants were deficient in the early steps of gravitropismbecause they had defects in their statocytes.e. In roots. (7) found that a starchless mutant (thus.PLANTS HAVE SENSE! Arabidopsis thaliana. and vascular tissue (VT). The hypothesis that the SCR protein in morning glory is necessary for the differentiation of the endodermis and gravity responses in stems was supported by complementation tests for P. yellow) are the statocytes with amyloplast (arrows) s functioning as statoliths. 10) were defective in the oscillatory movements. starch-filled plastids). ref. these results do not support the hypothesis originally proposed by Darwin and Darwin (1) that circumnutation had an internal driving force and regulating apparatus but add support to the idea that oscillatory movements in plants are directly coupled to gravity sensing. 6 suggests that the endodermal cells also play an important role in circumnutation. Ref. The present study reported in PNAS (6) builds on the previous 2003 paper (7) in several interesting ways. a cortical zone b (C). [A bit ofbackground: the SCR gene belongs to a gene family that encodes putative transcription factors required for asymmetric cell divisions that are essential for endodermal differentiation in roots and shoots (9).So it seems that the Darwins (1) may not have been right on this issue. an endodermal layer (En).

circumnutations) persist in the absence of significant gravitational accelerations. Kitazawa and coworkers (6) propose that because in these spaceflight experiments (11. H.. these results are in conflict with the study reported by Kitazawa et al. Suge. Plant Biol. 14. Thus. H. Alternatively. Fujisawa. Darwin. (2000) Crit. H. 712±718. However. which is published as supporting information on the PNAS web site). S. & Takahashi. Nevertheless. &Driss-Ecole. 464±473. R. A.and two hypotheses about the relationship between gravitropism and circumnutation? One unsatisfying suggestion is that the differences are caused by plantspecies differences. Acad. 94. (1984)Science 225. 6. Brown. Z. 2.. 533±538. (1880) The Power of Movement in Plants (John Murray. Goto. & Masson. H. some of the plants studied developed from seeds planted in orbit by the astronauts. 12. Y. Hatakeda.. and then circumnutation continued atthe reduced amplitude and period reported (11). (1990) Plant Physiol. Brown. D. 12) the sunflower seedlings were germinated on the ground. Opin. Kato. Kiss. 9. one must assume that most of these seedlings exhibited circumnutation because 93% of the total plants had these oscillatory movements. J. Perbal. L. Edelmann assisted in the preparation of the figures. 156±165. & Darwin. Morita. A. D. T. M. H. J. M. F. (2000) Am. K. As a space biologist. M. 3. E. 10. USA 102. 551±573. K. 133. J. Blancaflor. Tasaka. P. 19... 4. (2003)Plant Physiol. 11. C.. the oscillatory movement was established on the ground. C. (6). Correll. F. M. Benfey. Brown and coworkers (11) studied 4-day-old seedlings in microgravity and found that 93% of these plants circumnutated compared with 100% of the ground control. H. The circumnutation of the spaceflight seedlings had a reduced amplitude and period compared with the ground plants. (2003) Physiol. Tasaka. et al. 8. is there more support for the endogenous model of Darwin and Darwin (1) or the hypothesis that suggests that circumnutation depends on gravitropism most recently advocated by Kitazawa et al. 21. N. 87. M. H. T. Kamada. Hoshino. Plant GrowthRegul.. & Chapman. 7. (2002) J. Although their studies do not distinguish the results obtained from plants that first developed on the ground from those plants that were completely grown in microgravity. Fukaki. M. (1993) Plant Physiol. Mullen for helpful comments on the manuscript. 13. &Tasaka. Z.. H..Natl. J. there is strong support for the latter hypothesis provided for by the elegant study with multiple molecular appr aches outlined in ref. 5. Wysocka-Diller. Y. A. Chapman. Thus. 7. P. Hatakeda.. G. &Tasaka. 118. H..e. Hangarter for providing Movie 1 and Richard C.. 230±232. &Venditti. N. Kitazawa. M.. (2002)J. Lewis.. Department of Biotechnology Feb11 -Mar11 Page 52 . A.. B. Bot. Based on the evidence to date.. 21. Fukaki. Miyazawa. M. Fukaki. Larson. K. Y. Brown... REFERENCES 1. 6 and o theprevious studies from this group that used more structural methods (7). Plant Growth Regul. A. the spaceflight study seemed to be definitive and provides strong evidence that periodic growth oscillations (i. Maria Palmieri and Richard E. Kamada. H. 101. (2004)Curr. I hope there will be opportunities to confirm and extend these experiments by using the microgravity environment aboard orbiting spacecraftas a unique research tool (13). D. M. the plants sensed gravity before and during the launch. in the space experiments.. Rev. (2005) Proc. Morita.. 345±348. Moore and Jack L. N. 89±101.PLANTS HAVE SENSE! used in these experiments because these plants exhibit a vigorous circumnutation(see Movie 1. London). T. 18742±18747. (6)? Clearly. D. 233±238.. I thank Roger P.. & Kiss. Plant Sci. J. 1677±1690. (1998)Plant J. Fujii. Iida. Sci. 425±430. Plant.

