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Fluorescence is the emission of
light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 tera ...
by a substance that has absorbed light or other
electromagnetic radiation In physics, electromagnetic radiation (EMR) consists of waves of the electromagnetic field, electromagnetic (EM) field, which propagate through space and carry momentum and electromagnetic radiant energy. It includes radio waves, microwaves, inf ...
. It is a form of
luminescence Luminescence is spontaneous emission of light by a substance not resulting from heat; or "cold light". It is thus a form of cold-body radiation. It can be caused by chemical reactions, electrical energy, subatomic motions or stress on a cryst ...
. In most cases, the emitted light has a longer
wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tro ...
, and therefore a lower
photon A photon () is an elementary particle that is a quantum of the electromagnetic field, including electromagnetic radiation such as light and radio waves, and the force carrier for the electromagnetic force. Photons are massless, so they always ...
energy In physics, energy (from Ancient Greek: ἐνέργεια, ''enérgeia'', “activity”) is the quantitative property that is transferred to a body or to a physical system, recognizable in the performance of work and in the form of heat a ...
, than the absorbed radiation. A perceptible example of fluorescence occurs when the absorbed radiation is in the
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
region of the
electromagnetic spectrum The electromagnetic spectrum is the range of frequencies (the spectrum) of electromagnetic radiation and their respective wavelengths and photon energies. The electromagnetic spectrum covers electromagnetic waves with frequencies ranging from ...
(invisible to the human eye), while the emitted light is in the
visible region The visible spectrum is the portion of the electromagnetic spectrum that is visual perception, visible to the human eye. Electromagnetic radiation in this range of wavelengths is called ''visible light'' or simply light. A typical human eye wil ...
; this gives the fluorescent substance a distinct
color Color (American English) or colour (British English) is the visual perceptual property deriving from the spectrum of light interacting with the photoreceptor cells of the eyes. Color categories and physical specifications of color are associ ...
that can only be seen when the substance has been exposed to
UV light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...
. Fluorescent materials cease to glow nearly immediately when the radiation source stops, unlike
phosphorescent Phosphorescence is a type of photoluminescence related to fluorescence. When exposed to light (radiation) of a shorter wavelength, a phosphorescent substance will glow, absorbing the light and reemitting it at a longer wavelength. Unlike fluor ...
materials, which continue to emit light for some time after. Fluorescence has many practical applications, including
mineralogy Mineralogy is a subject of geology specializing in the scientific study of the chemistry, crystal structure, and physical (including optical) properties of minerals and mineralized artifacts. Specific studies within mineralogy include the proces ...
,
gemology Gemology or gemmology is the science dealing with natural and artificial gemstone materials. It is a geoscience and a branch of mineralogy. Some jewelers (and many non-jewelers) are academically trained gemologists and are qualified to identify a ...
,
medicine Medicine is the science and practice of caring for a patient, managing the diagnosis, prognosis, prevention, treatment, palliation of their injury or disease, and promoting their health. Medicine encompasses a variety of health care pract ...
, chemical sensors (
fluorescence spectroscopy Fluorescence spectroscopy (also known as fluorimetry or spectrofluorometry) is a type of electromagnetic spectroscopy that analyzes fluorescence from a sample. It involves using a beam of light, usually ultraviolet light, that excites the electron ...
),
fluorescent labelling In molecular biology and biotechnology, a fluorescent tag, also known as a fluorescent label or fluorescent probe, is a molecule that is attached chemically to aid in the detection of a biomolecule such as a protein, antibody, or amino acid. Gener ...
,
dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...
s, biological detectors, cosmic-ray detection,
vacuum fluorescent display A vacuum fluorescent display (VFD) is a display device once commonly used on consumer electronics equipment such as video cassette recorders, car radios, and microwave ovens. A VFD operates on the principle of cathodoluminescence, roughly s ...
s, and
cathode-ray tube A cathode-ray tube (CRT) is a vacuum tube containing one or more electron guns, which emit electron beams that are manipulated to display images on a phosphorescent screen. The images may represent electrical waveforms (oscilloscope), pictur ...
s. Its most common everyday application is in ( gas-discharge)
fluorescent lamp A fluorescent lamp, or fluorescent tube, is a low-pressure mercury-vapor gas-discharge lamp that uses fluorescence to produce visible light. An electric current in the gas excites mercury vapor, which produces short-wave ultraviolet lig ...
s and
LED lamps An LED lamp or LED light bulb is an electric light that produces light using light-emitting diodes (LEDs). LED lamps are significantly more energy-efficient than equivalent incandescent lamps and can be significantly more efficient than mos ...
, in which fluorescent coatings convert UV or blue light into longer-wavelengths resulting in white light which can even appear indistinguishable from that of the traditional but energy- inefficient
incandescent lamp An incandescent light bulb, incandescent lamp or incandescent light globe is an electric light with a wire filament heated until it glows. The filament is enclosed in a glass bulb with a vacuum or inert gas to protect the filament from oxid ...
. Fluorescence also occurs frequently in nature in some minerals and in many biological forms across all kingdoms of life. The latter may be referred to as ''biofluorescence'', indicating that the
fluorophore A fluorophore (or fluorochrome, similarly to a chromophore) is a fluorescent chemical compound that can re-emit light upon light excitation. Fluorophores typically contain several combined aromatic groups, or planar or cyclic molecules with se ...
is part of or is extracted from a living organism (rather than an inorganic
dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...
or
stain A stain is a discoloration that can be clearly distinguished from the surface, material, or medium it is found upon. They are caused by the chemical or physical interaction of two dissimilar materials. Accidental staining may make materials app ...
). But since fluorescence is due to a specific chemical, which can also be synthesized artificially in most cases, it is sufficient to describe the substance itself as ''fluorescent''.


History

An early observation of fluorescence was described in 1560 by
Bernardino de Sahagún Bernardino de Sahagún, OFM (; – 5 February 1590) was a Franciscan friar, missionary priest and pioneering ethnographer who participated in the Catholic evangelization of colonial New Spain (now Mexico). Born in Sahagún, Spain, in 1499, he ...
and in 1565 by
Nicolás Monardes Nicolás Bautista Monardes (1493 – 10 October 1588) was a Spanish physician and botanist. Monardes published several books of varying importance. In ''Diálogo llamado pharmacodilosis'' (1536), he examines humanism and suggests studying s ...
in the
infusion Infusion is the process of extracting chemical compounds or flavors from plant material in a solvent such as water, oil or alcohol, by allowing the material to remain suspended in the solvent over time (a process often called steeping). An inf ...
known as ''
lignum nephriticum ''Lignum nephriticum'' (Latin for "kidney wood") is a traditional diuretic that was derived from the wood of two tree species, the narra (''Pterocarpus indicus'') and the Mexican kidneywood (''Eysenhardtia polystachya''). The wood is capable of t ...
'' (
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for "kidney wood"). It was derived from the wood of two tree species, ''
Pterocarpus indicus ''Pterocarpus indicus'' (commonly known as Amboyna wood, Malay padauk, Papua New Guinea rosewood, Philippine mahogany, Andaman redwood, Burmese rosewood, narra and asana in the Philippines, angsana, or Pashu padauk) is a species of ''Pterocarpus ...
'' and ''
Eysenhardtia polystachya ''Eysenhardtia polystachya'', the kidneywood, is a tree from Mexico, growing along forest edges and water courses at elevations of 150–3000 m. Previously it was used as a source of lignum nephriticum. References * polystachya ...
''. The chemical compound responsible for this fluorescence is matlaline, which is the oxidation product of one of the
flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
s found in this wood. In 1819, E.D. Clarke and in 1822
René Just Haüy René Just Haüy () FRS MWS FRSE (28 February 1743 – 1 June 1822) was a French priest and mineralogist, commonly styled the Abbé Haüy after he was made an honorary canon of Notre Dame. Due to his innovative work on crystal structure and hi ...
described fluorescence in
fluorite Fluorite (also called fluorspar) is the mineral form of calcium fluoride, CaF2. It belongs to the halide minerals. It crystallizes in isometric cubic habit, although octahedral and more complex isometric forms are not uncommon. The Mohs sca ...
s,
Sir David Brewster Sir David Brewster KH PRSE FRS FSA Scot FSSA MICE (11 December 178110 February 1868) was a British scientist, inventor, author, and academic administrator. In science he is principally remembered for his experimental work in physical optics ...
described the phenomenon for
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
in 1833 and
Sir John Herschel Sir John Frederick William Herschel, 1st Baronet (; 7 March 1792 – 11 May 1871) was an English polymath active as a mathematician, astronomer, chemist, inventor, experimental photographer who invented the blueprint and did botanical wor ...
did the same for
quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg cr ...
in 1845. In his 1852 paper on the "Refrangibility" (
wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tro ...
change) of light,
George Gabriel Stokes Sir George Gabriel Stokes, 1st Baronet, (; 13 August 1819 – 1 February 1903) was an Irish migration to Great Britain, Irish English physicist and mathematician. Born in County Sligo, Ireland, Stokes spent all of his career at the University ...
described the ability of fluorspar and
uranium glass Uranium glass is glass which has had uranium, usually in oxide diuranate form, added to a glass mix before melting for colouration. The proportion usually varies from trace levels to about 2% uranium by weight, although some 20th-century piec ...
to change invisible light beyond the violet end of the visible spectrum into blue light. He named this phenomenon ''fluorescence'' : "I am almost inclined to coin a word, and call the appearance ''fluorescence'', from fluor-spar .e., fluorite as the analogous term ''opalescence'' is derived from the name of a mineral." The name was derived from the mineral
fluorite Fluorite (also called fluorspar) is the mineral form of calcium fluoride, CaF2. It belongs to the halide minerals. It crystallizes in isometric cubic habit, although octahedral and more complex isometric forms are not uncommon. The Mohs sca ...
(calcium difluoride), some examples of which contain traces of divalent
europium Europium is a chemical element with the symbol Eu and atomic number 63. Europium is the most reactive lanthanide by far, having to be stored under an inert fluid to protect it from atmospheric oxygen or moisture. Europium is also the softest lanth ...
, which serves as the fluorescent activator to emit blue light. In a key experiment he used a prism to isolate ultraviolet radiation from sunlight and observed blue light emitted by an ethanol solution of quinine exposed by it.


