Green fluorescent protein - Knowledge (XXG)

1228:; that is, the gene's regulatory sequence now controls the production of GFP, in addition to the tagged protein(s). In cells where the gene is expressed, and the tagged proteins are produced, GFP is produced at the same time. Thus, only those cells in which the tagged gene is expressed, or the target proteins are produced, will fluoresce when observed under fluorescence microscopy. Analysis of such time lapse movies has redefined the understanding of many biological processes including protein folding, protein transport, and RNA dynamics, which in the past had been studied using fixed (i.e., dead) material. Obtained data are also used to calibrate mathematical models of intracellular systems and to estimate rates of gene expression. Similarly, GFP can be used as an indicator of protein expression in heterologous systems. In this scenario, fusion proteins containing GFP are introduced indirectly, using RNA of the construct, or directly, with the tagged protein itself. This method is useful for studying structural and functional characteristics of the tagged protein on a macromolecular or single-molecule scale with fluorescence microscopy. 1159:-formaldehyde, phenol, triclosan, and paraben. GFP is great as a reporter protein because it has no effect on the host when introduced to the host's cellular environment. Due to this ability, no external visualization stain, ATP, or cofactors are needed. With regards to pollutant levels, the fluorescence was measured in order to gauge the effect that the pollutants have on the host cell. The cellular density of the host cell was also measured. Results from the study conducted by Song, Kim, & Seo (2016) showed that there was a decrease in both fluorescence and cellular density as pollutant levels increased. This was indicative of the fact that cellular activity had decreased. More research into this specific application in order to determine the mechanism by which GFP acts as a pollutant marker. Similar results have been observed in zebrafish because zebrafish that were injected with GFP were approximately twenty times more susceptible to recognize cellular stresses than zebrafish that were not injected with GFP. 1211:

method is to use a GFP that contains a mutation where the fluorescence will change from green to yellow over time, which is referred to as a fluorescent timer. With the fluorescent timer, researchers can study the state of protein production such as recently activated, continuously activated, or recently deactivated based on the color reported by the fluorescent protein. In yet another example, scientists have modified GFP to become active only after exposure to irradiation giving researchers a tool to selectively activate certain portions of a cell and observe where proteins tagged with the GFP move from the starting location. These are only two examples in a burgeoning field of fluorescent microcopy and a more complete review of biosensors utilizing GFP and other fluorescent proteins can be found here

822: 462: 837:, eqFP611, Dronpa, TagRFPs, KFP, EosFP/IrisFP, Dendra, and so on. Having been developed from proteins in different organisms, these proteins can sometimes display unanticipated approaches to chromophore formation. Some of these, such as KFP, are developed from naturally non- or weakly-fluorescent proteins to be greatly improved upon by mutagenesis. When GFP-like barrels of different spectra characteristics are used, the excitation spectra of one chromophore can be used to power another chromophore (FRET), allowing for conversion between wavelengths of light. 619: 1039:

residues of Gln94, Arg96, and His148 are able to stabilize by delocalizing the chromophore charge. Arg96 is the most important stabilizing residue due to the fact that it prompts the necessary structural realignments that are necessary from the HBI ring to occur. Any mutation to the Arg96 residue would result in a decrease in the development rate of the chromophore because proper electrostatic and steric interactions would be lost. Tyr66 is the recipient of hydrogen bonds and does not ionize in order to produce favorable electrostatics.

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binds Zn(II) and Cu(II) has been developed. BFPms1 have several important mutations including and the BFP chromophore (Y66H),Y145F for higher quantum yield, H148G for creating a hole into the beta-barrel and several other mutations that increase solubility. Zn(II) binding increases fluorescence intensity, while Cu(II) binding quenches fluorescence and shifts the absorbance maximum from 379 to 444 nm. Therefore, they can be used as Zn biosensor.

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of blue light. At a certain pulse threshold, the eGFP's optical output becomes brighter and completely uniform in color of pure green with a wavelength of 516 nm. Before being emitted as laser light, the light bounces back and forth within the resonator cavity and passes the cell numerous times. By studying the changes in optical activity, researchers may better understand cellular processes.

715: 452: 843:(FbFPs) were developed in 2007 and are a class of small (11–16 kDa), oxygen-independent fluorescent proteins that are derived from blue-light receptors. They are intended especially for the use under anaerobic or hypoxic conditions, since the formation and binding of the Flavin chromophore does not require molecular oxygen, as it is the case with the synthesis of the GFP chromophore. 1126: 1087: 574: 47: 1224:, where expression of GFP can be used as a marker for a particular characteristic. GFP can also be expressed in different structures enabling morphological distinction. In such cases, the gene for the production of GFP is incorporated into the genome of the organism in the region of the DNA that codes for the target proteins and that is controlled by the same 1331: 1380:, a German-born artist specializing in "protein sculptures," created sculptures based on the structure of GFP, including the 1.70 metres (5 feet 7 inches) tall "Green Fluorescent Protein" (2004) and the 1.40 metres (4 feet 7 inches) tall "Steel Jellyfish" (2006). The latter sculpture is located at the place of GFP's discovery by 2340: 1353:) that were initially developed to detect pollution in waterways. NeonPets, a US-based company has marketed green fluorescent mice to the pet industry as NeonMice. Green fluorescent pigs, known as Noels, were bred by a group of researchers led by Wu Shinn-Chih at the Department of Animal Science and Technology at 1210:

There are many techniques to utilize GFP in a live cell imaging experiment. The most direct way of utilizing GFP is to directly attach it to a protein of interest. For example, GFP can be included in a plasmid expressing other genes to indicate a successful transfection of a gene of interest. Another

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structure is a nearly perfect cylinder, 42Å long and 24Å in diameter (some studies have reported a diameter of 30Å), creating what is referred to as a "β-can" formation, which is unique to the GFP-like family. HBI, the spontaneously modified form of the tripeptide Ser65–Tyr66–Gly67, is nonfluorescent

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appears to evolutionarily prefer the less-efficient, dual-peaked excitation spectrum. Roger Tsien has speculated that varying hydrostatic pressure with depth may affect serine 65's ability to donate a hydrogen to the chromophore and shift the ratio of the two excitation peaks. Thus, the jellyfish may

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of GFP in jellyfish is unknown. GFP is co-expressed with aequorin in small granules around the rim of the jellyfish bell. The secondary excitation peak (480 nm) of GFP does absorb some of the blue emission of aequorin, giving the bioluminescence a more green hue. The serine 65 residue of the GFP

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derivatives (YFP, Citrine, Venus, YPet). BFP derivatives (except mKalama1) contain the Y66H substitution. They exhibit a broad absorption band in the ultraviolet centered close to 380 nanometers and an emission maximum at 448 nanometers. A green fluorescent protein mutant (BFPms1) that preferentially

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GFP can be used to analyse the colocalization of proteins. This is achieved by "splitting" the protein into two fragments which are able to self-assemble, and then fusing each of these to the two proteins of interest. Alone, these incomplete GFP fragments are unable to fluoresce. However, if the two

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The biggest advantage of GFP is that it can be heritable, depending on how it was introduced, allowing for continued study of cells and tissues it is expressed in. Visualizing GFP is noninvasive, requiring only illumination with blue light. GFP alone does not interfere with biological processes, but

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is responsible for the dual-peaked excitation spectra of wild-type GFP. It is conserved in all three GFP isoforms originally cloned by Prasher. Nearly all mutations of this residue consolidate the excitation spectra to a single peak at either 395 nm or 480 nm. The precise mechanism of this

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rather than phenol component. Several additional compensatory mutations in the surrounding barrel are required to restore brightness to this modified chromophore due to the increased bulk of the indole group. In ECFP and Cerulean, the N-terminal half of the seventh strand exhibits two conformations.

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filter sets, increasing the practicality of use by the general researcher. A 37 °C folding efficiency (F64L) point mutant to this scaffold, yielding enhanced GFP (EGFP), was discovered in 1995 by the laboratories of Thastrup and Falkow. EGFP allowed the practical use of GFPs in mammalian cells.

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in 1994. Frederick Tsuji's lab independently reported the expression of the recombinant protein one month later. Remarkably, the GFP molecule folded and was fluorescent at room temperature, without the need for exogenous cofactors specific to the jellyfish. Although this near-wtGFP was fluorescent,

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A novel possible use of GFP includes using it as a sensitive monitor of intracellular processes via an eGFP laser system made out of a human embryonic kidney cell line. The first engineered living laser is made by an eGFP expressing cell inside a reflective optical cavity and hitting it with pulses

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fluorescence from quenching by water. In addition to the auto-cyclization of the Ser65-Tyr66-Gly67, a 1,2-dehydrogenation reaction occurs at the Tyr66 residue. Besides the three residues that form the chromophore, residues such as Gln94, Arg96, His148, Thr203, and Glu222 all act as stabilizers. The

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Fluorescent proteins with other chromophores, such as UnaG with bilirubin, can display unique properties like red-shifted emission above 600 nm or photoconversion from a green-emitting state to a red-emitting state. They can have excitation and emission wavelengths far enough apart to achieve

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Additional site-directed random mutagenesis in combination with fluorescence lifetime based screening has further stabilized the seventh β-strand resulting in a bright variant, mTurquoise2, with a quantum yield (QY) of 0.93. The red-shifted wavelength of the YFP derivatives is accomplished by the

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when used in live cells, fluorescent proteins such as GFP are usually much less harmful when illuminated in living cells. This has triggered the development of highly automated live-cell fluorescence microscopy systems, which can be used to observe cells over time expressing one or more proteins

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microscope using the SPDM Phymod technology uses the so-called "reversible photobleaching" effect of fluorescent dyes like GFP and its derivatives to localize them as single molecules in an optical resolution of 10 nm. This can also be performed as a co-localization of two GFP derivatives

639:(FRET) experiments. Genetically encoded FRET reporters sensitive to cell signaling molecules, such as calcium or glutamate, protein phosphorylation state, protein complementation, receptor dimerization, and other processes provide highly specific optical readouts of cell activity in real time. 442:

Most commercially available genes for GFP and similar fluorescent proteins are around 730 base-pairs long. The natural protein has 238 amino acids. Its molecular mass is 27 kD. Therefore, fusing the GFP gene to the gene of a protein of interest can significantly increase the protein's size and

590:. This mutation dramatically improved the spectral characteristics of GFP, resulting in increased fluorescence, photostability, and a shift of the major excitation peak to 488 nm, with the peak emission kept at 509 nm. This matched the spectral characteristics of commonly available 1131:

Mechanistically, the process involves base-mediated cyclization followed by dehydration and oxidation. In the reaction of 7a to 8 involves the formation of an enamine from the imine, while in the reaction of 7b to 9 a proton is abstracted. The formed HBI fluorophore is highlighted in green.