however. In mimosa calcium can cause actomyosin microfilaments to contract.4 Toriyamaand Satô investigated a fine fibrillar structure in the central vacuole of motor cells. scanning electron and transmission electron microscopy) andelectrophysiological techniques.. i.2 It is known that movements of mimosa are governed by quick turgor pressure changes through vacuolar systems. We alsoprovide evidence that these red cells are derived from stomatal subsidiary cells and notguard cells. the fine fibrillar structure becomes an aggregated structure.13 Hollins and Jaffe. Most of the authors agree that mechanical stimuli are translated to membrane potential changes and at the end this electrical signal is necessary for movements of motor cells.16-19 Some results. and 3Plant Anatomy. As a result of this spectacular movement. 2 Chloride and potassium efflux were 1 measured on the primerypulvinus successfully. These cells could be one ofunidentified mechanoreceptors of mimosa.2.7 This double vacuolar system is a characteristic sign of excitable motor cells. Faculty of Sciences. that does not contain tannin. and this contraction can open putative potassium channels. In that. what cells are the mechanoreceptors. derives from osmotic pressure of potassium and chloride ions. for example. mimosa has been the subject of many studies since the 18th century. which is transformed into leaf movements.20. tannin containing and tannin less vacuoles.11 The power of motor cells. EötvösLoránd University.9 As electron micrographs show the osmiophilic tannin vacuolar contents and colloidal contents are sometimes associated (colloidal theory). Introduction Mimosa is famous for its rapid leaf movements. hormones (turgorins) may be the most important factors in this signal transduction. As histochemical studies show red cells contain tannin.6They reported on the cooperation between tannin and the central vacuole. detectable potassium salts are found in the motor cells before seismonastic response.15 but it is not known whether these interactions are artefactsof fixation or whether they are physiologically significant. we have demonstrated that these red cells are realmechanoreceptor cells. The leaves close up and droop when touched and reopen within a few minutes. after stimulation. using comparative histochemistry have shown that tannin vacuoles can also store calcium. Cells of pulvinar motor tissue have two different types of vacuoles.10 Histochemically.Using anatomical (light.5.11. with microelectrodes.14 These tannin vacuoles are analogous to sarcoplasmatic reticulum in muscle sarcomers that release Ca2+ into the cytoplasm. Institute of Biology.1 but there are still many questions in connection with the movement of mimosa. The contractile mechanism can be involved in the seismonastic reaction of the pulvinarmotor cells.21. They can generate receptor potential following mechanical stimuliand they are in connection with excitable motor cells (through plasmodesmata).9 Most data show that potassium salts caused turgor pressure changes in motor cells.3 Several data demonstrate that the large central vacuole. during seismonastic reaction.13.22 Department of Biotechnology Feb11 -Mar11 Page 53 .2. which is important indevelopment of action potentials and movements of plants. indicate that the role of organic molecules. Hungary Abstract Special red cells were found on the adaxial surface of tertiary pulvini of Mimosa pudicaand experiments performed to determine the origin and function of these cells.e. is a contractile vacuole.8 and describes movements of plants involving a process based not primarily on contractile proteins.* Ildikó Világi2 Petra Varró2 Zoltán Kristóf3 Departments of 1Plant Physiology and Molecular Plant Biology. 2 Physiology and Neurobiology. but the large crystals of the salts appear in the intercellular spaces after response. Budapest.PLANTS HAVE SENSE! 8.Mechanoreceptor Cells on the Tertiary Pulvini of Mimosa pudica Tamás Visnovitz1.