Physical principles


Mechanism

Fluorescence occurs when an excited molecule, atom, or
nanostructure A nanostructure is a structure of intermediate size between microscopic and molecular structures. Nanostructural detail is microstructure at nanoscale. In describing nanostructures, it is necessary to differentiate between the number of dimens ...
, relaxes to a lower energy state (usually the
ground state The ground state of a quantum-mechanical system is its stationary state of lowest energy; the energy of the ground state is known as the zero-point energy of the system. An excited state is any state with energy greater than the ground state. ...
) through emission of a
photon A photon () is an elementary particle that is a quantum of the electromagnetic field, including electromagnetic radiation such as light and radio waves, and the force carrier for the electromagnetic force. Photons are massless, so they always ...
without a change in
electron spin In atomic physics, the electron magnetic moment, or more specifically the electron magnetic dipole moment, is the magnetic moment of an electron resulting from its intrinsic properties of spin (physics), spin and electric charge. The value of the ...
. When the initial and final states have different multiplicity (spin), the phenomenon is termed
phosphorescence Phosphorescence is a type of photoluminescence related to fluorescence. When exposed to light (radiation) of a shorter wavelength, a phosphorescent substance will glow, absorbing the light and reemitting it at a longer wavelength. Unlike fluo ...
. The ground state of most molecules is a
singlet state In quantum mechanics, a singlet state usually refers to a system in which all electrons are paired. The term 'singlet' originally meant a linked set of particles whose net angular momentum is zero, that is, whose overall spin quantum number s=0. A ...
, denoted as S0. A notable exception is
molecular oxygen There are several known allotropes of oxygen. The most familiar is molecular oxygen (O2), present at significant levels in Earth's atmosphere and also known as dioxygen or triplet oxygen. Another is the highly reactive ozone (O3). Others are: *A ...
, which has a triplet ground state. Absorption of a photon of energy h \nu_ results in an excited state of the same multiplicity (spin) of the ground state, usually a singlet (Sn with n > 0). In solution, states with n > 1 relax rapidly to the lowest vibrational level of the first excited state (S1) by transferring energy to the solvent molecules through non-radiative processes, including internal conversion followed by vibrational relaxation, in which the energy is dissipated as
heat In thermodynamics, heat is defined as the form of energy crossing the boundary of a thermodynamic system by virtue of a temperature difference across the boundary. A thermodynamic system does not ''contain'' heat. Nevertheless, the term is al ...
. Therefore, most commonly, fluorescence occurs from the first singlet excited state, S1. Fluorescence is the emission of a photon accompanying the relaxation of the excited state to the ground state. Fluorescence photons are lower in energy (h \nu_ ) compared to the energy of the photons used to generate the excited state (h \nu_ ) * Excitation: \mathrm_0 + h \nu_ \to \mathrm_1 * Fluorescence (emission): \mathrm_1 \to \mathrm_0 + h \nu_ In each case the photon energy E is proportional to its
frequency Frequency is the number of occurrences of a repeating event per unit of time. It is also occasionally referred to as ''temporal frequency'' for clarity, and is distinct from ''angular frequency''. Frequency is measured in hertz (Hz) which is eq ...
\nu according to E=h\nu, where h is Planck's constant. The excited state S1 can relax by other mechanisms that do not involve the emission of light. These processes, called non-radiative processes, compete with fluorescence emission and decrease its efficiency. Examples include
internal conversion Internal conversion is a non-radioactive, atomic decay process where an excited nucleus interacts electromagnetically with one of the orbital electrons of an atom. This causes the electron to be emitted (ejected) from the atom. Thus, in internal ...
,
intersystem crossing Intersystem crossing (ISC) is an isoenergetic radiationless process involving a transition between the two electronic states with different spin multiplicity. Excited Singlet and Triplet States When an electron in a molecule with a singlet ground ...
to the triplet state, and energy transfer to another molecule. An example of energy transfer is
Förster resonance energy transfer Förster resonance energy transfer (FRET), fluorescence resonance energy transfer, resonance energy transfer (RET) or electronic energy transfer (EET) is a mechanism describing energy transfer between two light-sensitive molecules ( chromophores). ...
. Relaxation from an excited state can also occur through collisional
quenching In materials science, quenching is the rapid cooling of a workpiece in water, oil, polymer, air, or other fluids to obtain certain material properties. A type of heat treating, quenching prevents undesired low-temperature processes, such as pha ...
, a process where a molecule (the quencher) collides with the fluorescent molecule during its excited state lifetime. Molecular
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
(O2) is an extremely efficient quencher of fluorescence just because of its unusual triplet ground state.


Quantum yield

The fluorescence
quantum yield The quantum yield (Φ) of a radiation-induced process is the number of times a specific event occurs per photon absorbed by the system. Applications Fluorescence spectroscopy The fluorescence quantum yield is defined as the ratio of the numb ...
gives the efficiency of the fluorescence process. It is defined as the ratio of the number of photons emitted to the number of photons absorbed. : \Phi = \frac The maximum possible fluorescence quantum yield is 1.0 (100%); each
photon A photon () is an elementary particle that is a quantum of the electromagnetic field, including electromagnetic radiation such as light and radio waves, and the force carrier for the electromagnetic force. Photons are massless, so they always ...
absorbed results in a photon emitted. Compounds with quantum yields of 0.10 are still considered quite fluorescent. Another way to define the quantum yield of fluorescence is by the rate of excited state decay: : \Phi = \frac where _ is the rate constant of
spontaneous emission Spontaneous emission is the process in which a quantum mechanical system (such as a molecule, an atom or a subatomic particle) transits from an excited energy state to a lower energy state (e.g., its ground state) and emits a quantized amount of ...
of radiation and : \sum__ is the sum of all rates of excited state decay. Other rates of excited state decay are caused by mechanisms other than photon emission and are, therefore, often called "non-radiative rates", which can include: * dynamic collisional quenching * near-field dipole-dipole interaction (or
resonance energy transfer Resonance describes the phenomenon of increased amplitude that occurs when the frequency of an applied Periodic function, periodic force (or a Fourier analysis, Fourier component of it) is equal or close to a natural frequency of the system ...
) * internal conversion *
intersystem crossing Intersystem crossing (ISC) is an isoenergetic radiationless process involving a transition between the two electronic states with different spin multiplicity. Excited Singlet and Triplet States When an electron in a molecule with a singlet ground ...
Thus, if the rate of any pathway changes, both the excited state lifetime and the fluorescence quantum yield will be affected. Fluorescence quantum yields are measured by comparison to a standard. The
quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg cr ...
salt ''quinine sulfate'' in a
sulfuric acid Sulfuric acid (American spelling and the preferred IUPAC name) or sulphuric acid ( Commonwealth spelling), known in antiquity as oil of vitriol, is a mineral acid composed of the elements sulfur, oxygen and hydrogen, with the molecular formu ...
solution was regarded as the most common fluorescence standard, however, a recent study revealed that the fluorescence quantum yield of this solution is strongly affected by the temperature, and should no longer be used as the standard solution. The quinine in 0.1  M perchloric acid (Φ=0.60) shows no temperature dependence up to 45°C, therefore it can be considered as a reliable standard solution.


Lifetime

The fluorescence lifetime refers to the average time the molecule stays in its excited state before emitting a photon. Fluorescence typically follows
first-order kinetics In chemistry, the rate law or rate equation for a reaction is an equation that links the initial or forward reaction rate with the concentrations or pressures of the reactants and constant parameters (normally rate coefficients and partial reactio ...
: : \left _1 \right= \left _1 \right0 e^ where \left _1 \right/math> is the concentration of excited state molecules at time t, \left _1 \right0 is the initial concentration and \Gamma is the decay rate or the inverse of the fluorescence lifetime. This is an instance of
exponential decay A quantity is subject to exponential decay if it decreases at a rate proportional to its current value. Symbolically, this process can be expressed by the following differential equation, where is the quantity and (lambda) is a positive rate ...
. Various radiative and non-radiative processes can de-populate the excited state. In such case the total decay rate is the sum over all rates: : \Gamma_=\Gamma_ + \Gamma_ where \Gamma_ is the total decay rate, \Gamma_ the radiative decay rate and \Gamma_ the non-radiative decay rate. It is similar to a first-order chemical reaction in which the first-order rate constant is the sum of all of the rates (a parallel kinetic model). If the rate of spontaneous emission, or any of the other rates are fast, the lifetime is short. For commonly used fluorescent compounds, typical excited state decay times for photon emissions with energies from the UV to
near infrared Infrared (IR), sometimes called infrared light, is electromagnetic radiation (EMR) with wavelengths longer than those of visible light. It is therefore invisible to the human eye. IR is generally understood to encompass wavelengths from around ...
are within the range of 0.5 to 20
nanoseconds A nanosecond (ns) is a unit of time in the International System of Units (SI) equal to one billionth of a second, that is, of a second, or 10 seconds. The term combines the SI prefix ''nano-'' indicating a 1 billionth submultiple of an SI unit ( ...
. The fluorescence lifetime is an important parameter for practical applications of fluorescence such as
fluorescence resonance energy transfer Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, tha ...
and
fluorescence-lifetime imaging microscopy Fluorescence-lifetime imaging microscopy or FLIM is an imaging technique based on the differences in the exponential decay rate of the photon emission of a fluorophore from a sample. It can be used as an imaging technique in confocal microscopy, t ...
.


Jablonski diagram

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."Animation for the Principle of Fluorescence and UV-Visible Absorbance"
. ''PharmaXChange.info''.


Fluorescence anisotropy

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. 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 this means that 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 = where I_\parallel is the emitted intensity parallel to polarization of the excitation light and I_\perp is the emitted intensity perpendicular to the polarization of the excitation light.


