Thermotropism

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Thermotropism or thermotropic movement is the movement of an organism or a part of an organism in response to heat or changes from the environment's temperature. A common example is the curling of Rhododendron leaves in response to cold temperatures. Mimosa pudica also show thermotropism by the collapsing of leaf petioles leading to the folding of leaflets, when temperature drops. [1]

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The term "thermotropism" was originated by French botanist Philippe Van Tieghem in his 1884 textbook Traité de botanique. Van Tieghem stated that a plant irradiated with an optimum growth temperature on one side laterally, and a much higher or lower temperature on the opposite side, would exhibit faster growth on the side exposed to optimum temperature. [2]

The definition of thermotropism can sometimes be confused with the term, thermotaxis, a mechanism by which temperature gradients can alter the behavior of cells, such as moving toward the cold environment. [3] The difference between them is that thermotropism is more commonly used in botany because it could not only represent the movement in organism level, thermotropism could also represent an organ level of movement, such as movement of leaves and roots toward or away from heat; but thermotaxis can only represent locomotion at the organism level, such as the movement of a mouse away from a warm environment.

The precise physiological mechanism enabling plant thermotropism is not yet understood. [4] It has been noted that one of the earliest physiological responses by plants to cooling is an influx of calcium ions from the cell walls into the cytosol, which increases calcium ion concentration in the intracellular space. This calcium influx is dependent upon mechanical changes in the actin cytoskeleton that alter the fluidity of the cell membrane, which allows calcium ion channels to open. From this information, a hypothesis has formed that the plant cell plasma membrane is an important site of plant temperature perception. [5]

Thermotropism in leaves

Rhododendron leaves curling in response to cold temperatures. Nature's Thermometers.jpg
Rhododendron leaves curling in response to cold temperatures.

Gardening hobbyists have frequently noted the dramatic change in the shape of Rhododendron or "Rhodie" leaves during warm versus cold weather. In warm weather, the leaf has a flat oblong shape. As the temperature of the leaf drops, the blade curls inward, giving the leaf a tubular, cigar-like shape.

Research on Rhododendron leaf thermotropism suggests that the curling response might help prevent damage to cell membranes caused by rapid thawing after a freeze. During the winter months, wild Rhododendrons in the Appalachian Mountains regularly drop to freezing temperatures at night, then thaw again in the early morning. Because a curled leaf has less of its surface area exposed to the sunlight, the leaf will thaw more slowly than it would if it were unfurled. Slower thawing minimizes damage caused to leaf cell membranes by ice crystal formation. [4]

Although there is little known about the molecular mechanisms of this rolling behavior, turgor pressure is responsible for the leaf movement. The exact stimulus for this output is not understood, but it is known that freezing cold temperatures causes an influx of water to the leaf petiole. As the turgor pressure increases, the leaves roll up, making it tighter to the stem. The leaf also droops perpendicular to the ground. There are predictions on the mechanism of this behavior. Regional changes of cell hydration can cause the inward curling. Another prediction is a change in cell wall physiology. [6] These predictions are very broad, indicating the need for further research.

There are currently two hypotheses to why Rhododendrons do this. The first is that the shape is more effective for snow shedding and better protects the more sensitive areas. Another hypothesis for leaf rolling called the desiccation theory, circulating in recent years, is to prevent membrane and light damage. [7]

In a 2017 study about cold stressed Rhododendron leaves showed that photosynthetic proteins decreased, while proteins for cell permeability increased. The same study showed the highest increase in proteins were responsible for transcription and translation regulation. [8] Thermotropic response in rhododendron leaves protects cells by changing leaf shape and protein levels.

Thermotropism in roots

The roots of some plants, including Zea mays , have been shown to bend differently when exposed to different temperature conditions. In general, growing roots tend to bend away from warmer temperatures, and towards cooler temperatures, within a normal range. It has been suggested that this growth behavior is beneficial because in most natural environments, soil closer to the ground's surface is warmer in temperature, while deeper soil is cooler. [9]

Experimentation with maize has demonstrated the existence of thermotropic responses in roots, with stronger responses seen when the thermal gradient increases. Positive thermotropism, or growth towards higher temperatures, was shown to occur at lower temperatures, with the strongest response observed at a temperature of 15 C. As the temperature increases, the strength of the response decreases. With continually temperature increases, a lack of thermotropic response is observed and occurs once a temperature threshold is reached. This threshold is dependent on the thermal gradient, with the threshold being colder with smaller gradients. For example, a gradient of 4.2 C per cm had a threshold value of 30 C while a gradient of 0.5 C per cm had a threshold value of 24 C. It is thought that this lack of thermotropic response is due to the lack of sufficient stimuli to induce root curvature. Negative thermotropic behavior was recorded and was shown to occur at higher temperature, but the conditions to establish such behavior is less defined. [10]

