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. [1]
The photomorphogenesis of plants is often studied by using tightly frequency-controlled light sources to grow the plants. There are at least three stages of plant development where photomorphogenesis occurs: seed germination, seedling development, and the switch from the vegetative to the flowering stage (photoperiodism). [2]
Most research on photomorphogenesis is derived from plants studies involving several kingdoms: Fungi, Monera, Protista, and Plantae. [3]
Theophrastus of Eresus (371 to 287 BC) may have been the first to write about photomorphogenesis. He described the different wood qualities of fir trees grown in different levels of light, likely the result of the photomorphogenic "shade-avoidance" effect. In 1686, John Ray wrote "Historia Plantarum" which mentioned the effects of etiolation (grow in the absence of light). Charles Bonnet introduced the term "etiolement" to the scientific literature in 1754 when describing his experiments, commenting that the term was already in use by gardeners. [4]
Light has profound effects on the development of plants. The most striking effects of light are observed when a germinating seedling emerges from the soil and is exposed to light for the first time.
Normally the seedling radicle (root) emerges first from the seed, and the shoot appears as the root becomes established. Later, with growth of the shoot (particularly when it emerges into the light) there is increased secondary root formation and branching. In this coordinated progression of developmental responses are early manifestations of correlative growth phenomena where the root affects the growth of the shoot and vice versa. To a large degree, the growth responses are hormone mediated.
In the absence of light, plants develop an etiolated growth pattern. Etiolation of the seedling causes it to become elongated, which may facilitate it emerging from the soil.
A seedling that emerges in darkness follows a developmental program known as skotomorphogenesis (dark development), which is characterized by etiolation. Upon exposure to light, the seedling switches rapidly to photomorphogenesis (light development). [5]
There are differences when comparing dark-grown (etiolated) and light-grown (de-etiolated) seedlings
Etiolated characteristics:
De-etiolated characteristics:
The developmental changes characteristic of photomorphogenesis shown by de-etiolated seedlings, are induced by light.
Some plants rely on light signals to determine when to switch from the vegetative to the flowering stage of plant development. This type of photomorphogenesis is known as photoperiodism and involves using red photoreceptors (phytochromes) to determine the daylength. As a result, photoperiodic plants only start making flowers when the days have reached a "critical daylength," allowing these plants to initiate their flowering period according to the time of year. For example, "long day" plants need long days to start flowering, and "short day" plants need to experience short days before they will start making flowers. [2]
Photoperiodism also has an effect on vegetative growth, including on bud dormancy in perennial plants, though this is not as well-documented as the effect of photoperiodism on the switch to the flowering stage. [2]
Typically, plants are responsive to wavelengths of light in the blue, red and far-red regions of the spectrum through the action of several different photosensory systems. The photoreceptors for red and far-red wavelengths are known as phytochromes. There are at least 5 members of the phytochrome family of photoreceptors. There are several blue light photoreceptors known as cryptochromes. The combination of phytochromes and cryptochromes mediate growth and the flowering of plants in response to red light, far-red light, and blue light.
Plants use phytochrome to detect and respond to red and far-red wavelengths. Phytochromes are signaling proteins that promote photomorphogenesis in response to red light and far-red light. [6] Phytochrome is the only known photoreceptor that absorbs light in the red/far red spectrum of light (600-750 nm) specifically and only for photosensory purposes. [1] Phytochromes are proteins with a light absorbing pigment attached called a chromophore. The chromophore is a linear tetrapyrrole called phytochromobilin. [7]
There are two forms of phytochromes: red light absorbing, Pr, and far-red light absorbing, Pfr. Pfr, which is the active form of phytochromes, can be reverted to Pr, which is the inactive form, slowly by inducing darkness or more rapidly by irradiation by far-red light. [6] The phytochrome apoprotein, a protein that together with a prosthetic group forms a particular biochemical molecule such as a hormone or enzyme, is synthesized in the Pr form. Upon binding the chromophore, the holoprotein, an apoprotein combined with its prosthetic group, becomes sensitive to light. If it absorbs red light it will change conformation to the biologically active Pfr form. [6] The Pfr form can absorb far red light and switch back to the Pr form. The Pfr promotes and regulates photomorphogenesis in response to FR light, whereas Pr regulates de-etiolation in response to R light. [6]
Most plants have multiple phytochromes encoded by different genes. The different forms of phytochrome control different responses but there is also redundancy so that in the absence of one phytochrome, another may take on the missing functions. [6] There are five genes that encode phytochromes in the Arabidopsis thaliana genetic model, PHYA-PHYE. [7] PHYA is involved in the regulation of photomorphogenesis in response to far-red light. [6] PHYB is involved in regulating photoreversible seed germination in response to red light. PHYC mediates the response between PHYA and PHYB. PHYD and PHYE mediate elongation of the internode and control the time in which the plant flowers. [7]
Molecular analyses of phytochrome and phytochrome-like genes in higher plants (ferns, mosses, algae) and photosynthetic bacteria have shown that phytochromes evolved from prokaryotic photoreceptors that predated the origin of plants. [4]
Takuma Tanada observed that the root tips of barley adhered to the sides of a beaker with a negatively charged surface after being treated with red light, yet released after being exposed to far-red light. [8] For mung bean it was the opposite, where far-red light exposure caused the root tips to adhere, and red light caused the roots to detach. [9] This effect of red and far-red light on root tips is now known as the Tanada effect.
Plants contain multiple blue light photoreceptors which have different functions. Based on studies with action spectra, mutants and molecular analyses, it has been determined that higher plants contain at least 4, and probably 5, different blue light photoreceptors.