Fluorescence

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 Hermann Ludwig Ferdinand von Helmholtz (31 August 1821 – 8 September 1894) was a German physicist and physician who made significant contributions in several scientific fields, particularly hydrodynamic stability. The Helmholtz Association, ...
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.


Rules

There are several general
rules Rule or ruling may refer to: Education * Royal University of Law and Economics (RULE), a university in Cambodia Human activity * The exercise of political or personal control by someone with authority or power * Business rule, a rule pert ...
that deal with fluorescence. Each of the following rules has exceptions but they are useful guidelines for understanding fluorescence (these rules do not necessarily apply to
two-photon absorption Two-photon absorption (TPA or 2PA) or two-photon excitation or non-linear absorption is the simultaneous absorption of two photons of identical or different frequencies in order to excite a molecule from one state (usually the ground state) to a hi ...
).


Kasha's rule

Kasha's rule dictates that the quantum yield of luminescence is independent of the wavelength of exciting radiation. This occurs because excited molecules usually decay to the lowest vibrational level of the excited state before fluorescence emission takes place. The Kasha–Vavilov rule does not always apply and is violated severely in many simple molecules. 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.


Mirror image rule

For many fluorophores the absorption spectrum is a mirror image of the emission spectrum.
This is known as the mirror image rule and is related to the
Franck–Condon principle The Franck–Condon principle (named for James Franck and Edward Condon) is a rule in spectroscopy and quantum chemistry that explains the intensity of vibronic transitions (the simultaneous changes in electronic and vibrational energy levels o ...
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

In general, emitted fluorescence light has a longer wavelength and lower energy than the absorbed light. This phenomenon, known as
Stokes shift __NOTOC__ Stokes shift is the difference (in energy, wavenumber or frequency units) between positions of the band maxima of the absorption and emission spectra (fluorescence and Raman being two examples) of the same electronic transition. It ...
, 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

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.


Compared to bioluminescence and biophosphorescence


Fluorescence

Fluorescence is the phenomenon of absorption of
electromagnetic In physics, electromagnetism is an interaction that occurs between particles with electric charge. It is the second-strongest of the four fundamental interactions, after the strong force, and it is the dominant force in the interactions of a ...
radiation, typically from ultraviolet or
visible light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 te ...
, by a molecule and the subsequent emission of a photon of a lower energy (smaller frequency, longer wavelength). This causes the light that is emitted to be a different color than the light that is absorbed. Stimulating light excites an
electron The electron ( or ) is a subatomic particle with a negative one elementary electric charge. Electrons belong to the first generation of the lepton particle family, and are generally thought to be elementary particles because they have no kn ...
to an excited state. When the molecule returns to the ground state, it releases a photon, which is the fluorescent emission. The excited state lifetime is short, so emission of light is typically only observable when the absorbing light is on. Fluorescence can be of any wavelength but is often more significant when emitted photons are in the visible spectrum. When it occurs in a living organism, it is sometimes called biofluorescence. Fluorescence should not be confused with bioluminescence and biophosphorescence. Pumpkin toadlets that live in the Brazilian Atlantic forest are fluorescent.


Bioluminescence

Bioluminescence Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some b ...
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.
Fireflies The Lampyridae are a family of elateroid beetles with more than 2,000 described species, many of which are light-emitting. They are soft-bodied beetles commonly called fireflies, lightning bugs, or glowworms for their conspicuous production ...
and
anglerfish The anglerfish are fish of the teleost order Lophiiformes (). They are bony fish named for their characteristic mode of predation, in which a modified luminescent fin ray (the esca or illicium) acts as a lure for other fish. The luminescence ...
are two examples of bioluminescent organisms. To add to the potential confusion, some organisms are both bioluminescent and fluorescent, like the sea pansy
Renilla reniformis The sea pansy, ''Renilla reniformis,'' is a species of colonial cnidarian in the family Renillidae, part of an octocoral subclass of Anthozoa that inhabit an expansive range of environments. It is native to warm continental shelf waters of th ...
, where bioluminescence serves as the light source for fluorescence.


Phosphorescence

Phosphorescence Phosphorescence is a type of photoluminescence related to fluorescence. When exposed to light (radiation) of a shorter wavelength, a phosphorescent substance will glow, absorbing the light and reemitting it at a longer wavelength. Unlike fluo ...
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 electron. Unlike with fluorescence, in phosphorescence the electron retains stability, emitting light that continues to "glow-in-the-dark" even after the stimulating light source has been removed. For example, glow-in-the-dark stickers are phosphorescent, but there are no truly ''biophosphorescent'' animals known.


Mechanisms


Epidermal chromatophores

Pigment cells that exhibit fluorescence are called fluorescent chromatophores, and function somatically similar to regular
chromatophore Chromatophores are cells that produce color, of which many types are Biological pigment, pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and ...
s. 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. Fluorescent cells are innervated the same as other chromatophores, like melanophores, pigment cells that contain
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
. Short term fluorescent patterning and signaling is controlled by the nervous system. 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 A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., Endogeny (biology), endogeno ...
. Fish may also be sensitive to
cortisol Cortisol is a steroid hormone, in the glucocorticoid class of hormones. When used as a medication, it is known as hydrocortisone. It is produced in many animals, mainly by the ''zona fasciculata'' of the adrenal cortex in the adrenal gland ...
induced
stress response The fight-or-flight or the fight-flight-or-freeze response (also called hyperarousal or the acute stress response) is a physiological reaction that occurs in response to a perceived harmful event, attack, or threat to survival. It was first des ...
s to environmental stimuli, such as interaction with a predator or engaging in a mating ritual.


Phylogenetics


Evolutionary origins

The incidence of fluorescence across the
tree of life The tree of life is a fundamental archetype in many of the world's mythological, religious, and philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The Assyrian Sacred Tree: A History ...
is widespread, and has been studied most extensively in cnidarians and fish. The phenomenon appears to have evolved multiple times in multiple
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
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. 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.


Adaptive functions

Currently, relatively little is known about the functional significance of fluorescence and fluorescent proteins. However, it is suspected that fluorescence may serve important functions in signaling and communication,
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
, lures,
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
,
UV protection Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...
and antioxidation, photoacclimation,
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
regulation, and in coral health.


Aquatic

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 In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tro ...
and its reemission at another, lower energy wavelength. 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. 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.


Photic zone


=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 Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
, especially given the great phenotypic variance of the phenomenon. Many fish that exhibit fluorescence, such as
sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...
,
lizardfish The Synodontidae or lizardfishes(or typical lizardfish to distinguish them from the Bathysauridae and Pseudotrichonotidae) are benthic (bottom-dwelling) marine and estuarine bony fishes that belong to the aulopiform fish order, a diverse group ...
,
scorpionfish The Scorpaenidae (also known as scorpionfish) are a family of mostly marine fish that includes many of the world's most venomous species. As their name suggests, scorpionfish have a type of "sting" in the form of sharp spines coated with venom ...
,
wrasses The wrasses are a family, Labridae, of marine fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into 9 subgroups or tribes. They are typically small, most of them le ...
, and flatfishes, also possess yellow intraocular filters. Yellow intraocular filters in the
lenses A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
and
cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical power ...
of certain fishes function as long-pass filters. These filters enable the species that 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. 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. 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. 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. Current research suspects that this red fluorescence is used for private communication between members of the same species. 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. 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.


=Sharks

= A new
fluorophore A fluorophore (or fluorochrome, similarly to a chromophore) is a fluorescent chemical compound that can re-emit light upon light excitation. Fluorophores typically contain several combined aromatic groups, or planar or cyclic molecules with se ...
was described in two species of sharks, wherein it was due to an undescribed group of brominated tryptophane-kynurenine small molecule metabolites.


=Coral

= 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 Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
. Also, the proteins may fluctuate in number as more or less light becomes available as a means of photoacclimation. Similarly, these fluorescent proteins may possess antioxidant capacities to eliminate oxygen radicals produced by photosynthesis. Finally, through modulating photosynthesis, the fluorescent proteins may also serve as a means of regulating the activity of the coral's photosynthetic algal symbionts.


=Cephalopods

= ''Alloteuthis subulata'' and ''Loligo vulgaris'', two types of nearly transparent squid, have fluorescent spots above their eyes. These spots reflect incident light, which may serve as a means of camouflage, but also for signaling to other squids for schooling purposes.


=Jellyfish

= Another, well-studied example of fluorescence in the ocean is the
hydrozoan Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specialize ...
''
Aequorea victoria ''Aequorea victoria'', also sometimes called the crystal jelly, is a bioluminescent hydrozoan jellyfish, or hydromedusa, that is found off the west coast of North America. The species is best known as the source of two proteins involved in biolu ...
''. This jellyfish lives in the photic zone off the west coast of North America and was identified as a carrier of
green fluorescent protein The green fluorescent protein (GFP) is a protein that exhibits bright green fluorescence when exposed to light in the blue to ultraviolet range. The label ''GFP'' traditionally refers to the protein first isolated from the jellyfish ''Aequorea ...
(GFP) by
Osamu Shimomura was a Japanese organic chemist and marine biologist, and Professor Emeritus at Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts and Boston University School of Medicine. He was awarded the Nobel Prize in Chemistry in 2008 for th ...
. The gene for these green fluorescent proteins has been isolated and is scientifically significant because it is widely used in genetic studies to indicate the expression of other genes.


=Mantis shrimp

= Several species of
mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in lengt ...
, which are stomatopod
crustaceans Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
, including ''Lysiosquillina glabriuscula'', have yellow fluorescent markings along their antennal scales and
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
(shell) that males present during threat displays to predators and other males. The display involves raising the head and thorax, spreading the striking appendages and other maxillipeds, and extending the prominent, oval antennal scales laterally, which makes the animal appear larger and accentuates its yellow fluorescent markings. Furthermore, as depth increases, mantis shrimp fluorescence accounts for a greater part of the visible light available. During mating rituals, mantis shrimp actively fluoresce, and the wavelength of this fluorescence matches the wavelengths detected by their eye pigments.