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Macro-scale biological processes, such as the spread of virus infections, can be followed using GFP labeling. In the past, mutagenic ultra violet light (UV) has been used to illuminate living organisms (e.g., see) to detect and photograph the GFP expression. Recently, a technique using

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It has also been found that new lines of transgenic GFP rats can be relevant for gene therapy as well as regenerative medicine. By using "high-expresser" GFP, transgenic rats display high expression in most tissues, and many cells that have not been characterized or have been only poorly

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Maiti, Atanu; Buffalo, Cosmo Z.; Saurabh, Saumya; Montecinos-Franjola, Felipe; Hachey, Justin S.; Conlon, William J.; Tran, Geraldine N.; Hassan, Bakar; Walters, Kylie J.; Drobizhev, Mikhail; Moerner, W. E.; Ghosh, Partho; Matsuo, Hiroshi; Tsien, Roger Y.; Lin, John Y. (2023-07-12).

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sensitivity is complex, but, it seems, involves donation of a hydrogen from serine 65 to glutamate 222, which influences chromophore ionization. Since a single mutation can dramatically enhance the 480 nm excitation peak, making GFP a much more efficient partner of aequorin,

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These conformations both have a complex set of van der Waals interactions with the chromophore. The Y145A and H148D mutations in Cerulean stabilize these interactions and allow the chromophore to be more planar, better packed, and less prone to collisional quenching.

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formation and neighboring residue interactions. Researchers have modified these residues by directed and random mutagenesis to produce the wide variety of GFP derivatives in use today. Further research into GFP has shown that it is resistant to detergents, proteases,

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in the absence of the properly folded GFP scaffold and exists mainly in the un-ionized phenol form in wtGFP. Inward-facing sidechains of the barrel induce specific cyclization reactions in Ser65–Tyr66–Gly67 that induce ionization of HBI to the phenolate form and

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proteins of interest colocalize, then the two GFP fragments assemble together to form a GFP-like structure which is able to fluoresce. Therefore, by measuring the level of fluorescence it is possible to determine whether the two proteins of interest colocalize.

1034:. The hydrogen-bonding network and electron-stacking interactions with these sidechains influence the color, intensity and photostability of GFP and its numerous derivatives. The tightly packed nature of the barrel excludes solvent molecules, protecting the 3682:

Pan Y, Leifert A, Graf M, Schiefer F, Thoröe-Boveleth S, Broda J, Halloran MC, Hollert H, Laaf D, Simon U, Jahnen-Dechent W (March 2013). "High-sensitivity real-time analysis of nanoparticle toxicity in green fluorescent protein-expressing zebrafish".

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Remy S, Tesson L, Usal C, Menoret S, Bonnamain V, Nerriere-Daguin V, Rossignol J, Boyer C, Nguyen TH, Naveilhan P, Lescaudron L, Anegon I (Oct 2010). "New lines of GFP transgenic rats relevant for regenerative medicine and gene therapy".

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ions, inducing a blue glow. Some of this luminescent energy is transferred to the GFP, shifting the overall color towards green. However, its utility as a tool for molecular biologists did not begin to be realized until 1992 when

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Semirational mutagenesis of a number of residues led to pH-sensitive mutants known as pHluorins, and later super-ecliptic pHluorins. By exploiting the rapid change in pH upon synaptic vesicle fusion, pHluorins tagged to

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Lelimousin M, Noirclerc-Savoye M, Lazareno-Saez C, Paetzold B, Le Vot S, Chazal R, Macheboeuf P, Field MJ, Bourgeois D, Royant A (Oct 2009). "Intrinsic dynamics in ECFP and Cerulean control fluorescence quantum yield".

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Due to the potential for widespread usage and the evolving needs of researchers, many different mutants of GFP have been engineered. The first major improvement was a single point mutation (S65T) reported in 1995 in

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Rosenow MA, Huffman HA, Phail ME, Wachter RM (April 2004). "The crystal structure of the Y66L variant of green fluorescent protein supports a cyclization-oxidation-dehydration mechanism for chromophore maturation".

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when fused to proteins of interest, careful design of linkers is required to maintain the function of the protein of interest. Moreover, if used with a monomer it is able to diffuse readily throughout cells.

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3D reconstruction of confocal image of VEGF-overexpressing neural progenitors (red) and GFP-positive control neural progenitor cells (green) in the rat olfactory bulb. RECA-1-positive blood vessels - blue

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it had several drawbacks, including dual peaked excitation spectra, pH sensitivity, chloride sensitivity, poor fluorescence quantum yield, poor photostability and poor folding at 37 °C (99 °F).

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Many other mutations have been made, including color mutants; in particular, blue fluorescent protein (EBFP, EBFP2, Azurite, mKalama1), cyan fluorescent protein (ECFP, Cerulean, CyPet, mTurquoise2), and

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For example, GFP can be used as a reporter for environmental toxicity levels. This protein has been shown to be an effective way to measure the toxicity levels of various chemicals including ethanol,

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with fibroblast cells from sea anemones. The dogs give off a red fluorescent light, and they are meant to allow scientists to study the genes that cause human diseases like narcolepsy and blindness.