Plants of Mimosa pudicaL.37-40 Data have been published about the importance of electrical signals in reproductive systems41 and also in the electrical perception and µdeath message¶ in Characells.30 The characteristics of action potential can be modified by changing of Cl or K+ concentration. the impulse can control motor cell movements through voltage proton ATPase. Plasmodesmata are nearly identical to gap junctions of animal tissues. with µall or none¶ manner.32. This electrical stimulus spreads and coordinates the movements of motor cells in mimosa.17.26 The first review about this topic was published in 1973. and the K+ efflux repolarizes the plasma membrane. the action potential passes in the phloemtubes. stretch activated ion channels in Characells have been identified as mechanoreceptors. and investigate how they generate a receptor potential that is adequate for motor cell stimulation.Microscopy. However.29 After arrival of the stimulus. This variation potential can be a connection with wound hormone of Ricca. 33 In plants.42.18.35 Electrical activity in plants (action potentials. and this connection has high solute permeability and electrical conductivity.43 In case of mimosa. Depolarisation occurs due to Ca2+ activation of Ca2+ dependent anion channels and massive efflux of Cl .32 Plasmodesmata can be ³synapses of plants. the occurrence of action potentials in plants is a known phenomenon and described frequently.23 In plants (like animals) action potentials can be measured after external stimuli.28 These two types of potential changes of mimosa were measured exactly by the techniques of aphid stylet. through voltage dependent ion channels and from vacuoles via secondary transduction pathways. leaflets were fixed in2. and theseleaflets were placed on the surface of ¼ strength Hoagland solutionin Petri dishes. similar to the glutamate receptors in the nervous system of animals. for example in Arabidopsis thaliana. This Ca2+ originates from extracellular and intracellular spaces.34 These glutamate receptors. The mechanoreceptors responsible for the receptor potential in mimosa are unknown. variation potential and voltage transients) can be a mechanism for signal propagation and takes place in numerous physiological responses at the molecular and systemic level. Leaves for electrophysiologicalstudies were cut off at the petiole between the primary and secondarypulvinus or between the secondary and tertiary pulvinus. In plants. Materials and Methods Plant material.31 Electrical signals may pass through gap junctions (electrical synapses) at speed of 2 to 500 mm/s in animals. action potential (evoked by mechanical stimulation or with cold water application) and variation potential or slow wave potential (after wounding).PLANTS HAVE SENSE! Nowadays.45 The potential change and its characteristic is one of the most important factors in the control of motor cells. Ex vivo leaves were left for one or two days floatingon the ¼ strength Hoagland solution to stabilize all spontaneous ionfluxes before measurements were made.36 The role of action potentials is described in case of induction of systemic proteinase inhibitor.44 When the action potential arrives to the motor cells.19. Depolarization leads to opening of K+ channels. Experimental data indicate that adding 1 mM glutamate into Arabidopsis seedlings¶ growth medium causes a rapid increase in cytosolic free Ca2+ and also membrane depolarization. most cells have cell to cell conduction through plasmodesmata.5% Department of Biotechnology Feb11 -Mar11 Page 54 . action potentials were demonstrated in 1962.16 In that article two different types of membrane potential changes of mimosa were described.25 In case of Mimosa pudica. In plants the most important ions in this process are Ca2+ and Cl . can contribute to electrical signal transduction in plants. were grown in thegreenhouse at 25°C in soil culture. The membrane potential changes are generated in unidentified mechanoreceptor cells as a receptor potential. For electron microscopy. but action potentials can also be generated spontaneously.17. Action potential develops and propagates as an impulse in the axon of neurons. ionotropic glutamate receptors were also found.24 The first description of an µaction potential¶ was an electric disturbance following stimulation of a leaf of Dionaea muscipulata.19 There is a very good understanding about the mechanism of action potential generation in animals.´ and from a special point of view auxin can be the mediator/neurotransmitter. Na+ an K+ ions d play a key role in it.46 The goal of the present study was to describe mechanosensitive cells on the tertiary pulvinus of Mimosa pudicaL. free Ca2+ concentration in the cytoplasm increases.27.