Aphotic zone


=Siphonophores

= ''
Siphonophorae Siphonophorae (from Greek ''siphōn'' 'tube' + ''pherein'' 'to bear') is an order within Hydrozoa, which is a class of marine organisms within the phylum Cnidaria. According to the World Register of Marine Species, the order contains 175 specie ...
'' is an order of marine animals from the phylum
Hydrozoa Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specializ ...
that consist of a specialized medusoid and polyp
zooid A zooid or zoöid is a single animal that is part of a colonial animal. This lifestyle has been adopted by animals from separate unrelated taxa. Zooids are multicellular; their structure is similar to that of other solitary animals. The zooi ...
. Some siphonophores, including the genus Erenna that live in the aphotic zone between depths of 1600 m and 2300 m, exhibit yellow to red fluorescence in the
photophores A photophore is a glandular organ that appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors, ...
of their tentacle-like
tentilla In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
. This fluorescence occurs as a by-product of bioluminescence from these same photophores. The siphonophores exhibit the fluorescence in a flicking pattern that is used as a lure to attract prey.


=Dragonfish

= The predatory deep-sea dragonfish ''Malacosteus niger'', the closely related genus ''
Aristostomias ''Aristostomias'' is a genus of barbeled dragonfishes native to the ocean depths in the Pacific, Atlantic and Indian oceans. Red Light Bioluminescence Production of red light bioluminescence Similar to other deep-sea organisms that are capa ...
'' and the species '' Pachystomias microdon'' use fluorescent red accessory pigments to convert the blue light emitted from their own bioluminescence to red light from suborbital
photophores A photophore is a glandular organ that appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors, ...
. This red luminescence is invisible to other animals, which allows these dragonfish extra light at dark ocean depths without attracting or signaling predators.


Terrestrial


Amphibians

Fluorescence is widespread among
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s and has been documented in several families of
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s,
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
s and
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
s, but the extent of it varies greatly. The
polka-dot tree frog The polka-dot tree frog (''Boana punctata''), also known as the dotted tree frog, is a frog species in the family (biology), family Hylidae found in much of South America and also in Trinidad and Tobago. It is fairly small with a snout–to–Clo ...
(''Hypsiboas punctatus''), widely found in South America, was unintentionally discovered to be the first fluorescent amphibian in 2017. The fluorescence was traced to a new compound found in the
lymph Lymph (from Latin, , meaning "water") is the fluid that flows through the lymphatic system, a system composed of lymph vessels (channels) and intervening lymph nodes whose function, like the venous system, is to return fluid from the tissues to ...
and skin glands. The main fluorescent compound is Hyloin-L1 and it gives a blue-green glow when exposed to violet or
ultraviolet light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...
. The scientists behind the discovery suggested that the fluorescence can be used for communication. They speculated that fluorescence possibly is relatively widespread among frogs. Only a few months later, fluorescence was discovered in the closely related ''
Hypsiboas atlanticus ''Boana atlantica'' is a small-sized species of tree frog in the family Hylidae. It is endemic to the Atlantic Forest from Pernambuco to Bahia in northeastern Brazil. It is a very abundant species inhabiting primary and secondary forest and for ...
''. Because it is linked to secretions from skin glands, they can also leave fluorescent markings on surfaces where they have been. In 2019, two other frogs, the tiny
pumpkin toadlet The pumpkin toadlet (''Brachycephalus ephippium''), or Spix's saddleback toad, is a small and brightly coloured species of frog in the family Brachycephalidae. This diurnal species is endemic to southeastern Brazil where it is found among leaf ...
(''Brachycephalus ephippium'') and
red pumpkin toadlet ''Brachycephalus pitanga'', the red pumpkin toadlet, is a small and brightly coloured species of anuran in the family Brachycephalidae. It is endemic to Atlantic rainforests in São Paulo state of southeastern Brazil, and only known from four lo ...
(''B. pitanga'') of southeastern Brazil, were found to be have naturally fluorescent skeletons, which is visible through their skin when exposed to ultraviolet light. It was initially speculated that the fluorescence supplemented their already
aposematic Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
colours (they are toxic) or that it was related to
mate choice Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior.Bateson, Paul Patrick Gordon. "Mate Choice." Mate Choic ...
(
species recognition Intra-species recognition is the recognition by a member of a species of a conspecific (another member of the same species). In many species, such recognition is necessary for procreation. Different species may employ different methods, but all ...
or determining fitness of a potential partner), but later studies indicate that the former explanation is unlikely, as predation attempts on the toadlets appear to be unaffected by the presence/absence of fluorescence. In 2020 it was confirmed that green or yellow fluorescence is widespread not only in adult frogs that are exposed to blue or ultraviolet light, but also among
tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found i ...
s, salamanders and caecilians. The extent varies greatly depending on species; in some it is highly distinct and in others it is barely noticeable. It can be based on their skin pigmentation, their mucous or their bones.


Butterflies

Swallowtail (''Papilio'') butterflies have complex systems for emitting fluorescent light. Their wings contain pigment-infused crystals that provide directed fluorescent light. These crystals function to produce fluorescent light best when they absorb
radiance In radiometry, radiance is the radiant flux emitted, reflected, transmitted or received by a given surface, per unit solid angle per unit projected area. Radiance is used to characterize diffuse emission and reflection of electromagnetic radiati ...
from sky-blue light (wavelength about 420 nm). The wavelengths of light that the butterflies see the best correspond to the absorbance of the crystals in the butterfly's wings. This likely functions to enhance the capacity for signaling.


Parrots

Parrots Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genus (biology), genera comprising the order (biology), order Psittaciformes (), found mostly in tropics, tropical and subtropics, subtropical regions. The order ...
have fluorescent
plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
that may be used in mate signaling. A study using mate-choice experiments on
budgerigars The budgerigar ( ; ''Melopsittacus undulatus''), also known as the common parakeet or shell parakeet, is a small, long-tailed, seed-eating parrot usually nicknamed the budgie ( ), or in American English, the parakeet. Budgies are the only spe ...
(''Melopsittacus undulates'') found compelling support for fluorescent sexual signaling, with both males and females significantly preferring birds with the fluorescent experimental stimulus. This study suggests that the fluorescent plumage of parrots is not simply a by-product of
pigmentation A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
, but instead an adapted sexual signal. Considering the intricacies of the pathways that produce fluorescent pigments, there may be significant costs involved. Therefore, individuals exhibiting strong fluorescence may be honest indicators of high individual quality, since they can deal with the associated costs.


Arachnids

Spiders fluoresce under UV light and possess a huge diversity of fluorophores. Remarkably, spiders are the only known group in which fluorescence is: : "taxonomically widespread, variably expressed, evolutionarily labile, and probably under selection and potentially of ecological importance for intraspecific and interspecific signaling". Andrews, Reed, & Masta showed that fluorescence evolved multiple times across spider taxa, with novel fluorophores evolving during spider diversification. In some spiders, ultraviolet cues are important for predator-prey interactions, intraspecific communication, and camouflage-matching with fluorescent flowers. Differing ecological contexts could favor inhibition or enhancement of fluorescence expression, depending upon whether fluorescence helps spiders be cryptic or makes them more conspicuous to predators. Therefore, natural selection could be acting on expression of fluorescence across spider species. Scorpions are also fluorescent, in their case due to the presence of
beta carboline β-Carboline (9''H''- pyrido ,4-''b'' ndole) represents the basic chemical structure for more than one hundred alkaloids and synthetic compounds. The effects of these substances depend on their respective substituent. Natural β-carbolines prima ...
in their cuticles.


Platypus

In 2020 fluorescence was reported for several platypus specimens.


Plants

Many plants are fluorescent due to the presence of
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
, which is probably the most widely-distributed fluorescent molecule, producing red emission under a range of excitation wavelengths. This attribute of chlorophyll is commonly used by ecologists to measure photosynthetic efficiency. The ''Mirabilis jalapa'' flower contains violet, fluorescent betacyanins and yellow, fluorescent betaxanthins. Under white light, parts of the flower containing only betaxanthins appear yellow, but in areas where both betaxanthins and betacyanins are present, the visible fluorescence of the flower is faded due to internal light-filtering mechanisms. Fluorescence was previously suggested to play a role in
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
attraction, however, it was later found that the visual signal by fluorescence is negligible compared to the visual signal of light reflected by the flower.


