Phototropism

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The light from the lamp (1.) functions as a detectable change in the plant's environment. As a result, the plant exhibits a reaction of phototropism--directional growth (2.) toward the light stimulus. Phototrophic Response to Stimulus.svg
The light from the lamp (1.) functions as a detectable change in the plant's environment. As a result, the plant exhibits a reaction of phototropism--directional growth (2.) toward the light stimulus.
Auxin distribution controls phototropism. 1. Sunlight strikes the plant from directly above. Auxin (pink dots) encourages growth straight up. 2, 3, 4. Sunlight strikes the plant at an angle. Auxin is concentrated on one side, encouraging growth at an angle from the preceding stem. Phototropism Diagram.svg
Auxin distribution controls phototropism. 1. Sunlight strikes the plant from directly above. Auxin (pink dots) encourages growth straight up. 2, 3, 4. Sunlight strikes the plant at an angle. Auxin is concentrated on one side, encouraging growth at an angle from the preceding stem.
Phototropism in Solanum lycopersicum.

In biology, phototropism is the growth of an organism in response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi. The cells on the plant that are farthest from the light contain a hormone called auxin that reacts when phototropism occurs. This causes the plant to have elongated cells on the furthest side from the light. Phototropism is one of the many plant tropisms, or movements, which respond to external stimuli. Growth towards a light source is called positive phototropism, while growth away from light is called negative phototropism. Negative phototropism is not to be confused with skototropism, which is defined as the growth towards darkness, whereas negative phototropism can refer to either the growth away from a light source or towards the darkness. [1] Most plant shoots exhibit positive phototropism, and rearrange their chloroplasts in the leaves to maximize photosynthetic energy and promote growth. [2] [3] Some vine shoot tips exhibit negative phototropism, which allows them to grow towards dark, solid objects and climb them. The combination of phototropism and gravitropism allow plants to grow in the correct direction. [4]

Contents

Mechanism

There are several signaling molecules that help the plant determine where the light source is coming from, and these activate several genes, which change the hormone gradients allowing the plant to grow towards the light. The very tip of the plant is known as the coleoptile, which is necessary in light sensing. [2] The middle portion of the coleoptile is the area where the shoot curvature occurs. The Cholodny–Went hypothesis, developed in the early 20th century, predicts that in the presence of asymmetric light, auxin will move towards the shaded side and promote elongation of the cells on that side to cause the plant to curve towards the light source. [5] Auxins activate proton pumps, decreasing the pH in the cells on the dark side of the plant. This acidification of the cell wall region activates enzymes known as expansins which disrupt hydrogen bonds in the cell wall structure, making the cell walls less rigid. In addition, increased proton pump activity leads to more solutes entering the plant cells on the dark side of the plant, which increases the osmotic gradient between the symplast and apoplast of these plant cells. [6] Water then enters the cells along its osmotic gradient, leading to an increase in turgor pressure. The decrease in cell wall strength and increased turgor pressure above a yield threshold [7] causes cells to swell, exerting the mechanical pressure that drives phototropic movement.

Proteins encoded by a second group of genes, PIN genes, have been found to play a major role in phototropism. They are auxin transporters, and it is thought that they are responsible for the polarization of auxin location. Specifically PIN3 has been identified as the primary auxin carrier. [8] It is possible that phototropins receive light and inhibit the activity of PINOID kinase (PID), which then promotes the activity of PIN3. This activation of PIN3 leads to asymmetric distribution of auxin, which then leads to asymmetric elongation of cells in the stem. pin3 mutants had shorter hypocotyls and roots than the wild-type, and the same phenotype was seen in plants grown with auxin efflux inhibitors. [9] Using anti-PIN3 immunogold labeling, movement of the PIN3 protein was observed. PIN3 is normally localized to the surface of hypocotyl and stem, but is also internalized in the presence of Brefeldin A (BFA), an exocytosis inhibitor. This mechanism allows PIN3 to be repositioned in response to an environmental stimulus. PIN3 and PIN7 proteins were thought to play a role in pulse-induced phototropism. The curvature responses in the "pin3" mutant were reduced significantly, but only slightly reduced in "pin7" mutants. There is some redundancy among "PIN1", "PIN3", and "PIN7", but it is thought that PIN3 plays a greater role in pulse-induced phototropism. [10]