advanced Intrasysprogram (Experimetria) and plotted in Origin 5. Arrows point to red receptor cells. Supertech. Histochemical studies revealed that these special red cells contain tannin.P.For light microscopy. Therefore. stars indicate guard cells (out of work). Figure 1. Instrutech Corp. To identify tannin.1 M potassium sodiumphosphate. previously undescribed cells on the tertiary pulvinus of Mimosa pudicaL. which is a typical reaction of tannin.0 (Microcal) program. These cells have bright red color without any staining. Hungary) and displayedon a digital oscilloscope (Gould DSO 420.E. Thus we were interested in their role with respect to the seismonastic reaction. we determined to examine the nature and role of these cells. Stored signals were analysedwith S. Red receptor cells (without staining) on tertiary pulvinus of Mimosa pudicaresemble stomatal guard cells.) and stored on a VHS tape for further analysis. E. After insertion of recording electrode into the cell there was a 20 minute wait before initiating mechanical stimulation. samples were stained with FeSO4 according to Ruzin48 and were studied with Zeiss SMX stereo microscope and Olympus BH 2light microscope for examination. Arrows show these bright red cells (A) Receptor cells and receptor complex on tertiary pulvinus of mimosa by light microscope (B) and by scanning electron microscope (C). Recorded signals were digitized online with an A/D converter (VR 10.PLANTS HAVE SENSE! glutaraldehyde buffered with 0. elongated excitable motor cells. the fixed and dehydrated leaf pieces were transferred to iso amyl acetate and than dried with a Polaron CPD 7501 critical point dryer. The presence of red stomatal guard cells on the adaxial part of the tertiary pulvini was very strange. For scanning electron microscopy. pH 7. and embedded in Spurr or Durcupanepoxi re sin. then stained with toluidine blue (pH 4. and at first sight they resemble stomatal guard cells (Fig. Recording electrodes were positioned with a Märzhausermicromanipulator under stereomicroscope control (Zeiss SMX).75 mm diameter borosilicate microcapillaries(Medical System).Reference electrode was placed in the bottom of recording chamber in contact with the ¼ strength Hoagland solution surrounding the plant material. M. We discovered that the location of these cells completely overlapped the area that is sensible to fine mechanical stimulation.2. Tissues were postfixed in 1% OsO4. dehydrated in an ethanol series. motor cells. Measures were performed in a Faraday cage. 1A).4) and examined using Olympus BH 2 light microscope. Sections were stained with 5% uranyl acetate and lead citrate47 and examined using a Hitachi 7100 transmission electron microscope at 75 kV. semi thin sections (1 m) were cut with Zeiss Hm 360 microtome. we identified some special. Thin sections (70 nm) were cut by Reichert Ultracut E ultramicrotome.L. Gould Electronics). Results Upon examination of the seismonastic reactions of the sensitive plant. Samples finally were coated with gold in a Zeiss vacuum evaporator. Department of Biotechnology Feb11 -Mar11 Page 55 . One-fourthstrengthHoagland solution was supplied with peristaltic pump (Pharmacia). Electrophysiology. During the whole experiment the same light intensity was ensured with Tungsram bulb (Tungsraflex R63). Observations were made by Hitachi 2360N scanning electron microscope at 18 kV. Recording microelectrodes were fabricated with a horizontal electrode puller Sutter P 97 from 0. Signals were amplified (Bioamp. Receptor potentials in the red cells were recorded with glass microelectrodes (8±10 M ) filled with 3 M KCl and connected through an Ag/AgCl electrode to the amplifier. The tertiary pulvini were fixed in the presence of FeSO4 and the bright red coloration changed into black or deep blue.

