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r infrared are within the range of 0 5 to 20 nanoseconds the fluorescence lifetime is an important parameter for practical applications of fluorescence such as fluorescence resonance energy transfer and fluorescence lifetime imaging microscopy jablonski diagram edit the jablonski diagram describes most of the relaxation mechanisms for excited state molecules the diagram alongside shows how fluorescence occurs due to the relaxation of certain excited electrons of a molecule 19 fluorescence anisotropy edit fluorophores are more likely to be excited by photons if the transition moment of the fluorophore is parallel to the electric vector of the photon 15 pp12 13 the polarization of the emitted light will also depend on the transition moment the transition moment is dependent on the physical orientation of the fluorophore molecule for fluorophores in solution the intensity and polarization of the emitted light is dependent on rotational diffusion therefore anisotropy measurements can be used to investigate how freely a fluorescent molecule moves in a particular environment fluorescence anisotropy can be defined quantitatively as r i i i 2 i displaystyle r i_ parallel i_ perp over i_ parallel 2i_ perp where i displaystyle i_ parallel is the emitted intensity parallel to the polarization of the excitation light and i displaystyle i_ perp is the emitted intensity perpendicular to the polarization of the excitation light 13 anisotropy is independent of the intensity of the absorbed or emitted light it is a property of the light so photobleaching of the dye will not affect the anisotropy value as long as the signal is detectable fluorescence edit fluorescent security strip in a us twenty dollar bill under uv light strongly fluorescent pigments often have an unusual appearance which is often described colloquially as a neon color originally day glo in the late 1960s early 1970s this phenomenon was termed farbenglut by hermann von helmholtz and fluorence by ralph m evans it is generally thought to be related to the high brightness of the color relative to what it would be as a component of white fluorescence shifts energy in the incident illumination from shorter wavelengths to longer such as blue to yellow and thus can make the fluorescent color appear brighter more saturated than it could possibly be by reflection alone 20 rules edit there are several general rules that deal with fluorescence each of the following rules have exceptions but they are useful guidelines for understanding fluorescence these rules do not necessarily apply to two photon absorption kasha s rule edit kasha s rule states that the luminesce fluorescence or phosphorescence of a molecule will be emitted only from the lowest excited state of its given multiplicity 21 vavilov s rule a logical extension of kasha s rule thusly called kasha vavilov rule dictates that the quantum yield of luminescence is independent of the wavelength of exciting radiation and is proportional to the absorbance of the excited wavelength 22 kasha s rule does not always apply and is violated by simple molecules such an example is azulene 23 a somewhat more reliable statement although still with exceptions would be that the fluorescence spectrum shows very little dependence on the wavelength of exciting radiation 24 mirror image rule edit the fluorescent dye rhodamine 6g is commonly used in applications such as highlighter pens dye lasers and automotive leak detection the absorption profile is a mirror of the emission profile for many fluorophores the absorption spectrum is a mirror image of the emission spectrum 15 pp6 8 this is known as the mirror image rule and is related to the franck condon principle which states that electronic transitions are vertical that is energy changes without distance changing as can be represented with a vertical line in jablonski diagram this means the nucleus does not move and the vibration levels of the excited state resemble the vibration levels of the ground state stokes shift edit main article stokes shift in general emitted fluorescence light has a longer wavelength and lower energy than the absorbed light 15 pp6 7 this phenomenon known as stokes shift is due to energy loss between the time a photon is absorbed and when a new one is emitted the causes and magnitude of stokes shift can be complex and are dependent on the fluorophore and its environment however there are some common causes it is frequently due to non radiative decay to the lowest vibrational energy level of the excited state another factor is that the emission of fluorescence frequently leaves a fluorophore in a higher vibrational level of the ground state in nature edit fluorescent coral main article fluorescence in the life sciences there are many natural compounds that exhibit fluorescence and they have a number of applications some deep sea animals such as the greeneye have fluorescent structures fluorescence edit when fluorescence occurs in a living