Abiotic


Gemology, mineralogy and geology

Gemstone A gemstone (also called a fine gem, jewel, precious stone, or semiprecious stone) is a piece of mineral crystal which, in cut and polished form, is used to make jewelry or other adornments. However, certain rocks (such as lapis lazuli, opal, ...
s,
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. ( ...
s, may have a distinctive fluorescence or may fluoresce differently under short-wave ultraviolet, long-wave ultraviolet, visible light, or
X-ray An X-ray, or, much less commonly, X-radiation, is a penetrating form of high-energy electromagnetic radiation. Most X-rays have a wavelength ranging from 10  picometers to 10  nanometers, corresponding to frequencies in the range 30&nb ...
s. Many types of
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
and
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
will fluoresce under shortwave UV, longwave UV and visible light.
Rubies A ruby is a pinkish red to blood-red colored gemstone, a variety of the mineral corundum ( aluminium oxide). Ruby is one of the most popular traditional jewelry gems and is very durable. Other varieties of gem-quality corundum are called sap ...
,
emerald Emerald is a gemstone and a variety of the mineral beryl (Be3Al2(SiO3)6) colored green by trace amounts of chromium or sometimes vanadium.Hurlbut, Cornelius S. Jr. and Kammerling, Robert C. (1991) ''Gemology'', John Wiley & Sons, New York, p ...
s, and
diamonds Diamond is a solid form of the element carbon with its atoms arranged in a crystal structure called diamond cubic. Another solid form of carbon known as graphite is the chemically stable form of carbon at room temperature and pressure, b ...
exhibit red fluorescence under long-wave UV, blue and sometimes green light; diamonds also emit light under
X-ray An X-ray, or, much less commonly, X-radiation, is a penetrating form of high-energy electromagnetic radiation. Most X-rays have a wavelength ranging from 10  picometers to 10  nanometers, corresponding to frequencies in the range 30&nb ...
radiation. Fluorescence in minerals is caused by a wide range of activators. In some cases, the concentration of the activator must be restricted to below a certain level, to prevent quenching of the fluorescent emission. Furthermore, the mineral must be free of impurities such as
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
or
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
, to prevent quenching of possible fluorescence. Divalent
manganese Manganese is a chemical element with the symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese is a transition metal with a multifaceted array of industrial alloy use ...
, in concentrations of up to several percent, is responsible for the red or orange fluorescence of
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
, the green fluorescence of
willemite Willemite is a zinc silicate mineral () and a minor ore of zinc. It is highly fluorescent (green) under shortwave ultraviolet light. It occurs in a variety of colors in daylight, in fibrous masses and apple-green gemmy masses. Troostite is a vari ...
, the yellow fluorescence of
esperite Esperite is a rare complex calcium lead zinc silicate (PbCa3Zn4(SiO4)4) related to beryllonite and trimerite that used to be called ''calcium larsenite''. Esperite has a white, greasy appearance in daylight and is much prized for its brilliant ...
, and the orange fluorescence of
wollastonite Wollastonite is a calcium inosilicate mineral ( Ca Si O3) that may contain small amounts of iron, magnesium, and manganese substituting for calcium. It is usually white. It forms when impure limestone or dolomite is subjected to high temperature ...
and
clinohedrite Clinohedrite is a rare silicate mineral. Its chemical composition is a hydrous calcium-zinc silicate; CaZn(SiO4)·H2O. It crystallizes in the monoclinic system and typically occurs as veinlets and fracture coatings. It is commonly colorless, wh ...
. Hexavalent
uranium Uranium is a chemical element with the symbol U and atomic number 92. It is a silvery-grey metal in the actinide series of the periodic table. A uranium atom has 92 protons and 92 electrons, of which 6 are valence electrons. Uranium is weak ...
, in the form of the uranyl cation (), fluoresces at all concentrations in a yellow green, and is the cause of fluorescence of minerals such as
autunite Autunite (hydrated calcium uranyl phosphate), with formula Ca(UO2)2(PO4)2·10–12H2O, is a yellow-greenish fluorescent phosphate mineral with a hardness of 2–. Autunite crystallizes in the orthorhombic system and often occurs as tabular square ...
or
andersonite Andersonite, Na2Ca(UO2)(CO3)3·6H2O, or hydrated sodium calcium uranyl carbonate is a rare uranium carbonate mineral that was first described in 1948. Named after Charles Alfred Anderson (1902–1990) of the United States Geological Survey, who ...
, and, at low concentration, is the cause of the fluorescence of such materials as some samples of
hyalite Hyalite is a transparent form of opal with a glassy lustre. It may exhibit an internal play of colors if natural inclusions are present. It is also called Muller's glass, water opal, and jalite. Müller's glass is named after its discoverer, Fr ...
opal Opal is a hydrated amorphous form of silica (SiO2·''n''H2O); its water content may range from 3 to 21% by weight, but is usually between 6 and 10%. Due to its amorphous property, it is classified as a mineraloid, unlike crystalline forms ...
. Trivalent
chromium Chromium is a chemical element with the symbol Cr and atomic number 24. It is the first element in group 6. It is a steely-grey, lustrous, hard, and brittle transition metal. Chromium metal is valued for its high corrosion resistance and hardne ...
at low concentration is the source of the red fluorescence of
ruby A ruby is a pinkish red to blood-red colored gemstone, a variety of the mineral corundum ( aluminium oxide). Ruby is one of the most popular traditional jewelry gems and is very durable. Other varieties of gem-quality corundum are called sa ...
. Divalent
europium Europium is a chemical element with the symbol Eu and atomic number 63. Europium is the most reactive lanthanide by far, having to be stored under an inert fluid to protect it from atmospheric oxygen or moisture. Europium is also the softest lanth ...
is the source of the blue fluorescence, when seen in the mineral
fluorite Fluorite (also called fluorspar) is the mineral form of calcium fluoride, CaF2. It belongs to the halide minerals. It crystallizes in isometric cubic habit, although octahedral and more complex isometric forms are not uncommon. The Mohs sca ...
. Trivalent
lanthanide The lanthanide () or lanthanoid () series of chemical elements comprises the 15 metallic chemical elements with atomic numbers 57–71, from lanthanum through lutetium. These elements, along with the chemically similar elements scandium and yttr ...
s such as
terbium Terbium is a chemical element with the symbol Tb and atomic number 65. It is a silvery-white, rare earth metal that is malleable, and ductile. The ninth member of the lanthanide series, terbium is a fairly electropositive metal that reacts with wa ...
and
dysprosium Dysprosium is the chemical element with the symbol Dy and atomic number 66. It is a rare-earth element in the lanthanide series with a metallic silver luster. Dysprosium is never found in nature as a free element, though, like other lanthanides, it ...
are the principal activators of the creamy yellow fluorescence exhibited by the yttrofluorite variety of the mineral fluorite, and contribute to the orange fluorescence of
zircon Zircon () is a mineral belonging to the group of nesosilicates and is a source of the metal zirconium. Its chemical name is zirconium(IV) silicate, and its corresponding chemical formula is Zr SiO4. An empirical formula showing some of the r ...
.
Powellite Powellite is a calcium molybdate mineral with formula CaMoO4. Powellite crystallizes with tetragonal - dipyramidal crystal structure as transparent adamantine blue, greenish brown, yellow to grey typically anhedral forms. It exhibits distinct cle ...
( calcium molybdate) and
scheelite Scheelite is a calcium tungstate mineral with the chemical formula Ca W O4. It is an important ore of tungsten (wolfram). Scheelite is originally named after Swedish chemist K. Scheele (1742-1786). Well-formed crystals are sought by collectors a ...
(calcium tungstate) fluoresce intrinsically in yellow and blue, respectively. When present together in
solid solution A solid solution, a term popularly used for metals, is a homogenous mixture of two different kinds of atoms in solid state and have a single crystal structure. Many examples can be found in metallurgy, geology, and solid-state chemistry. The word ...
, energy is transferred from the higher-energy
tungsten Tungsten, or wolfram, is a chemical element with the symbol W and atomic number 74. Tungsten is a rare metal found naturally on Earth almost exclusively as compounds with other elements. It was identified as a new element in 1781 and first isolat ...
to the lower-energy
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
, such that fairly low levels of
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
are sufficient to cause a yellow emission for
scheelite Scheelite is a calcium tungstate mineral with the chemical formula Ca W O4. It is an important ore of tungsten (wolfram). Scheelite is originally named after Swedish chemist K. Scheele (1742-1786). Well-formed crystals are sought by collectors a ...
, instead of blue. Low-iron
sphalerite Sphalerite (sometimes spelled sphaelerite) is a sulfide mineral with the chemical formula . It is the most important ore of zinc. Sphalerite is found in a variety of deposit types, but it is primarily in Sedimentary exhalative deposits, sedimen ...
(zinc sulfide), fluoresces and phosphoresces in a range of colors, influenced by the presence of various trace impurities. Crude oil (
petroleum Petroleum, also known as crude oil, or simply oil, is a naturally occurring yellowish-black liquid mixture of mainly hydrocarbons, and is found in geological formations. The name ''petroleum'' covers both naturally occurring unprocessed crud ...
) fluoresces in a range of colors, from dull-brown for heavy oils and tars through to bright-yellowish and bluish-white for very light oils and condensates. This phenomenon is used in
oil exploration Hydrocarbon exploration (or oil and gas exploration) is the search by petroleum geologists and geophysicists for deposits of hydrocarbons, particularly petroleum and natural gas, in the Earth using petroleum geology. Exploration methods Vis ...
drilling to identify very small amounts of oil in drill cuttings and core samples.
Humic acid Humic substances (HS) are organic compounds that are important components of humus, the major organic fraction of soil, peat, and coal (and also a constituent of many upland streams, dystrophic lakes, and ocean water). For a long era in the 19th ...
s and
fulvic acid Humic substances (HS) are organic compounds that are important components of humus, the major organic fraction of soil, peat, and coal (and also a constituent of many upland streams, dystrophic lakes, and ocean water). For a long era in the 19th an ...
s produced by the degradation of
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
in soils (
humus In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
) may also fluoresce because of the presence of aromatic cycles in their complex
molecular structure Molecular geometry is the three-dimensional arrangement of the atoms that constitute a molecule. It includes the general shape of the molecule as well as bond lengths, bond angles, torsional angles and any other geometrical parameters that deter ...
s. Humic substances dissolved in
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidate ...
can be detected and characterized by
spectrofluorimetry Fluorescence spectroscopy (also known as fluorimetry or spectrofluorometry) is a type of electromagnetic spectroscopy that analyzes fluorescence from a sample. It involves using a beam of light, usually ultraviolet light, that excites the electro ...
.