Finally. Results from light microscopy (Fig. 1B). first hyper than depolarisation and finally repolarisation was detected. Under the receptors. 3B).2). the receptor cells and excitable cells are connected through plasmodesmata (Fig. In our electrophysiological investigations changes of membrane potential were measured after mechanical stimulation. Development of receptor potential through the plasma membrane was demonstrated (Fig.PLANTS HAVE SENSE! These red cells seemed to be good candidates for the unknown mechanoreceptor cells. and scanning electron microscopy (Fig. Motor cell movement requires special membrane potential changes.19 These data strongly implicate these red cells as real mechanoreceptor cells. the leaflet closed. Figure 2. When trichomes or epidermis cells were touched. after the artifact of stimulation. The receptor potential measured in agreement with the potential changes measured in motor cells. The answer was yielded by transmission electron microscopy. direct mechanical stimuli different types of epidermal cells (trichome. As the diagram shows. We then explored the anatomical structure of these mechanosensitive cells. but when the red cells were stimulated.13 Several of these recordings (more than 15) were made. Moreover. The receptor complex consists of two guard cells between two subsidiary cells (Fig. epidermis cells and the specialisedredcells) were stimulated with a micromanipulator needle. 1C) studies certified that the red cells do not derive from stomatal guard cells. Department of Biotechnology Feb11 -Mar11 Page 56 . 3A). there was no reaction. They appear to derive from stomatal subsidiary cells. and that receptor potential should propagate to the motor cells. A representative receptor potential on red receptor cells aftermechanical stimuli (A) and electrical record on an epidermal cell neartoadaxial part of tertiary pulvinus of mimosa. as the electron micrographs show. potential changes varied between 10±12 mV. so a real mechanoreceptor cell has to produce a receptor potential. Electrical signals can e asily pass down this path. motor cell like elongated cells are situated. with typical double vacuolar system. we investigated whether there were connections between the receptor cells and the motor cells and what the nature of this connection was. To produce fine. (B) The resting membranepotential of receptor cells was about 90±100 mV and on a simple epidermis cell that was about 30±40 mV.

after mechanical stimulus a receptor potential is generated in these cells. Tannin vacuoles can function similarly to the sarcoplasmatic reticulum of animal muscle cells. the red cells would be able to produce a receptor potential whi h is essential for functioning as a c mechanoreceptor cell. which are able to control the movements of motor cells. are modified to control the pulvinar motor cells.46 Probably this mechanism is present also in mechanoreceptor cells of mimosa. As our electrophysiological measurements demonstrated.14 Similarly. This receptor potential can pass through plasmodesmata.52 so it seems logical that stomatal subsidiary cells. One of the most important questions was whether the red receptor cells can produce a receptor potential that can pass down to motor cells. Department of Biotechnology Feb11 -Mar11 Page 57 .9. Additionally. E.PLANTS HAVE SENSE! Discussion We have found red mechanoreceptive cells on the tertiary pulvini of Mimosa pudicaL. Their presence has been demonstrated in the plasma membrane of stomatal guard cells. However. the receptor and motor cells are in connection through plasmodesmata. Figure 3. These types of molecular receptors have been characterised several times. are real receptor cells. guard cells (out of work). and after stimulation they are able to release calcium ions.45 These results supported our original hypothesis that the red cells on the adaxial part of the tertiary pulvini in Mimosa pudicaL. elongated excitable motor cells. that normally coordinate the movements of stomatal guard cells. the question of how they became sensitive for mechanical stimuli remains unanswered. normal epidermis cell. After the stimulation of these cells the tertiary pulvinus closes. A question which arises is why stomatal subsidiary cells have transformed to mechanoreceptor cells. They sense the mechanical stimuli and operate the closure mechanism of leaflets. By means of changesin membrane potential the receptor cells are able to control the pulvinar motor cells. Histochemical studies show that the red coloration of the receptor cells is due to their tannin content. Subsidiary cells in the stomatal complex play a role in the control of guard cells taking part in ion channel mediated opening and closing of the stomatal pore.32 Both electron microscopy studies and stimulation with glass pin confirmed that these cells differ from the surrounding epidermal cells. red receptor cells. They are huge calcium stores. Receptor complex on tertiary pulvinus of mimosa by transmission electron microscope (A) and plasmodesmata (arrow) between the red receptor cell and the elongated excitable motor cell (B). The stimulation of these red cells with a micromanipulator needle proved that the stimulus evoked the closure of leaflets.50 Potassium and chloride flux between guard cells and subsidiary cells were also observed.51 Stomatal guard cells and pulvinar motor cells in mimosa operate with the same mechanism. G. which is an electric junction between these two different types of cells. R.49 It is well known that electrical changes can be measured on the plasma membrane of subsidiary cells during stomatal movements.53 and it has also been proven that stretch activated (mechanosensitive) channels translate mechanical stimuli (mechanical energy) into electrical code. Ep. as electron micrographs reveal. Stretch activated ion channels are likelyto play an essential role in mechanoperception.18.

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