organism it is sometimes called biofluorescence fluorescence is distinct from bioluminescence and biophosphorescence 25 pumpkin toadlets that live in the brazilian atlantic forest are fluorescent 26 bioluminescence edit main article bioluminescence bioluminescence differs from fluorescence in that it is the natural production of light by chemical reactions within an organism whereas fluorescence is the absorption and reemission of light from the environment 25 fireflies and anglerfish are two examples of bioluminescent organisms 27 some organisms are both bioluminescent and fluorescent like the sea pansy renilla reniformis where bioluminescence serves as the light source for fluorescence 28 phosphorescence edit main article phosphorescence phosphorescence is similar to fluorescence in its requirement of light wavelengths as a provider of excitation energy the difference here lies in the relative stability of the energized molecule unlike with fluorescence in phosphorescence the energized molecule is metastable emitting light that continues to glow in the dark even after the stimulating light source has been removed 25 for example glow in the dark stickers are phosphorescent citation needed biophosphorescence in living organisms is exceptionally rare only reported in a few millipedes 29 mechanisms edit epidermal chromatophores edit pigment cells that exhibit fluorescence are called fluorescent chromatophores and function somatically similar to regular chromatophores these cells are dendritic and contain pigments called fluorosomes these pigments contain fluorescent proteins which are activated by k potassium ions and it is their movement aggregation and dispersion within the fluorescent chromatophore that cause directed fluorescence patterning 30 31 fluorescent cells are innervated the same as other chromatophores like melanophores pigment cells that contain melanin short term fluorescent patterning and signaling is controlled by the nervous system 30 fluorescent chromatophores can be found in the skin e g in fish just below the epidermis amongst other chromatophores epidermal fluorescent cells in fish also respond to hormonal stimuli by the α msh and mch hormones much the same as melanophores this suggests that fluorescent cells may have color changes throughout the day that coincide with their circadian rhythm 32 fish may also be sensitive to cortisol induced stress responses to environmental stimuli such as interaction with a predator or engaging in a mating ritual 30 phylogenetics edit evolutionary origins edit the incidence of fluorescence across the tree of life is widespread and has been studied most extensively in cnidarians and fish the phenomenon appears to have evolved multiple times in multiple taxa such as in the anguilliformes eels gobioidei gobies and cardinalfishes and tetradontiformes triggerfishes along with the other taxa discussed later in the article fluorescence is highly genotypically and phenotypically variable even within ecosystems in regards to the wavelengths emitted the patterns displayed and the intensity of the fluorescence generally the species relying upon camouflage exhibit the greatest diversity in fluorescence likely because camouflage may be one of the uses of fluorescence 33 fluorescence has multiple origins in the tree of life this diagram displays the origins within actinopterygians ray finned fish it is suspected by some scientists that gfps and gfp like proteins began as electron donors activated by light these electrons were then used for reactions requiring light energy functions of fluorescent proteins such as protection from the sun conversion of light into different wavelengths or for signaling are thought to have evolved secondarily 34 adaptive functions edit currently relatively little is known about the functional significance of fluorescence and fluorescent proteins 34 however it is suspected that fluorescence may serve important functions in signaling and communication mating lures camouflage uv protection and antioxidation photoacclimation dinoflagellate regulation and in coral health 35 aquatic edit water absorbs light of long wavelengths so less light from these wavelengths reflects back to reach the eye therefore warm colors from the visual light spectrum appear less vibrant at increasing depths water scatters light of shorter wavelengths above violet meaning cooler colors dominate the visual field in the photic zone light intensity decreases 10 fold with every 75 m of depth so at depths of 75 m light is 10 as intense as it is on the surface and is only 1 as intense at 150 m as it is on the surface because the water filters out the wavelengths and intensity of water reaching certain depths different proteins because of the wavelengths and intensities of light they are capable of absorbing are better suited to different depths theoretically some fish eyes can detect light as deep as 1000 m at these depths of the aphotic zone the only sources of light are organisms themselves giving off light