There are phototropins that are highly expressed in the upper region of coleoptiles. There are two main phototropism they are phot1 and phot2. phot2 single mutants have phototropic responses like that of the wild-type, but phot1 phot2 double mutants do not show any phototropic responses. [4] The amounts of PHOT1 and PHOT2 present are different depending on the age of the plant and the intensity of the light. There is a high amount of PHOT2 present in mature Arabidopsis leaves and this was also seen in rice orthologs. The expression of PHOT1 and PHOT2 changes depending on the presence of blue or red light. There was a downregulation of PHOT1 mRNA in the presence of light, but upregulation of PHOT2 transcript. The levels of mRNA and protein present in the plant were dependent upon the age of the plant. This suggests that the phototropin expression levels change with the maturation of the leaves. [11] Mature leaves contain chloroplasts that are essential in photosynthesis. Chloroplast rearrangement occurs in different light environments to maximize photosynthesis. There are several genes involved in plant phototropism including the NPH1 and NPL1 gene. They are both involved in chloroplast rearrangement. [3] The nph1 and npl1 double mutants were found to have reduced phototropic responses. In fact, the two genes are both redundant in determining the curvature of the stem.

Recent studies reveal that multiple AGC kinases, except for PHOT1 and PHOT2, are involved in plant phototropism. Firstly, PINOID, exhibiting a light-inducible expression pattern, determines the subcellular relocation of PIN3 during phototropic responses via a direct phosphorylation. Secondly, D6PK and its D6PKL homologs modulates the auxin transport activity of PIN3, likely through phosphorylation as well. Third, upstream of D6PK/D6PKLs, PDK1.1 and PDK1.2 acts an essential activator for these AGC kinases. Interestingly, different AGC kinases might participate in different steps during the progression of a phototropic response. D6PK/D6PKLs exhibit an ability to phosphorylate more phosphosites than PINOID.

Five models of auxin distribution in phototropism

In 2012, Sakai and Haga [12] outlined how different auxin concentrations could be arising on shaded and lighted side of the stem, giving birth to phototropic response. Five models in respect to stem phototropism have been proposed, using Arabidopsis thaliana as the study plant.

Five models showing how auxin is transported in the plant Arabidopsis. Model for auxin transport.png
Five models showing how auxin is transported in the plant Arabidopsis.
First model

In the first model incoming light deactivates auxin on the light side of the plant allowing the shaded part to continue growing and eventually bend the plant over towards the light. [12]

Second model

In the second model light inhibits auxin biosynthesis on the light side of the plant, thus decreasing the concentration of auxin relative to the unaffected side. [12]

Third model

In the third model there is a horizontal flow of auxin from both the light and dark side of the plant. Incoming light causes more auxin to flow from the exposed side to the shaded side, increasing the concentration of auxin on the shaded side and thus more growth occurring. [12]

Fourth model

In the fourth model it shows the plant receiving light to inhibit auxin basipetal down to the exposed side, causing the auxin to only flow down the shaded side. [12]

Fifth model

Model five encompasses elements of both model 3 and 4. The main auxin flow in this model comes from the top of the plant vertically down towards the base of the plant with some of the auxin travelling horizontally from the main auxin flow to both sides of the plant. Receiving light inhibits the horizontal auxin flow from the main vertical auxin flow to the irradiated exposed side. And according to the study by Sakai and Haga, the observed asymmetric auxin distribution and subsequent phototropic response in hypocotyls seems most consistent with this fifth scenario. [12]

Effects of wavelength

Phototropism in plants such as Arabidopsis thaliana is directed by blue light receptors called phototropins. [13] Other photosensitive receptors in plants include phytochromes that sense red light [14] and cryptochromes that sense blue light. [15] Different organs of the plant may exhibit different phototropic reactions to different wavelengths of light. Stem tips exhibit positive phototropic reactions to blue light, while root tips exhibit negative phototropic reactions to blue light. Both root tips and most stem tips exhibit positive phototropism to red light.[ citation needed ] Cryptochromes are photoreceptors that absorb blue/ UV-A light, and they help control the circadian rhythm in plants and timing of flowering. Phytochromes are photoreceptors that sense red/far-red light, but they also absorb blue light; they can control flowering in adult plants and the germination of seeds, among other things. The combination of responses from phytochromes and cryptochromes allow the plant to respond to various kinds of light. [16] Together phytochromes and cryptochromes inhibit gravitropism in hypocotyls and contribute to phototropism. [2]

See also

Related Research Articles

<i>Arabidopsis thaliana</i> Model plant species in the family Brassicaceae

Arabidopsis thaliana, the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally considered a weed.