Organic liquids

Organic solutions such
anthracene Anthracene is a solid polycyclic aromatic hydrocarbon (PAH) of formula C14H10, consisting of three fused benzene rings. It is a component of coal tar. Anthracene is used in the Economic production, production of the red dye alizarin and other dyes ...
or
stilbene Stilbene may refer to one of the two stereoisomers of 1,2-diphenylethene: * (''E'')-Stilbene (''trans'' isomer) * (''Z'')-Stilbene (''cis'' isomer) See also * Stilbenoids, a class of molecules found in plants * 1,1-Diphenylethylene 1,1-Diphenyl ...
, dissolved in
benzene Benzene is an organic chemical compound with the molecular formula C6H6. The benzene molecule is composed of six carbon atoms joined in a planar ring with one hydrogen atom attached to each. Because it contains only carbon and hydrogen atoms, ...
or
toluene Toluene (), also known as toluol (), is a substituted aromatic hydrocarbon. It is a colorless, water-insoluble liquid with the smell associated with paint thinners. It is a mono-substituted benzene derivative, consisting of a methyl group (CH3) at ...
, fluoresce with
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
or
gamma ray A gamma ray, also known as gamma radiation (symbol γ or \gamma), is a penetrating form of electromagnetic radiation arising from the radioactive decay of atomic nuclei. It consists of the shortest wavelength electromagnetic waves, typically ...
irradiation Irradiation is the process by which an object is exposed to radiation. The exposure can originate from various sources, including natural sources. Most frequently the term refers to ionizing radiation, and to a level of radiation that will serve ...
. The decay times of this fluorescence are on the order of nanoseconds, since the duration of the light depends on the lifetime of the excited states of the fluorescent material, in this case anthracene or stilbene. Scintillation is defined a flash of light produced in a transparent material by the passage of a particle (an electron, an alpha particle, an ion, or a high-energy photon). Stilbene and derivatives are used in
scintillation counter A scintillation counter is an instrument for detecting and measuring ionizing radiation by using the excitation effect of incident radiation on a scintillating material, and detecting the resultant light pulses. It consists of a scintillator w ...
s to detect such particles. Stilbene is also one of the
gain medium The active laser medium (also called gain medium or lasing medium) is the source of optical gain within a laser. The gain results from the stimulated emission of photons through electronic or molecular transitions to a lower energy state from a ...
s used in
dye lasers A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution and ...
.


Atmosphere

Fluorescence is observed in the atmosphere when the air is under energetic electron bombardment. In cases such as the natural
aurora An aurora (plural: auroras or aurorae), also commonly known as the polar lights, is a natural light display in Earth's sky, predominantly seen in high-latitude regions (around the Arctic and Antarctic). Auroras display dynamic patterns of bri ...
, high-altitude nuclear explosions, and rocket-borne electron gun experiments, the molecules and ions formed have a fluorescent response to light.


Common materials that fluoresce

*
Vitamin B2 Riboflavin, also known as vitamin B2, is a vitamin found in food and sold as a dietary supplement. It is essential to the formation of two major coenzymes, flavin mononucleotide and flavin adenine dinucleotide. These coenzymes are involved in e ...
fluoresces yellow. *
Tonic water Tonic water (or Indian tonic water) is a carbonated soft drink in which quinine is dissolved. Originally used as a prophylactic against malaria, tonic water usually has a significantly lower quinine content and is consumed for its distinctive b ...
fluoresces blue due to the presence of
quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg cr ...
. *
Highlighter A highlighter is a type of writing device used to mark attention to sections of text by marking them with a vivid, translucent colour. A typical highlighter is fluorescent yellow, colored with pyranine. Different compounds, such as rhodamines ( ...
ink is often fluorescent due to the presence of
pyranine Pyranine is a hydrophilic, pH-sensitive fluorescent dye from the group of chemicals known as arylsulfonates. Pyranine is soluble in water and has applications as a coloring agent, biological stain, optical detecting reagent, and a pH indicator. On ...
. *
Banknote A banknote—also called a bill (North American English), paper money, or simply a note—is a type of negotiable instrument, negotiable promissory note, made by a bank or other licensed authority, payable to the bearer on demand. Banknotes w ...
s,
postage stamp A postage stamp is a small piece of paper issued by a post office, postal administration, or other authorized vendors to customers who pay postage (the cost involved in moving, insuring, or registering mail), who then affix the stamp to the fa ...
s and
credit card A credit card is a payment card issued to users (cardholders) to enable the cardholder to pay a merchant for goods and services based on the cardholder's accrued debt (i.e., promise to the card issuer to pay them for the amounts plus the o ...
s often have fluorescent security features.


In novel technology

In August 2020 researchers reported the creation of the brightest fluorescent solid optical materials so far by enabling the transfer of properties of highly fluorescent
dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...
s via spatial and electronic isolation of the dyes by mixing cationic dyes with anion-binding
cyanostar A cyanostar (pentacyanopentabenzo 5nnulene) is a shape-persistent macrocycle that binds anions. Synthesis The cyanostar structure is synthesized in a one-pot process among five equivalents of a benzaldehyde bearing a ''meta''- cyanomethyl subs ...
macrocycle Macrocycles are often described as molecules and ions containing a ring of twelve or more atoms. Classical examples include the crown ethers, calixarenes, porphyrins, and cyclodextrins. Macrocycles describe a large, mature area of chemistry. ...
s. According to a co-author these materials may have applications in areas such as solar energy harvesting, bioimaging, and lasers.


Applications


Lighting

The common
fluorescent lamp A fluorescent lamp, or fluorescent tube, is a low-pressure mercury-vapor gas-discharge lamp that uses fluorescence to produce visible light. An electric current in the gas excites mercury vapor, which produces short-wave ultraviolet lig ...
relies on fluorescence. Inside the
glass Glass is a non-crystalline, often transparent, amorphous solid that has widespread practical, technological, and decorative use in, for example, window panes, tableware, and optics. Glass is most often formed by rapid cooling (quenching) of ...
tube is a partial vacuum and a small amount of
mercury Mercury commonly refers to: * Mercury (planet), the nearest planet to the Sun * Mercury (element), a metallic chemical element with the symbol Hg * Mercury (mythology), a Roman god Mercury or The Mercury may also refer to: Companies * Merc ...
. An electric discharge in the tube causes the mercury atoms to emit mostly ultraviolet light. The tube is lined with a coating of a fluorescent material, called the ''
phosphor A phosphor is a substance that exhibits the phenomenon of luminescence; it emits light when exposed to some type of radiant energy. The term is used both for fluorescent or phosphorescent substances which glow on exposure to ultraviolet or vi ...
'', which absorbs ultraviolet light and re-emits visible light. Fluorescent
lighting Lighting or illumination is the deliberate use of light to achieve practical or aesthetic effects. Lighting includes the use of both artificial light sources like lamps and light fixtures, as well as natural illumination by capturing daylig ...
is more energy-efficient than
incandescent Incandescence is the emission of electromagnetic radiation (including visible light) from a hot body as a result of its high temperature. The term derives from the Latin verb ''incandescere,'' to glow white. A common use of incandescence is ...
lighting elements. However, the uneven
spectrum A spectrum (plural ''spectra'' or ''spectrums'') is a condition that is not limited to a specific set of values but can vary, without gaps, across a continuum. The word was first used scientifically in optics to describe the rainbow of colors i ...
of traditional fluorescent lamps may cause certain colors to appear different from when illuminated by incandescent light or
daylight Daylight is the combination of all direct and indirect sunlight during the daytime. This includes direct sunlight, diffuse sky radiation, and (often) both of these reflected by Earth and terrestrial objects, like landforms and buildings. Sunligh ...
. The mercury vapor emission spectrum is dominated by a short-wave UV line at 254 nm (which provides most of the energy to the phosphors), accompanied by visible light emission at 436 nm (blue), 546 nm (green) and 579 nm (yellow-orange). These three lines can be observed superimposed on the white continuum using a hand spectroscope, for light emitted by the usual white fluorescent tubes. These same visible lines, accompanied by the emission lines of trivalent europium and trivalent terbium, and further accompanied by the emission continuum of divalent europium in the blue region, comprise the more discontinuous light emission of the modern trichromatic phosphor systems used in many
compact fluorescent lamp A compact fluorescent lamp (CFL), also called compact fluorescent light, energy-saving light and compact fluorescent tube, is a fluorescent lamp designed to replace an incandescent light bulb; some types fit into light fixtures designed for inca ...
and traditional lamps where better color rendition is a goal. Fluorescent lights were first available to the public at the
1939 New York World's Fair The 1939–40 New York World's Fair was a world's fair held at Flushing Meadows–Corona Park in Queens, New York, United States. It was the second-most expensive American world's fair of all time, exceeded only by St. Louis's Louisiana Purchas ...
. Improvements since then have largely been better phosphors, longer life, and more consistent internal discharge, and easier-to-use shapes (such as compact fluorescent lamps). Some high-intensity discharge (HID) lamps couple their even-greater electrical efficiency with phosphor enhancement for better color rendition. White
light-emitting diode A light-emitting diode (LED) is a semiconductor device that emits light when current flows through it. Electrons in the semiconductor recombine with electron holes, releasing energy in the form of photons. The color of the light (cor ...
s (LEDs) became available in the mid-1990s as
LED lamp An LED lamp or LED light bulb is an electric light that produces light using light-emitting diodes (LEDs). LED lamps are significantly more energy-efficient than equivalent incandescent lamps and can be significantly more efficient than mos ...
s, in which blue light emitted from the
semiconductor A semiconductor is a material which has an electrical resistivity and conductivity, electrical conductivity value falling between that of a electrical conductor, conductor, such as copper, and an insulator (electricity), insulator, such as glas ...
strikes phosphors deposited on the tiny chip. The combination of the blue light that continues through the phosphor and the green to red fluorescence from the phosphors produces a net emission of white light.
Glow stick A glow stick, also known as a light stick, chem light, light wand, light rod, and rave light, is a self-contained, short-term light-source. It consists of a translucent plastic tube containing isolated substances that, when combined, make light ...
s sometimes utilize fluorescent materials to absorb light from the
chemiluminescent Chemiluminescence (also chemoluminescence) is the emission of light (luminescence) as the result of a chemical reaction. There may also be limited emission of heat. Given reactants A and B, with an excited intermediate ◊, : + -> lozenge - ...
reaction and emit light of a different color.