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GFP is used widely in cancer research to label and track cancer cells. GFP-labelled cancer cells have been used to model metastasis, the process by which cancer cells spread to distant organs.

423:, and maintained in their genome and that of their offspring. GFP has been expressed in many species, including bacteria, yeasts, fungi, fish and mammals, including in human cells. Scientists 4104:

Livet J, Weissman TA, Kang H, Draft RW, Lu J, Bennis RA, Sanes JR, Lichtman JW (Nov 2007). "Transgenic strategies for combinatorial expression of fluorescent proteins in the nervous system".

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The diversity of genetic mutations is illustrated by this San Diego beach scene drawn with living bacteria expressing 8 different colors of fluorescent proteins (derived from GFP and dsRed).

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Chudakov DM, Belousov VV, Zaraisky AG, Novoselov VV, Staroverov DB, Zorov DB, Lukyanov S, Lukyanov KA (February 2003). "Kindling fluorescent proteins for precise in vivo photolabeling".

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T203Y mutation and is due to π-electron stacking interactions between the substituted tyrosine residue and the chromophore. These two classes of spectral variants are often employed for

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non-mutagenic LED lights have been developed for macro-photography. The technique uses an epifluorescence camera attachment based on the same principle used in the construction of

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tendency at concentrations above 5 mg/mL. mGFP also stands for "modified GFP," which has been optimized through amino acid exchange for stable expression in plant cells.

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Wiens MD, Shen Y, Li X, Salem MA, Smisdom N, Zhang W, Brown A, Campbell RE (December 2016). "A Tandem Green-Red Heterodimeric Fluorescent Protein with High FRET Efficiency".

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agent in the larvae, preventing damage caused by high-intensity blue light by converting it into lower-intensity green light. However, these theories have not been tested.

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Ma Y, Yu JG, Sun Q, Li Z, Smith SC (2015-07-01). "The mechanism of dehydration in chromophore maturation of wild-type green fluorescent protein: A theoretical study".

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Shimomura O, Johnson FH, Saiga Y (Jun 1962). "Extraction, purification and properties of aequorin, a bioluminescent protein from the luminous hydromedusan, Aequorea".

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that a gene can be expressed throughout a given organism, in selected organs, or in cells of interest. GFP can be introduced into animals or other species through

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Bokman SH, Ward WW (1982). "Reversible denaturation of Aequorea green-fluorescent protein: physical separation and characterization of the renatured protein".

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Cabantous S, Terwilliger TC, Waldo GS (January 2005). "Protein tagging and detection with engineered self-assembling fragments of green fluorescent protein".

1190:(GFP-Snf2H and RFP-H2A), Co-localisation studies (2CLM) in the nucleus of a bone cancer cell. 120.000 localized molecules in a widefield area (470 μm). 4432:

Fakhrudin N, Ladurner A, Atanasov AG, Heiss EH, Baumgartner L, Markt P, Schuster D, Ellmerer EP, Wolber G, Rollinger JM, Stuppner H, Dirsch VM (Apr 2010).

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Terskikh A, Fradkov A, Ermakova G, Zaraisky A, Tan P, Kajava AV, et al. (November 2000). ""Fluorescent timer": protein that changes color with time".

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Another powerful use of GFP is to express the protein in small sets of specific cells. This allows researchers to optically detect specific types of cells

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Superfolder GFP (sfGFP), a series of mutations that allow GFP to rapidly fold and mature even when fused to poorly folding peptides, was reported in 2006.

5398: 5144: 4434:"Computer-aided discovery, validation, and mechanistic characterization of novel neolignan activators of peroxisome proliferator-activated receptor gamma" 3647:

Song YH, Kim CS, Seo JH (April 2016). "Noninvasive monitoring of environmental toxicity through green fluorescent protein expressing Escherichia coli".

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Ormö M, Cubitt AB, Kallio K, Gross LA, Tsien RY, Remington SJ (September 1996). "Crystal structure of the Aequorea victoria green fluorescent protein".

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Uckert W, Pedersen L, Günzburg W (2000). "Green fluorescent protein retroviral vector: generation of high-titer producer cells and virus supernatant".

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Prendergast FG, Mann KG (Aug 1978). "Chemical and physical properties of aequorin and the green fluorescent protein isolated from Aequorea forskålea".

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Drepper T, Eggert T, Circolone F, Heck A, Krauss U, Guterl JK, et al. (April 2007). "Reporter proteins for in vivo fluorescence without oxygen".

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Pédelacq JD, Cabantous S, Tran T, Terwilliger TC, Waldo GS (Jan 2006). "Engineering and characterization of a superfolder green fluorescent protein".

2203: 376:) has a single major excitation peak at 498 nm. GFP makes for an excellent tool in many forms of biology due to its ability to form an internal 5485: 2605:

Miesenböck G, De Angelis DA, Rothman JE (Jul 1998). "Visualizing secretion and synaptic transmission with pH-sensitive green fluorescent proteins".