Within the same experiment, roots were capable of undergoing positive thermotropism away from gravitational force. The inhibition of normal gravitropic curvature was seen when temperatures were 18 C and lower, with stronger curvature away from gravity seen with lower temperature. This overriding behavior indicates integration of the plant's gravitropic and thermotropic system and suggests that the sensory systems are an interconnected network of responses rather than separate stimulation response pathways. [11]

Thermotropism's relation with Heliotropism

Para-heliotropic movements in the Phaseolus genus (beans) coincided with regulating leaf temperatures to improve photosynthesis efficiency and heat avoidance in hot, sunny, and arid environments. These movements worked to avoid photo-inhibition and keep leaf temperature lower than the air temperature. [12] In sunflowers, we find a different relation involving floral warming. The floral heads of these plants follow the sun from east to west causing increased solar irradiation heating the plant. This resulted in more pollinators being attracted. A study showed this by forcing some floral heads to the west leaving other floral heads to warm illustrating varied pollinator choice. [13] Though these findings are in correlation with heliotropisms, these heat avoidance and acquisition strategies are entwined with thermotropism as well. With further research, more examples can be found that can definitively detail the thermotropic role in heat avoidance and acquisition.

Related Research Articles

<span class="mw-page-title-main">Thigmotropism</span> Directed growth of plants in response to touch

In plant biology, thigmotropism is a directional growth movement which occurs as a mechanosensory response to a touch stimulus. Thigmotropism is typically found in twining plants and tendrils, however plant biologists have also found thigmotropic responses in flowering plants and fungi. This behavior occurs due to unilateral growth inhibition. That is, the growth rate on the side of the stem which is being touched is slower than on the side opposite the touch. The resultant growth pattern is to attach and sometimes curl around the object which is touching the plant. However, flowering plants have also been observed to move or grow their sex organs toward a pollinator that lands on the flower, as in Portulaca grandiflora.

Cold hardening is the physiological and biochemical process by which an organism prepares for cold weather.

Shade avoidance is a set of responses that plants display when they are subjected to the shade of another plant. It often includes elongation, altered flowering time, increased apical dominance and altered partitioning of resources. This set of responses is collectively called the shade-avoidance syndrome (SAS).

<span class="mw-page-title-main">Ampullae of Lorenzini</span> Sensory organs in some fish that detect electrical fields

Ampullae of Lorenzini are electroreceptors, sense organs able to detect electric fields. They form a network of mucus-filled pores in the skin of cartilaginous fish and of basal bony fishes such as reedfish, sturgeon, and lungfish. They are associated with and evolved from the mechanosensory lateral line organs of early vertebrates. Most bony fishes and terrestrial vertebrates have lost their ampullae of Lorenzini.

<span class="mw-page-title-main">Phenotypic plasticity</span> Trait change of an organism in response to environmental variation

Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompasses all types of environmentally induced changes that may or may not be permanent throughout an individual's lifespan.

<span class="mw-page-title-main">Neuronal noise</span> Random electric fluctuations in neurons

Neuronal noise or neural noise refers to the random intrinsic electrical fluctuations within neuronal networks. These fluctuations are not associated with encoding a response to internal or external stimuli and can be from one to two orders of magnitude. Most noise commonly occurs below a voltage-threshold that is needed for an action potential to occur, but sometimes it can be present in the form of an action potential; for example, stochastic oscillations in pacemaker neurons in suprachiasmatic nucleus are partially responsible for the organization of circadian rhythms.

<span class="mw-page-title-main">Thigmonasty</span> Undirected movement in response to touch or vibration

In biology, thigmonasty or seismonasty is the nastic (non-directional) response of a plant or fungus to touch or vibration. Conspicuous examples of thigmonasty include many species in the leguminous subfamily Mimosoideae, active carnivorous plants such as Dionaea and a wide range of pollination mechanisms.

Ecophysiology, environmental physiology or physiological ecology is a biological discipline that studies the response of an organism's physiology to environmental conditions. It is closely related to comparative physiology and evolutionary physiology. Ernst Haeckel's coinage bionomy is sometimes employed as a synonym.

<span class="mw-page-title-main">Plant perception (physiology)</span> Plants interaction to environment

Plant perception is the ability of plants to sense and respond to the environment by adjusting their morphology and physiology. Botanical research has revealed that plants are capable of reacting to a broad range of stimuli, including chemicals, gravity, light, moisture, infections, temperature, oxygen and carbon dioxide concentrations, parasite infestation, disease, physical disruption, sound, and touch. The scientific study of plant perception is informed by numerous disciplines, such as plant physiology, ecology, and molecular biology.