Analytical chemistry

Many analytical procedures involve the use of a
fluorometer A fluorometer, fluorimeter or fluormeter is a device used to measure parameters of visible spectrum fluorescence: its intensity and wavelength distribution of emission spectrum after excitation by a certain spectrum of light. These parameters ar ...
, usually with a single exciting wavelength and single detection wavelength. Because of the sensitivity that the method affords, fluorescent molecule concentrations as low as 1 part per trillion can be measured. Fluorescence in several wavelengths can be detected by an
array detector An array is a systematic arrangement of similar objects, usually in rows and columns. Things called an array include: {{TOC right Music * In twelve-tone and serial composition, the presentation of simultaneous twelve-tone sets such that the ...
, to detect compounds from
HPLC High-performance liquid chromatography (HPLC), formerly referred to as high-pressure liquid chromatography, is a technique in analytical chemistry used to separate, identify, and quantify each component in a mixture. It relies on pumps to pa ...
flow. Also,
TLC TLC may refer to: Arts and entertainment Television * ''TLC'' (TV series), a 2002 British situational comedy television series that aired on BBC2 * TLC (TV network), formerly the Learning Channel, an American cable TV network ** TLC (Asia), an A ...
plates can be visualized if the compounds or a coloring reagent is fluorescent. Fluorescence is most effective when there is a larger ratio of atoms at lower energy levels in a
Boltzmann distribution In statistical mechanics and mathematics, a Boltzmann distribution (also called Gibbs distribution Translated by J.B. Sykes and M.J. Kearsley. See section 28) is a probability distribution or probability measure that gives the probability t ...
. There is, then, a higher probability of excitement and release of photons by lower-energy atoms, making analysis more efficient.


Spectroscopy

Usually the setup of a fluorescence assay involves a light source, which may emit many different wavelengths of light. In general, a single wavelength is required for proper analysis, so, in order to selectively filter the light, it is passed through an excitation monochromator, and then that chosen wavelength is passed through the sample cell. After absorption and re-emission of the energy, many wavelengths may emerge due to
Stokes shift __NOTOC__ Stokes shift is the difference (in energy, wavenumber or frequency units) between positions of the band maxima of the absorption and emission spectra (fluorescence and Raman being two examples) of the same electronic transition. It ...
and various electron transitions. To separate and analyze them, the fluorescent radiation is passed through an emission
monochromator A monochromator is an optical device that transmits a mechanically selectable narrow band of wavelengths of light or other radiation chosen from a wider range of wavelengths available at the input. The name is from the Greek roots ''mono-'', "s ...
, and observed selectively by a detector.


Biochemistry and medicine

Fluorescence in the life sciences is used generally as a non-destructive way of tracking or analysis of biological molecules by means of the fluorescent emission at a specific frequency where there is no background from the excitation light, as relatively few cellular components are naturally fluorescent (called intrinsic or
autofluorescence Autofluorescence is the natural emission of light by biological structures such as mitochondria and lysosomes when they have absorbed light, and is used to distinguish the light originating from artificially added fluorescent markers (fluorophores) ...
). In fact, a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
or other component can be "labelled" with an extrinsic
fluorophore A fluorophore (or fluorochrome, similarly to a chromophore) is a fluorescent chemical compound that can re-emit light upon light excitation. Fluorophores typically contain several combined aromatic groups, or planar or cyclic molecules with se ...
, a fluorescent
dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...
that can be a small molecule, protein, or quantum dot, finding a large use in many biological applications. The quantification of a dye is done with a
spectrofluorometer A spectrofluorometer is an instrument which takes advantage of fluorescent properties of some compounds in order to provide information regarding their concentration and chemical environment in a sample. A certain excitation wavelength is selected, ...
and finds additional applications in:


Microscopy

* When scanning the fluorescence intensity across a plane one has
fluorescence microscopy A fluorescence microscope is an optical microscope that uses fluorescence instead of, or in addition to, scattering, reflection, and attenuation or absorption, to study the properties of organic or inorganic substances. "Fluorescence microscop ...
of tissues, cells, or subcellular structures, which is accomplished by labeling an antibody with a fluorophore and allowing the antibody to find its target antigen within the sample. Labelling multiple antibodies with different fluorophores allows visualization of multiple targets within a single image (multiple channels). DNA microarrays are a variant of this. * Immunology: An antibody is first prepared by having a fluorescent chemical group attached, and the sites (e.g., on a microscopic specimen) where the antibody has bound can be seen, and even quantified, by the fluorescence. * FLIM (
Fluorescence Lifetime Imaging Microscopy Fluorescence-lifetime imaging microscopy or FLIM is an imaging technique based on the differences in the exponential decay rate of the photon emission of a fluorophore from a sample. It can be used as an imaging technique in confocal microscopy, ...
) can be used to detect certain bio-molecular interactions that manifest themselves by influencing fluorescence lifetimes. * Cell and molecular biology: detection of
colocalization In fluorescence microscopy, colocalization refers to observation of the spatial overlap between two (or more) different fluorescent labels, each having a separate emission wavelength, to see if the different "targets" are located in the same area of ...
using fluorescence-labelled antibodies for selective detection of the antigens of interest using specialized software such as ImageJ.


Other techniques

* FRET (
Förster resonance energy transfer Förster resonance energy transfer (FRET), fluorescence resonance energy transfer, resonance energy transfer (RET) or electronic energy transfer (EET) is a mechanism describing energy transfer between two light-sensitive molecules ( chromophores). ...
, also known as
fluorescence resonance energy transfer Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, tha ...
) is used to study protein interactions, detect specific nucleic acid sequences and used as biosensors, while fluorescence lifetime (FLIM) can give an additional layer of information. * Biotechnology:
biosensors A biosensor is an analytical device, used for the detection of a chemical substance, that combines a biological component with a physicochemical detector. The ''sensitive biological element'', e.g. tissue, microorganisms, organelles, cell rece ...
using fluorescence are being studied as possible
Fluorescent glucose biosensors Fluorescent glucose biosensors are devices that measure the concentration of glucose in diabetic patients by means of sensitive protein that relays the concentration by means of fluorescence, an alternative to amperometric sension of glucose. Due ...
. * Automated sequencing of DNA by the
chain termination method Sanger sequencing is a method of DNA sequencing that involves electrophoresis and is based on the random incorporation of chain-terminating dideoxynucleotides by DNA polymerase during in vitro DNA replication. After first being developed by Fred ...
; each of four different chain terminating bases has its own specific fluorescent tag. As the labelled DNA molecules are separated, the fluorescent label is excited by a UV source, and the identity of the base terminating the molecule is identified by the wavelength of the emitted light. * FACS (
fluorescence-activated cell sorting Flow cytometry (FC) is a technique used to detect and measure physical and chemical characteristics of a population of cells or particles. In this process, a sample containing cells or particles is suspended in a fluid and injected into the flo ...
). One of several important cell sorting techniques used in the separation of different cell lines (especially those isolated from animal tissues). * DNA detection: the compound
ethidium bromide Ethidium bromide (or homidium bromide, chloride salt homidium chloride) is an intercalating agent commonly used as a fluorescent tag (nucleic acid stain) in molecular biology laboratories for techniques such as agarose gel electrophoresis. It i ...
, in aqueous solution, has very little fluorescence, as it is quenched by water. Ethidium bromide's fluorescence is greatly enhanced after it binds to DNA, so this compound is very useful in visualising the location of DNA fragments in
agarose gel electrophoresis Agarose gel electrophoresis is a method of gel electrophoresis used in biochemistry, molecular biology, genetics, and clinical chemistry to separate a mixed population of macromolecules such as DNA or proteins in a matrix of agarose, one of the ...
. Intercalated ethidium is in a hydrophobic environment when it is between the base pairs of the DNA, protected from quenching by water which is excluded from the local environment of the intercalated ethidium. Ethidium bromide may be carcinogenic – an arguably safer alternative is the dye
SYBR Green SYBR Green I (SG) is an asymmetrical cyanine dye used as a nucleic acid stain in molecular biology. The SYBR family of dyes is produced by Molecular Probes Inc., now owned by Thermo Fisher Scientific. SYBR Green I binds to DNA. The resulting DN ...
. * FIGS ( Fluorescence image-guided surgery) is a medical imaging technique that uses fluorescence to detect properly labeled structures during surgery. *
Intravascular fluorescence Intravascular fluorescence is a catheter-based molecular imaging technique that uses near-infrared fluorescence to detect artery wall autofluorescence (NIRAF) or fluorescence generated by molecular agents injected intravenously (NIRF) . No commerc ...
is a catheter-based medical imaging technique that uses fluorescence to detect high-risk features of atherosclerosis and unhealed vascular stent devices. Plaque autofluorescence has been used in a first-in-man study in coronary arteries in combination with
optical coherence tomography Optical coherence tomography (OCT) is an imaging technique that uses low-coherence light to capture micrometer-resolution, two- and three-dimensional images from within optical scattering media (e.g., biological tissue). It is used for medical ...
. Molecular agents has been also used to detect specific features, such as stent
fibrin Fibrin (also called Factor Ia) is a fibrous, non-globular protein involved in the clotting of blood. It is formed by the action of the protease thrombin on fibrinogen, which causes it to polymerize. The polymerized fibrin, together with platele ...
accumulation and enzymatic activity related to artery inflammation. * SAFI (species altered fluorescence imaging) an imaging technique in electrokinetics and
microfluidics Microfluidics refers to the behavior, precise control, and manipulation of fluids that are geometrically constrained to a small scale (typically sub-millimeter) at which surface forces dominate volumetric forces. It is a multidisciplinary field tha ...
. It uses non-electromigrating dyes whose fluorescence is easily quenched by migrating chemical species of interest. The dye(s) are usually seeded everywhere in the flow and differential quenching of their fluorescence by analytes is directly observed. * Fluorescence-based assays for screening
toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subst ...
chemicals. The optical assays consist of a mixture of environmental-sensitive fluorescent dyes and human skin cells that generate fluorescence spectra patterns. This approach can reduce the need for
laboratory animals Animals used by laboratories for testing purposes are largely supplied by dealers who specialize in selling them to universities, medical and veterinary schools, and companies that provide contract animal-testing services. It is comparatively rare ...
in biomedical research and pharmaceutical industry. *Bone-margin detection: Alizarin-stained specimens and certain fossils can be lit by fluorescent lights to view anatomical structures, including bone margins.