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Zacharias DA, Violin JD, Newton AC, Tsien RY (May 2002). "Partitioning of lipid-modified monomeric GFPs into membrane microdomains of live cells".

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Ormö M, Cubitt AB, Kallio K, Gross LA, Tsien RY, Remington SJ (Sep 1996). "Crystal structure of the Aequorea victoria green fluorescent protein".

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Prasher DC, Eckenrode VK, Ward WW, Prendergast FG, Cormier MJ (Feb 1992). "Primary structure of the Aequorea victoria green-fluorescent protein".

4770:"An epifluorescent attachment improves whole-plant digital photography of Arabidopsis thaliana expressing red-shifted green fluorescent protein" 4676:

Zhu YJ, Agbayani R, Moore PH (Apr 2004). "Green fluorescent protein as a visual selection marker for papaya (Carica papaya L.) transformation".

1017:-hydroxybenzylidene)imidazolidin-5-one (HBI) running through the center. Five shorter alpha helices form caps on the ends of the structure. The 5480: 4641:

Rodman MK, Yadav NS, Artus NN (2002-09-01). "Progression of geminivirus-induced transgene silencing is associated with transgene methylation".

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Chudakov DM, Matz MV, Lukyanov S, Lukyanov KA (Jul 2010). "Fluorescent proteins and their applications in imaging living cells and tissues".

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assays were 0.97. Another application is the use of GFP co-transfection as internal control for transfection efficiency in mammalian cells.

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Elliott G, McGrath J, Crockett-Torabi E (Jun 2000). "Green fluorescent protein: A novel viability assay for cryobiological applications".

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Baker BJ, Mutoh H, Dimitrov D, Akemann W, Perron A, Iwamoto Y, Jin L, Cohen LB, Isacoff EY, Pieribone VA, Hughes T, Knöpfel T (Aug 2008).

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The nomenclature of modified GFPs is often confusing due to overlapping mapping of several GFP versions onto a single name. For example,

1251:(in the living organism). Genetically combining several spectral variants of GFP is a useful trick for the analysis of brain circuitry ( 1345:

using GFP for purposes of art and social commentary. The US company Yorktown Technologies markets to aquarium shops green fluorescent

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Chromophore binding. The critical mutation in cyan derivatives is the Y66W substitution, which causes the chromophore to form with an

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The reactions are catalyzed by residues Glu222 and Arg96. An analogous mechanism is also possible with threonine in place of Ser65.

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Morise H, Shimomura O, Johnson FH, Winant J (Jun 1974). "Intermolecular energy transfer in the bioluminescent system of Aequorea".

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Patterson GH, Lippincott-Schwartz J (September 2002). "A photoactivatable GFP for selective photolabeling of proteins and cells".

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molecular mass, and can impair the protein's natural function or change its location or trajectory of transport within the cell.

3563:"Construction and use of a Cupriavidus necator H16 soluble hydrogenase promoter (PSH) fusion to gfp (green fluorescent protein)" 1009:

structure consisting of eleven β-strands with a pleated sheet arrangement, with an alpha helix containing the covalently bonded

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Baker SS, Vidican CB, Cameron DS, Greib HG, Jarocki CC, Setaputri AW, Spicuzza CH, Burr AA, Waqas MA, Tolbert DA (2012-01-01).

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Montecinos-Franjola F, Lin JY, Rodriguez EA (2020-11-16). "Fluorescent proteins for in vivo imaging, where's the biliverdin?".

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of 395 nm and a minor one at 475 nm. Its emission peak is at 509 nm, which is in the lower green portion of the

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Arun KH, Kaul CL, Ramarao P (2005). "Green fluorescent proteins in receptor research: an emerging tool for drug discovery".

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Chalfie M, Tu Y, Euskirchen G, Ward WW, Prasher DC (Feb 1994). "Green fluorescent protein as a marker for gene expression".

1552: 195: 4818:"PlantEdDL - Using SRL digital cameras in quantitative investigations of plants expressing green fluorescent protein (GFP)" 2548:

Goedhart J, von Stetten D, Noirclerc-Savoye M, Lelimousin M, Joosen L, Hink MA, van Weeren L, Gadella TW, Royant A (2012).

224: 5526: 5511: 5325: 5230: 959: 596: 4721:"Non-invasive quantitative detection and applications of non-toxic, S65T-type green fluorescent protein in living plants" 1687:

Phillips GJ (Oct 2001). "Green fluorescent protein—a bright idea for the study of bacterial protein localization".

5373: 3967:"Using a single fluorescent reporter gene to infer half-life of extrinsic noise and other parameters of gene expression" 1970:"Aequorea green fluorescent protein. Expression of the gene and fluorescence characteristics of the recombinant protein" 1198:

and the way it is used in cell biology and other biological disciplines. While most small fluorescent molecules such as

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Mattson S, Tran GN, Rodriguez EA (2023). "Directed Evolution of Fluorescent Proteins in Bacteria". In Sharma M (ed.).

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There are many GFP-like proteins that, despite being in the same protein family as GFP, are not directly derived from

3344:"Kinetic isotope effect studies on the de novo rate of chromophore formation in fast- and slow-maturing GFP variants" 2395:

McRae SR, Brown CL, Bushell GR (May 2005). "Rapid purification of EGFP, EYFP, and ECFP with high yield and purity".