<span class="mw-page-title-main">Uncoupling protein</span> Mitochondrial protein

An uncoupling protein (UCP) is a mitochondrial inner membrane protein that is a regulated proton channel or transporter. An uncoupling protein is thus capable of dissipating the proton gradient generated by NADH-powered pumping of protons from the mitochondrial matrix to the mitochondrial intermembrane space. The energy lost in dissipating the proton gradient via UCPs is not used to do biochemical work. Instead, heat is generated. This is what links UCP to thermogenesis. However, not every type of UCPs are related to thermogenesis. Although UCP2 and UCP3 are closely related to UCP1, UCP2 and UCP3 do not affect thermoregulatory abilities of vertebrates. UCPs are positioned in the same membrane as the ATP synthase, which is also a proton channel. The two proteins thus work in parallel with one generating heat and the other generating ATP from ADP and inorganic phosphate, the last step in oxidative phosphorylation. Mitochondria respiration is coupled to ATP synthesis, but is regulated by UCPs. UCPs belong to the mitochondrial carrier (SLC25) family.

Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues. By contrast, an animal embryo will very early produce all of the body parts that it will ever have in its life. When the animal is born, it has all its body parts and from that point will only grow larger and more mature. However, both plants and animals pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.

<span class="mw-page-title-main">Pulvinus</span> Swollen or thickened leaf base

A pulvinus is a joint-like thickening at the base of a plant leaf or leaflet that facilitates growth-independent movement. Pulvini are common, for example, in members of the bean family Fabaceae (Leguminosae) and the prayer plant family Marantaceae.

Sperm guidance is the process by which sperm cells (spermatozoa) are directed to the oocyte (egg) for the aim of fertilization. In the case of marine invertebrates the guidance is done by chemotaxis. In the case of mammals, it appears to be done by chemotaxis, thermotaxis and rheotaxis.

<span class="mw-page-title-main">Infrared sensing in vampire bats</span>

Vampire bats have developed a specialized system using infrared-sensitive receptors on their nose-leaf to prey on homeothermic (warm-blooded) vertebrates. Trigeminal nerve fibers that innervate these IR-sensitive receptors may be involved in detection of infrared thermal radiation emitted by their prey. This may aid bats in locating blood-rich areas on their prey. In addition, neuroanatomical and molecular research has suggested possible similarities of IR-sensing mechanisms between vampire bats and IR-sensitive snakes. Infrared sensing in vampire bats has not yet been hypothesized to be image forming, as it was for IR-sensitive snakes. While the literature on IR-sensing in vampire bats is thin, progress continues to be made in this field to identify how vampire bats can sense and use infrared thermal radiation.

<span class="mw-page-title-main">Sperm thermotaxis</span>

Sperm thermotaxis is a form of sperm guidance, in which sperm cells (spermatozoa) actively change their swimming direction according to a temperature gradient, swimming up the gradient. Thus far this process has been discovered in mammals only.

Leaf expansion is a process by which plants make efficient use of the space around them by causing their leaves to enlarge, or wither. This process enables a plant to maximize its own biomass, whether it be due to increased surface area; which enables more sunlight to be absorbed by chloroplasts, driving the rate of photosynthesis upward, or it enables more stomata to be created on the leaf surface, allowing the plant to increase its carbon dioxide intake.

In plant biology, thermonasty is a nondirectional response to temperature in plants. It is a form of nastic movement, not to be confused with thermotropism, which is a directional response in plants to temperature. A common example of this is in some Rhododendron species, but thermonasty has also been observed in other plants, such as Phryma leptostachya. Flower opening in certain crocus and tulip species is also known to be thermonastic. These movements are thought to be regulated by having unequal cell elongation in certain plant tissues, causing different tissues to bend. In other processes, like in the temperature regulation of flower openings, movement has instead been shown to be a result of irreversible cell growth, a growth type not typically associated with plant movement. Furthermore, thermonasty has been shown to be independent of other environmental signals, such as light and gravity.

In plant biology, plant memory describes the ability of a plant to retain information from experienced stimuli and respond at a later time. For example, some plants have been observed to raise their leaves synchronously with the rising of the sun. Other plants produce new leaves in the spring after overwintering. Many experiments have been conducted into a plant's capacity for memory, including sensory, short-term, and long-term. The most basic learning and memory functions in animals have been observed in some plant species, and it has been proposed that the development of these basic memory mechanisms may have developed in an early organismal ancestor.

Hydraulic signals in plants are detected as changes in the organism's water potential that are caused by environmental stress like drought or wounding. The cohesion and tension properties of water allow for these water potential changes to be transmitted throughout the plant.

<span class="mw-page-title-main">Protist locomotion</span> Motion system of a type of eukaryotic organism

Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly unicellular and microscopic. Many unicellular protists, particularly protozoans, are motile and can generate movement using flagella, cilia or pseudopods. Cells which use flagella for movement are usually referred to as flagellates, cells which use cilia are usually referred to as ciliates, and cells which use pseudopods are usually referred to as amoeba or amoeboids. Other protists are not motile, and consequently have no built-in movement mechanism.

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