Forensics

Fingerprint A fingerprint is an impression left by the friction ridges of a human finger. The recovery of partial fingerprints from a crime scene is an important method of forensic science. Moisture and grease on a finger result in fingerprints on surfac ...
s can be visualized with fluorescent compounds such as
ninhydrin Ninhydrin (2,2-dihydroxyindane-1,3-dione) is an organic compound with the formula C6H4(CO)2C(OH)2. It is used to detect ammonia and amines. Upon reaction with these amines, ninhydrin gets converted into deep blue or purple derivatives, which are ...
or DFO (
1,8-Diazafluoren-9-one 1,8-Diazafluoren-9-one (DFO) is an aromatic ketone first synthesized in 1950. It is used to find fingerprints on porous surfaces. It makes fingerprints glow when they are lit by blue-green light. DFO reacts with amino acids present in the finger ...
). Blood and other substances are sometimes detected by fluorescent reagents, like
fluorescein Fluorescein is an organic compound and dye based on the xanthene tricyclic structural motif, formally belonging to triarylmethine dyes family. It is available as a dark orange/red powder slightly soluble in water and alcohol. It is widely used ...
.
Fiber Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporate ...
s, and other materials that may be encountered in
forensics Forensic science, also known as criminalistics, is the application of science to criminal and civil laws, mainly—on the criminal side—during criminal investigation, as governed by the legal standards of admissible evidence and crimina ...
or with a relationship to various
collectible A collectable (collectible or collector's item) is any object regarded as being of value or interest to a collector. Collectable items are not necessarily monetarily valuable or uncommon. There are numerous types of collectables and terms t ...
s, are sometimes fluorescent.


Non-destructive testing

Fluorescent penetrant inspection Fluorescent penetrant inspection (FPI) is a type of dye penetrant inspection in which a fluorescent dye is applied to the surface of a non-porous material in order to detect defects that may compromise the integrity or quality of the part in que ...
is used to find cracks and other defects on the surface of a part.
Dye tracing Dye tracing is a method of tracking and tracing various flows using dye as a flow tracer when added to a liquid. Dye tracing may be used to analyse the flow of the liquid or the transport of objects within the liquid. Dye tracking may be either qu ...
, using fluorescent dyes, is used to find leaks in liquid and gas plumbing systems.


Signage

Fluorescent colors are frequently used in
signage Signage is the design or use of signs and symbols to communicate a message. A signage also means signs ''collectively'' or being considered as a group. The term ''signage'' is documented to have been popularized in 1975 to 1980. Signs are any ...
, particularly road signs. Fluorescent colors are generally recognizable at longer ranges than their non-fluorescent counterparts, with fluorescent orange being particularly noticeable.Hawkins, H. Gene; Carlson, Paul John and Elmquist, Michael (2000
"Evaluation of fluorescent orange signs"
, Texas Transportation Institute Report 2962-S.
This property has led to its frequent use in safety signs and labels.


Optical brighteners

Fluorescent compounds are often used to enhance the appearance of fabric and paper, causing a "whitening" effect. A white surface treated with an optical brightener can emit more visible light than that which shines on it, making it appear brighter. The blue light emitted by the brightener compensates for the diminishing blue of the treated material and changes the hue away from yellow or brown and toward white. Optical brighteners are used in laundry detergents, high brightness paper, cosmetics,
high-visibility clothing High-visibility clothing, sometimes shortened to hi vis or hi viz, is any clothing worn that is highly luminescent in its natural matt property or a color that is easily discernible from any background. It is most commonly worn on the torso and ...
and more.


See also

* Absorption-re-emission
atomic line filter An atomic line filter (ALF) is a more effective optical band-pass filter used in the physical sciences for filtering electromagnetic radiation with precision, accuracy, and minimal signal strength loss. Atomic line filters work via the absorptio ...
s use the phenomenon of fluorescence to filter light extremely effectively. *
Black light A blacklight, also called a UV-A light, Wood's lamp, or ultraviolet light, is a lamp that emits long-wave (UV-A) ultraviolet light and very little visible light. One type of lamp has a violet filter material, either on the bulb or in a separat ...
*
Blacklight paint Black light paint or black light fluorescent paint is luminous paint that glows under a black light. It is based on pigments that respond to light in the ultraviolet segment of the electromagnetic spectrum. The paint may or may not be colorful ...
*
Fiber photometry Fiber photometry is a calcium imaging technique that captures 'bulk' or population-level calcium (Ca2+) activity from specific cell-types within a brain region or functional network in order to study neural circuits Population-level calcium activi ...
*
Fluorescence-activating and absorption-shifting tag FAST (''Fluorescence-Activating and absorption-Shifting Tag'') is a small, genetically-encoded, protein tag which allows for fluorescence reporting of proteins of interest. Unlike natural fluorescent proteins and derivates such as GFP or mCherry, ...
*
Fluorescence correlation spectroscopy Fluorescence correlation spectroscopy (FCS) is a statistical analysis, via time correlation, of stationary fluctuations of the fluorescence intensity. Its theoretical underpinning originated from L. Onsager's regression hypothesis. The analysis p ...
* Fluorescence image-guided surgery * Fluorescence in plants *
Fluorescence spectroscopy Fluorescence spectroscopy (also known as fluorimetry or spectrofluorometry) is a type of electromagnetic spectroscopy that analyzes fluorescence from a sample. It involves using a beam of light, usually ultraviolet light, that excites the electron ...
*
Fluorescent lamp A fluorescent lamp, or fluorescent tube, is a low-pressure mercury-vapor gas-discharge lamp that uses fluorescence to produce visible light. An electric current in the gas excites mercury vapor, which produces short-wave ultraviolet lig ...
*
Fluorescent Multilayer Disc 3D optical data storage is any form of optical data storage in which information can be recorded or read with three-dimensional resolution (as opposed to the two-dimensional resolution afforded, for example, by CD). This innovation has the pote ...
*
Fluorometer A fluorometer, fluorimeter or fluormeter is a device used to measure parameters of visible spectrum fluorescence: its intensity and wavelength distribution of emission spectrum after excitation by a certain spectrum of light. These parameters ar ...
*
High-visibility clothing High-visibility clothing, sometimes shortened to hi vis or hi viz, is any clothing worn that is highly luminescent in its natural matt property or a color that is easily discernible from any background. It is most commonly worn on the torso and ...
*
Integrated fluorometer Photosynthesis systems are electronic scientific instruments designed for non-destructive measurement of photosynthetic rates in the field. Photosynthesis systems are commonly used in agronomic and environmental research, as well as studies of t ...
*
Laser-induced fluorescence Laser-induced fluorescence (LIF) or laser-stimulated fluorescence (LSF) is a spectroscopic method in which an atom or molecule is excited to a higher energy level by the absorption of laser light followed by spontaneous emission of light. It was f ...
*
List of light sources This is a list of sources of light, the visible part of the electromagnetic spectrum. Light sources produce photons from another energy source, such as heat, chemical reactions, or conversion of mass or a different frequency of electromagnetic ener ...
*
Microbial art Microbial art, agar art, or germ art is artwork created by culturing microorganisms in certain patterns. The microbes used can be bacteria, yeast fungi, or less commonly, protists. The microbes can be chosen for their natural colours, or can be ...
, using fluorescent bacteria *
Mössbauer effect The Mössbauer effect, or recoilless nuclear resonance fluorescence, is a physical phenomenon discovered by Rudolf Mössbauer in 1958. It involves the resonant and recoil-free emission and absorption of gamma radiation by atomic nuclei bound in ...
, resonant fluorescence of gamma rays *
Organic light-emitting diode An organic light-emitting diode (OLED or organic LED), also known as organic electroluminescent (organic EL) diode, is a light-emitting diode (LED) in which the emissive electroluminescent layer is a film of organic compound that emits light i ...
s can be fluorescent *
Phosphorescence Phosphorescence is a type of photoluminescence related to fluorescence. When exposed to light (radiation) of a shorter wavelength, a phosphorescent substance will glow, absorbing the light and reemitting it at a longer wavelength. Unlike fluo ...
*
Phosphor thermometry Phosphor thermometry is an optical method for surface temperature measurement. The method exploits luminescence emitted by phosphor material. Phosphors are fine white or pastel-colored inorganic powders which may be stimulated by any of a variety of ...
, the use of phosphorescence to measure temperature. *
Spectroscopy Spectroscopy is the field of study that measures and interprets the electromagnetic spectra that result from the interaction between electromagnetic radiation and matter as a function of the wavelength or frequency of the radiation. Matter wa ...
*
Two-photon absorption Two-photon absorption (TPA or 2PA) or two-photon excitation or non-linear absorption is the simultaneous absorption of two photons of identical or different frequencies in order to excite a molecule from one state (usually the ground state) to a hi ...
*
Vibronic spectroscopy Vibronic spectroscopy is a branch of molecular spectroscopy concerned with vibronic transitions: the simultaneous changes in electronic and vibrational energy levels of a molecule due to the absorption or emission of a photon of the appropriate ...
*
X-ray fluorescence X-ray fluorescence (XRF) is the emission of characteristic "secondary" (or fluorescent) X-rays from a material that has been excited by being bombarded with high-energy X-rays or gamma rays. The phenomenon is widely used for elemental analysis ...


Footnotes


References


Further reading

*


External links


Fluorophores.org
the database of fluorescent dyes

Basic Concepts in Fluorescence
"A nano-history of fluorescence" lecture by David Jameson



Database of fluorescent minerals with pictures, activators and spectra (fluomin.org)

"Biofluorescent Night Dive – Dahab/Red Sea (Egypt), Masbat Bay/Mashraba, "Roman Rock""
YouTube. 9 October 2012. * Steffen O. Beyer
"FluoPedia.org: Publications"
fluopedia.org. * Steffen O. Beyer
"FluoMedia.org: Science"
fluomedia.org. {{Authority control Dyes Molecular biology Radiochemistry