5123: 1434: 1407: 1389: 1354: 611: 5330: 5320: 5310: 754: 351: 846: 5531: 5521: 5506: 5350: 5223: 1385: 868: 461: 436: 1640:"Fluorescent proteins as biomarkers and biosensors: throwing color lights on molecular and cellular processes" 989: 183: 2338:, Thastrup O, Tullin S, Kongsbak Poulsen L, Bjørn S, "Fluorescent Proteins", published 2001-01-09 4549:

Kouros-Mehr H, Bechis SK, Slorach EM, Littlepage LE, Egeblad M, Ewald AJ, Pai SY, Ho IC, Werb Z (Feb 2008).

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Rodriguez EA, Campbell RE, Lin JY, Lin MZ, Miyawaki A, Palmer AE, Shu X, Zhang J, Tsien RY (February 2017).

2750:"The evolution of genes encoding for green fluorescent proteins: insights from cephalochordates (amphioxus)" 1319: 1220: 1195: 1177: 881: 821: 381: 618: 5421: 5383: 4069:

Chudakov DM, Lukyanov S, Lukyanov KA (Dec 2005). "Fluorescent proteins as a toolkit for in vivo imaging".

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change the color of its bioluminescence with depth. However, a collapse in the population of jellyfish in

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expressed the coding sequence of wtGFP, with the first few amino acids deleted, in heterologous cells of

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GFP, which is often achieved by the dimer interface breaking A206K mutation. Wild-type GFP has a weak

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Stepanenko OV, Verkhusha VV, Shavlovsky MM, Kuznetsova IM, Uversky VN, Turoverov KK (November 2008).

3163: 2820: 2761: 2706: 2614: 2561: 2474: 2293: 2156: 2025: 1926: 1467: 1067: 562: 136: 1255:). Other interesting uses of fluorescent proteins in the literature include using FPs as sensors of 552:

in 1996. One month later, the Phillips group independently reported the wild-type GFP structure in

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The first reported crystal structure of a GFP was that of the S65T mutant by the Remington group in

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Reviews on new classes of fluorescent proteins and applications can be found in the cited reviews.

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towards the mouth of the lancelet, serving as a passive hunting mechanism. It may also serve as a

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Gunkel M, Erdel F, Rippe K, Lemmer P, Kaufmann R, Hörmann C, Amberger R, Cremer C (Jun 2009).

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Goodman SR, ed. (2008). "Chapter 1 - Tools of the Cell Biologist: Green Fluorescent Protein".

1745: 1735: 1704: 1669: 1610: 1594: 1575: 1526: 1483: 1096: 1072: 943: 927: 766: 548: 538: 400: 315: 170: 5457: 4969: 4918: 4910: 4881: 4789: 4781: 4732: 4685: 4650: 4607: 4570: 4562: 4498: 4453: 4445: 4406: 4363: 4325: 4317: 4276: 4266: 4217: 4176: 4168: 4121: 4078: 4035: 3994: 3986: 3937: 3927: 3880: 3837: 3786: 3749: 3739: 3692: 3656: 3621: 3584: 3574: 3535: 3496: 3455: 3447: 3400: 3363: 3355: 3316: 3273: 3236: 3228: 3187: 3171: 3115: 3070: 3037: 3029: 2979: 2936: 2893: 2838: 2828: 2779: 2769: 2714: 2669: 2622: 2577: 2569: 2522: 2486: 2439: 2404: 2367: 2301: 2218: 2174: 2164: 2096: 2088: 2033: 1983: 1934: 1889: 1852: 1815: 1768: 1727: 1696: 1659: 1651: 1602: 1567: 1518: 1475: 873: 526: 416: 359: 5133: 3069:. Methods in Molecular Biology. Vol. 2564. New York, NY: Springer US. pp. 75–97. 2656:

Hanson GT, Aggeler R, Oglesbee D, Cannon M, Capaldi RA, Tsien RY, Remington SJ (Mar 2004).

162: 5426: 5250: 5163: 3152:"Structural and photophysical characterization of the small ultra-red fluorescent protein" 1411: 1381: 1361:

as proof of concept to use them potentially as model organisms for diseases, particularly

1286: 877: 825:

Different proteins produce different fluorescent colors when exposed to ultraviolet light.

779: 731: 509: 492: 428: 264: 31: 4023: 3436:"Understanding the role of Arg96 in structure and stability of green fluorescent protein" 439:

on 10 October 2008 for their discovery and development of the green fluorescent protein.

17: 4494: 4262: 4117: 3982: 3876: 3833: 3735: 3531: 3192: 3167: 3151: 2824: 2765: 2710: 2618: 2565: 2550:"Structure-guided evolution of cyan fluorescent proteins towards a quantum yield of 93%" 2297: 2160: 2029: 1930: 1471: 1334:

Mice expressing GFP under UV light (left & right), compared to normal mouse (center)

810:

of 0.92, making them nearly two-fold brighter than the commonly used EGFP isolated from

5365: 5282: 5171: 5167: 4923: 4898: 4794: 4769: 4575: 4551:"GATA-3 links tumor differentiation and dissemination in a luminal breast cancer model" 4550: 4458: 4433: 4330: 4305: 4181: 4156: 3999: 3966: 3942: 3915: 3754: 3719: 3589: 3562: 3460: 3435: 3368: 3343: 3241: 3216: 3042: 3017: 2843: 2809:"Very bright green fluorescent proteins from the Pontellid copepod Pontella mimocerami" 2808: 2784: 2749: 2582: 2549: 1880: 1772: 1664: 1639: 1365:. In 2009 a South Korean team from Seoul National University bred the first transgenic 1187: 1091: 679: 587: 521: 432: 424: 5149: 4281: 4246: 1700: 1395: 5500: 5462: 5452: 5393: 5204: 5040: 4894: 4737: 4720: 4654: 3806: 3293: 3119: 3016:

Rodriguez EA, Tran GN, Gross LA, Crisp JL, Shu X, Lin JY, Tsien RY (September 2016).