<span class="mw-page-title-main">Root</span> Basal organ of a vascular plant

In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the surface of the soil, but roots can also be aerial or aerating, that is, growing up above the ground or especially above water.

<span class="mw-page-title-main">Auxin</span> Plant hormone

Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essential for plant body development. The Dutch biologist Frits Warmolt Went first described auxins and their role in plant growth in the 1920s. Kenneth V. Thimann became the first to isolate one of these phytohormones and to determine its chemical structure as indole-3-acetic acid (IAA). Went and Thimann co-authored a book on plant hormones, Phytohormones, in 1937.

<span class="mw-page-title-main">Phytochrome</span> Protein used by plants, bacteria and fungi to detect light

Phytochromes are a class of photoreceptor proteins found in plants, bacteria and fungi. They respond to light in the red and far-red regions of the visible spectrum and can be classed as either Type I, which are activated by far-red light, or Type II that are activated by red light. Recent advances have suggested that phytochromes also act as temperature sensors, as warmer temperatures enhance their de-activation. All of these factors contribute to the plant's ability to germinate.

<span class="mw-page-title-main">Coleoptile</span> Protective sheath in certain plants

Coleoptile is the pointed protective sheath covering the emerging shoot in monocotyledons such as grasses in which few leaf primordia and shoot apex of monocot embryo remain enclosed. The coleoptile protects the first leaf as well as the growing stem in seedlings and eventually, allows the first leaf to emerge. Coleoptiles have two vascular bundles, one on either side. Unlike the flag leaves rolled up within, the pre-emergent coleoptile does not accumulate significant protochlorophyll or carotenoids, and so it is generally very pale. Some preemergent coleoptiles do, however, accumulate purple anthocyanin pigments.

<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.

<span class="mw-page-title-main">Gravitropism</span> Plant growth in reaction to gravity and bending of leaves and roots

Gravitropism is a coordinated process of differential growth by a plant in response to gravity pulling on it. It also occurs in fungi. Gravity can be either "artificial gravity" or natural gravity. It is a general feature of all higher and many lower plants as well as other organisms. Charles Darwin was one of the first to scientifically document that roots show positive gravitropism and stems show negative gravitropism. That is, roots grow in the direction of gravitational pull and stems grow in the opposite direction. This behavior can be easily demonstrated with any potted plant. When laid onto its side, the growing parts of the stem begin to display negative gravitropism, growing upwards. Herbaceous (non-woody) stems are capable of a degree of actual bending, but most of the redirected movement occurs as a consequence of root or stem growth outside. The mechanism is based on the Cholodny–Went model which was proposed in 1927, and has since been modified. Although the model has been criticized and continues to be refined, it has largely stood the test of time.

In developmental biology, photomorphogenesis is light-mediated development, where plant growth patterns respond to the light spectrum. This is a completely separate process from photosynthesis where light is used as a source of energy. Phytochromes, cryptochromes, and phototropins are photochromic sensory receptors that restrict the photomorphogenic effect of light to the UV-A, UV-B, blue, and red portions of the electromagnetic spectrum.

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">Cryptochrome</span> Class of photoreceptors in plants and animals

Cryptochromes are a class of flavoproteins found in plants and animals that are sensitive to blue light. They are involved in the circadian rhythms and the sensing of magnetic fields in a number of species. The name cryptochrome was proposed as a portmanteau combining the chromatic nature of the photoreceptor, and the cryptogamic organisms on which many blue-light studies were carried out.

Phototropins are photoreceptor proteins that mediate phototropism responses in various species of algae, fungi and higher plants. Phototropins can be found throughout the leaves of a plant. Along with cryptochromes and phytochromes they allow plants to respond and alter their growth in response to the light environment. Phototropins may also be important for the opening of stomata and the movement of chloroplasts. These blue light receptors are seen across the entire green plant lineage. When Phototropins are hit with blue light, they induce a signal transduction pathway that alters the plant cells' functions in different ways.