through chemical reactions in a process called bioluminescence fluorescence is simply defined as the absorption of electromagnetic radiation at one wavelength and its reemission at another lower energy wavelength 33 thus any type of fluorescence depends on the presence of external sources of light biologically functional fluorescence is found in the photic zone where there is not only enough light to cause fluorescence but enough light for other organisms to detect it 36 the visual field in the photic zone is naturally blue so colors of fluorescence can be detected as bright reds oranges yellows and greens green is the most commonly found color in the marine spectrum yellow the second most orange the third and red is the rarest fluorescence can occur in organisms in the aphotic zone as a byproduct of that same organism s bioluminescence some fluorescence in the aphotic zone is merely a byproduct of the organism s tissue biochemistry and does not have a functional purpose however some cases of functional and adaptive significance of fluorescence in the aphotic zone of the deep ocean is an active area of research 37 photic zone edit main article photic zone fish edit fluorescent marine fish bony fishes living in shallow water generally have good color vision due to their living in a colorful environment thus in shallow water fishes red orange and green fluorescence most likely serves as a means of communication with conspecifics especially given the great phenotypic variance of the phenomenon 33 many fish that exhibit fluorescence such as sharks lizardfish scorpionfish wrasses and flatfishes also possess yellow intraocular filters 38 yellow intraocular filters in the lenses and cornea of certain fishes function as long pass filters these filters enable the species to visualize and potentially exploit fluorescence in order to enhance visual contrast and patterns that are unseen to other fishes and predators that lack this visual specialization 33 fish that possess the necessary yellow intraocular filters for visualizing fluorescence potentially exploit a light signal from members of it fluorescent patterning was especially prominent in cryptically patterned fishes possessing complex camouflage many of these lineages also possess yellow long pass intraocular filters that could enable visualization of such patterns 38 another adaptive use of fluorescence is to generate orange and red light from the ambient blue light of the photic zone to aid vision red light can only be seen across short distances due to attenuation of red light wavelengths by water 39 many fish species that fluoresce are small group living or benthic aphotic and have conspicuous patterning this patterning is caused by fluorescent tissue and is visible to other members of the species however the patterning is invisible at other visual spectra these intraspecific fluorescent patterns also coincide with intra species signaling the patterns present in ocular rings to indicate directionality of an individual s gaze and along fins to indicate directionality of an individual s movement 39 current research suspects that this red fluorescence is used for private communication between members of the same species 30 33 39 due to the prominence of blue light at ocean depths red light and light of longer wavelengths are muddled and many predatory reef fish have little to no sensitivity for light at these wavelengths fish such as the fairy wrasse that have developed visual sensitivity to longer wavelengths are able to display red fluorescent signals that give a high contrast to the blue environment and are conspicuous to conspecifics in short ranges yet are relatively invisible to other common fish that have reduced sensitivities to long wavelengths thus fluorescence can be used as adaptive signaling and intra species communication in reef fish 39 40 additionally it is suggested that fluorescent tissues that surround an organism s eyes are used to convert blue light from the photic zone or green bioluminescence in the aphotic zone into red light to aid vision 39 sharks edit a new fluorophore was described in two species of sharks wherein it was due to an undescribed group of brominated tryptophane kynurenine small molecule metabolites 41 coral edit fluorescence serves a wide variety of functions in coral fluorescent proteins in corals may contribute to photosynthesis by converting otherwise unusable wavelengths of light into ones for which the coral s symbiotic algae are able to conduct photosynthesis 42 also the proteins may fluctuate in number as more or less light becomes available as a means of photoacclimation 43 similarly these fluorescent proteins may possess antioxidant capacities to eliminate oxygen radicals produced by photosynthesis 44 finally through modulating photosynthesis the fluorescent proteins may also serve as a means of regulating the activity of the coral s photosynthetic algal symbionts 45 cephalopods edit main ...
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