2372: 2355: 2179: 2138: 1988: 1969: 1893: 1624: 1571: 1338: 1149: 963: 923: 863: 807: 691: 683: 644: 600: 408: 363: 128: 111: 4981: 4705: 4627: 4510: 4383: 4055: 3900: 3668: 3420: 2913: 2734: 2321: 2238: 2118: 2053: 1790: 1495: 850:

White light image, or image seen by the eye, of fluorescent proteins in image above.

79: 5447: 5416: 5277: 4842: 4141: 3135: 2999: 2956: 2642: 2459: 2137:

Brejc K, Sixma TK, Kitts PA, Kain SR, Tsien RY, Ormö M, Remington SJ (March 1997).

1974: 1954: 1215: 1054: 739: 396: 299: 158: 4754: 4082: 3916:"Genetically encoded fluorescent biosensors illuminate kinase signaling in cancer" 3841: 2037: 1479: 654:) was engineered by introduction of cysteines into the beta barrel structure. The 92: 5215: 4960:

Voss-Andreae J (2005). "Protein Sculptures: Life's Building Blocks Inspire Art".

4222: 4205: 3539: 2833: 1638:

Stepanenko OV, Verkhusha VV, Kuznetsova IM, Uversky VN, Turoverov KK (Aug 2008).

1341:, a green-fluorescent rabbit, was created by a French laboratory commissioned by 603:(QY) of EGFP is 0.60. The relative brightness, expressed as ε•QY, is 33,000 Mcm. 104: 27:

Protein that exhibits bright green fluorescence when exposed to ultraviolet light

4206:"Photoinactivation of native AMPA receptors reveals their real-time trafficking" 3625: 3074: 2477:(Apr 2002). "Structural chemistry of a green fluorescent protein Zn biosensor". 1731: 1424: 1342: 1290: 1282: 1267: 1232: 1199: 1063: 1059: 1035: 1023: 1018: 1010: 1006: 947: 931: 896: 793: 786: 744: 714: 659: 591: 557: 451: 377: 328: 303: 274: 4251:

Proceedings of the National Academy of Sciences of the United States of America

3724:

Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

1655: 702: 5259: 4973: 4689: 4566: 4367: 4172: 3990: 3932: 3660: 3018:"A far-red fluorescent protein evolved from a cyanobacterial phycobiliprotein" 2408: 1203: 951: 899: 797: 355: 332: 240: 3547: 3183: 1058:

GFP molecules drawn in cartoon style, one fully and one with the side of the

5089: 5061: 4785: 4502: 4271: 3884: 3744: 2718: 2254:

Fluorescent Analogs of Biomolecular Building Blocks: Design and Applications

1938: 1346: 1271: 911: 892: 484: 415:, and many animals have been created that express GFP, which demonstrates a 412: 367: 311: 5182: 4932: 4803: 4746: 4697: 4662: 4619: 4584: 4467: 4449: 4418: 4410: 4375: 4339: 4290: 4231: 4190: 4133: 4090: 4047: 4039: 4008: 3951: 3892: 3849: 3798: 3763: 3704: 3696: 3633: 3598: 3508: 3469: 3412: 3377: 3285: 3250: 3201: 3092: 3051: 2991: 2948: 2940: 2905: 2852: 2793: 2726: 2683: 2674: 2657: 2591: 2534: 2498: 2451: 2416: 2230: 2169: 1827: 1819: 1749: 1708: 1673: 1125: 1086: 5176: 3790: 3328: 3127: 2634: 2381: 2313: 2188: 2110: 2045: 1997: 1946: 1901: 1864: 1606: 1579: 1487: 1046:

GFP Movie showing entire structure and zoom in to fluorescent chromophore.

573: 132: 46: 4306:"Visualization of targeted transduction by engineered lentiviral vectors" 3217:"The Growing and Glowing Toolbox of Fluorescent and Photoactive Proteins" 2092: 1252: 1241: 775: 770:, lancelets do not produce their own blue light, and the origin of their 761: 735: 720: 500: 476: 475:

In the 1960s and 1970s, GFP, along with the separate luminescent protein

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Video of 2008 Nobel Prize lecture of Roger Tsien on fluorescent proteins

4125: 3320: 3277: 1856: 1530: 1522: 1330: 30:"EGFP" redirects here. For the airport with that ICAO airport code, see 5246: 5209: 5199: 4914: 4321: 3579: 3451: 3033: 2866: 2573: 1350: 1247: 903: 789: 687: 504: 385: 336: 295: 3500: 3359: 2774: 2526: 2490: 2101: 1100: 1076: 2305: 2222: 1256: 1194:

The availability of GFP and its derivatives has thoroughly redefined

979: 955: 939: 935: 919: 889: 885: 627: 480: 190: 5014: 4611: 3342:

Pouwels LJ, Zhang L, Chan NH, Dorrestein PC, Wachter RM (Sep 2008).