<span class="mw-page-title-main">Lateral root</span> Plant root

Lateral roots, emerging from the pericycle, extend horizontally from the primary root (radicle) and over time makeup the iconic branching pattern of root systems. They contribute to anchoring the plant securely into the soil, increasing water uptake, and facilitate the extraction of nutrients required for the growth and development of the plant. Lateral roots increase the surface area of a plant's root system and can be found in great abundance in several plant species. In some cases, lateral roots have been found to form symbiotic relationships with rhizobia (bacteria) and mycorrhizae (fungi) found in the soil, to further increase surface area and increase nutrient uptake.

<span class="mw-page-title-main">Etiolation</span> Developmental pathway followed in flowering plants in absence of visible light

Etiolation is a process in flowering plants grown in partial or complete absence of light. It is characterized by long, weak stems; smaller leaves due to longer internodes; and a pale yellow color (chlorosis). The development of seedlings in the dark is known as "skotomorphogenesis" and leads to etiolated seedlings.

Photoreceptor proteins are light-sensitive proteins involved in the sensing and response to light in a variety of organisms. Some examples are rhodopsin in the photoreceptor cells of the vertebrate retina, phytochrome in plants, and bacteriorhodopsin and bacteriophytochromes in some bacteria. They mediate light responses as varied as visual perception, phototropism and phototaxis, as well as responses to light-dark cycles such as circadian rhythm and other photoperiodisms including control of flowering times in plants and mating seasons in animals.

<span class="mw-page-title-main">Cholodny–Went model</span> Botany model

In botany, the Cholodny–Went model, proposed in 1927, is an early model describing tropism in emerging shoots of monocotyledons, including the tendencies for the shoot to grow towards the light (phototropism) and the roots to grow downward (gravitropism). In both cases the directional growth is considered to be due to asymmetrical distribution of auxin, a plant growth hormone. Although the model has been criticized and continues to be refined, it has largely stood the test of time.

<span class="mw-page-title-main">Joanne Chory</span> American plant biologist

Joanne Chory is an American plant biologist and geneticist. Chory is a professor and director of the Plant Molecular and Cellular Biology Laboratory, at the Salk Institute for Biological Studies and an investigator of the Howard Hughes Medical Institute.

Circadian Clock Associated 1 (CCA1) is a gene that is central to the circadian oscillator of angiosperms. It was first identified in Arabidopsis thaliana in 1993. CCA1 interacts with LHY and TOC1 to form the core of the oscillator system. CCA1 expression peaks at dawn. Loss of CCA1 function leads to a shortened period in the expression of many other genes.

LUX or Phytoclock1 (PCL1) is a gene that codes for LUX ARRHYTHMO, a protein necessary for circadian rhythms in Arabidopsis thaliana. LUX protein associates with Early Flowering 3 (ELF3) and Early Flowering 4 (ELF4) to form the Evening Complex (EC), a core component of the Arabidopsis repressilator model of the plant circadian clock. The LUX protein functions as a transcription factor that negatively regulates Pseudo-Response Regulator 9 (PRR9), a core gene of the Midday Complex, another component of the Arabidopsis repressilator model. LUX is also associated with circadian control of hypocotyl growth factor genes PHYTOCHROME INTERACTING FACTOR 4 (PIF4) and PHYTOCHROME INTERACTING FACTOR 5 (PIF5).

Magnetotropism is the movement or plant growth in response to the stimulus provided by the magnetic field in plants around the world. As a natural environmental factor in the Earth, variations of magnetic field level causes many biological effects, including germination rate, flowering time, photosynthesis, biomass accumulation, activation of cryptochrome, and shoot growth.

EARLY FLOWERING 3 (ELF3) is a plant-specific gene that encodes the hydroxyproline-rich glycoprotein and is required for the function of the circadian clock. ELF3 is one of the three components that make up the Evening Complex (EC) within the plant circadian clock, in which all three components reach peak gene expression and protein levels at dusk. ELF3 serves as a scaffold to bind EARLY FLOWERING 4 (ELF4) and LUX ARRHYTHMO (LUX), two other components of the EC, and functions to control photoperiod sensitivity in plants. ELF3 also plays an important role in temperature and light input within plants for circadian clock entrainment. Additionally, it plays roles in light and temperature signaling that are independent from its role in the EC.

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