Cryptochromes were the first blue light receptors to be isolated and characterized from any organism, and are responsible for the blue light reactions in photomorphogenesis. [7] The proteins use a flavin as a chromophore. The cryptochromes have evolved from microbial DNA-photolyase, an enzyme that carries out light-dependent repair of UV damaged DNA. [10] There are two different forms of cryptochromes that have been identified in plants, CRY1 and CRY2, which regulate the inhibition of hypocotyl elongation in response to blue light. [10] Cryptochromes control stem elongation, leaf expansion, circadian rhythms and flowering time. In addition to blue light, cryptochromes also perceive long wavelength UV irradiation (UV-A). [10] Since the cryptochromes were discovered in plants, several labs have identified homologous genes and photoreceptors in a number of other organisms, including humans, mice and flies. [10]
There are blue light photoreceptors that are not a part of photomorphogenesis. For example, phototropin is the blue light photoreceptor that controls phototropism.
Plants show various responses to UV light. UVR8 has been shown to be a UV-B receptor. [11] Plants undergo distinct photomorphogenic changes as a result of UV-B radiation. They have photoreceptors that initiate morphogenetic changes in the plant embryo (hypocotyl, epicotyl, radicle) [12] Exposure to UV- light in plants mediates biochemical pathways, photosynthesis, plant growth and many other processes essential to plant development. The UV-B photoreceptor, UV Resistance Locus8 (UVR8) detects UV-B rays and elicits photomorphogenic responses. These response are important for initiating hypocotyl elongation, leaf expansion, biosynthesis of flavonoids and many other important processes that affect the root-shoot system. [13] Exposure to UV-B rays can be damaging to DNA inside of the plant cells, however, UVR8 induces genes required to acclimate plants to UV-B radiation, these genes are responsible for many biosynthesis pathways that involve protection from UV damage, oxidative stress, and photorepair of DNA damage. [14]
There is still much to be discovered about the mechanisms involved in UV-B radiation and UVR8. Scientists are working to understand the pathways responsible for plant UV receptors response to solar radiation in natural environments. [14]
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.
Photobiology is the scientific study of the beneficial and harmful interactions of light in living organisms. The field includes the study of photophysics, photochemistry, photosynthesis, photomorphogenesis, visual processing, circadian rhythms, photomovement, bioluminescence, and ultraviolet radiation effects.
Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants.
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.
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.
Photoperiodism is the physiological reaction of organisms to the length of light or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light and dark periods. They are classified under three groups according to the photoperiods: short-day plants, long-day plants, and day-neutral plants.
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).
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.
Far-red light is a range of light at the extreme red end of the visible spectrum, just before infrared light. Usually regarded as the region between 700 and 750 nm wavelength, it is dimly visible to human eyes. It is largely reflected or transmitted by plants because of the absorbance spectrum of chlorophyll, and it is perceived by the plant photoreceptor phytochrome. However, some organisms can use it as a source of energy in photosynthesis. Far-red light also is used for vision by certain organisms such as some species of deep-sea fishes and mantis shrimp.
A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle, the hypocotyl, and the cotyledons. The two classes of flowering plants (angiosperms) are distinguished by their numbers of seed leaves: monocotyledons (monocots) have one blade-shaped cotyledon, whereas dicotyledons (dicots) possess two round cotyledons. Gymnosperms are more varied. For example, pine seedlings have up to eight cotyledons. The seedlings of some flowering plants have no cotyledons at all. These are said to be acotyledons.
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.
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. Most plant shoots exhibit positive phototropism, and rearrange their chloroplasts in the leaves to maximize photosynthetic energy and promote growth. 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.
UV-B resistance 8 (UVR8) also known as ultraviolet-B receptor UVR8 is an UV-B – sensing protein found in plants and possibly other sources. It is responsible for sensing ultraviolet light in the range 280-315 nm and initiating the plant stress response. It is most sensitive at 285nm, near the lower limit of UVB. UVR8 was first identified as a crucial mediator of a plant's response to UV-B in Arabidopsis thaliana containing a mutation in this protein. This plant was found to have a hypersensitivity to UV-B which damages DNA. UVR8 is thought to be a unique photoreceptor as it doesn't contain a prosthetic chromophore but its light-sensing ability is intrinsic to the molecule. Tryptophan (Trp) residue 285 has been suggested to act the UV-B sensor, while other Trp residues have been also seen to be involved although in-vivo data suggests that Trp285 and Trp233 are most important.
Steve A. Kay is a British-born chronobiologist who mainly works in the United States. Dr. Kay has pioneered methods to monitor daily gene expression in real time and characterized circadian gene expression in plants, flies and mammals. In 2014, Steve Kay celebrated 25 years of successful chronobiology research at the Kaylab 25 Symposium, joined by over one hundred researchers with whom he had collaborated with or mentored. Dr. Kay, a member of the National Academy of Sciences, U.S.A., briefly served as president of The Scripps Research Institute. and is currently a professor at the University of Southern California. He also served on the Life Sciences jury for the Infosys Prize in 2011.
Takuma Akuma Tanada was an American plant biologist who made several discoveries related to the effects of light radiation on plants, including his discovery of the Tanada effect. He conducted research at the United States Department of Agriculture and in 2011 was awarded a Congressional Gold Medal, the highest civilian award in the United States, for his assistance to the U.S. military in World War II.
Aureochromes are blue light photoreceptors as well as transcription factors found only in stramenopiles so far.
Dmitri Nusinow is an American chronobiologist who studies plant circadian rhythms. He was born on November 7, 1976, in Inglewood, California. He currently resides in St. Louis, and his research focus includes a combination of molecular, biochemical, genetic, genomic, and proteomic tools to discover the molecular connections between signaling networks, circadian oscillators, and specific outputs. By combining these methods, he hopes to apply the knowledge elucidated from the Arabidopsis model to other plant species.
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.