2983: 2443: 2139:"Structural basis for dual excitation and photoisomerization of the 1767:(3rd ed.). Amsterdam: Elsevier/Academic Press. pp. 14–25. 5162:

dedicated to the 2008 Nobel Prize winners in Chemistry, Professors

2897: 5408: 2626: 1394: 1366: 1329: 975: 915: 907: 845: 834: 820: 764:

are known to produce GFP in various regions of their body. Unlike

713: 701: 663: 655: 651: 617: 324: 116: 5185:: an illustrated overview of GFP-like variants by David Goodsell. 5155:

Excitation and emission spectra for various fluorescent proteins

1726:. Methods in Molecular Medicine. Vol. 35. pp. 275–85. 1429: 1263: 967: 517: 404: 152: 86: 74: 5219: 5143:

Interactive Java applet demonstrating the chemistry behind the

5134:

A comprehensive article on fluorescent proteins at Scholarpedia

5042:

Aglow in the Dark: The Revolutionary Science of Biofluorescence

4899:"Antiviral restriction factor transgenesis in the domestic cat" 4157:"Genetically encoded fluorescent sensors of membrane potential" 4024:"Dual color localization microscopy of cellular nanostructures" 2356:"FACS-optimized mutants of the green fluorescent protein (GFP)" 4719:

Niwa Y, Hirano T, Yoshimoto K, Shimizu M, Kobayashi H (1999).

1362: 1358: 983: 516:. The funding for this project had run out, so Prasher sent 323:. However, GFPs have been found in other organisms including 5194:

Overview of all the structural information available in the

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Popular science book describing history and discovery of GFP

966:(0.20), which makes it comparable biophysical brightness to 310:

traditionally refers to the protein first isolated from the

4481:

Gather MC, Yun SH (2011). "Single-cell biological lasers".

647:

have been used to visualize synaptic activity in neurons.

487:, releasing light), was first purified from the jellyfish 4245:

Lakadamyali M, Rust MJ, Babcock HP, Zhuang X (Aug 2003).

1406:(2006). The image shows the stainless-steel sculpture at 785:

GFP-like proteins have been found in multiple species of

512:

reported the cloning and nucleotide sequence of wtGFP in

3106:

Tsien RY (1998-01-01). "The green fluorescent protein".

556:. These crystal structures provided vital background on 4860: 4247:"Visualizing infection of individual influenza viruses" 3561:

Jugder BE, Welch J, Braidy N, Marquis CP (2016-07-26).

2807:

Hunt ME, Scherrer MP, Ferrari FD, Matz MV (July 2010).

2748:

Yue JX, Holland ND, Holland LZ, Deheyn DD (June 2016).

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of various organisms for identification purposes as in

1214:

For example, GFP had been widely used in labelling the

2074:"The molecular structure of green fluorescent protein" 1601:. Boca Raton: Jenny Stanford Publishing. p. 122. 774:

GFP is still unknown. Some speculate that it attracts

3777:

Yuste R (Dec 2005). "Fluorescence microscopy today".

1357:. A Japanese-American Team created green-fluorescent 565:(GdmCl) treatments, and drastic temperature changes. 3614:

Journal of Pharmacological and Toxicological Methods

5471: 5435: 5407: 5364: 5258: 5179:: an illustrated overview of GFP by David Goodsell. 5160:

Green Fluorescent Protein Chem Soc Rev themed issue

4947:"Fluorescent puppy is world's first transgenic dog" 4526:"Green Fluorescent Protein Makes for Living Lasers" 1113:

Autocatalytic formation of the chromophore in wtGFP

982:spectral properties are similar to the organic dye 270: 260: 255: 239: 231: 219: 214: 209: 189: 169: 151: 146: 122: 110: 98: 85: 73: 65: 60: 39: 5039: 3691:(6). Weinheim an Der Bergstrasse, Germany: 863–9. 686:. However, the same term is also used to refer to 4882:Scientists in Taiwan breed fluorescent green pigs 993:E. coli colonies expressing fluorescent proteins. 3965:Komorowski M, Finkenstädt B, Rand D (Jun 2010). 1278:characterized in previous GFP-transgenic rats. 1119:3,5-Difluoro-4-hydroxybenzylidene imidazolinone 2202:Shaner NC, Steinbach PA, Tsien RY (Dec 2005). 1807:Journal of Cellular and Comparative Physiology 970:and ~2-fold brighter than most red or far-red 5231: 4204:Adesnik H, Nicoll RA, England PM (Dec 2005). 2132: 2130: 2128: 411:. It has been used in modified forms to make 8: 5072:Glowing Genes: A Revolution In Biotechnology 4893:Wongsrikeao P, Saenz D, Rinkoski T, Otoi T, 2067: 2065: 2063: 2011: 2009: 2007: 1546: 1544: 1542: 1540: 738:'s action on luciferin) and the (secondary) 599:(denoted ε) of 55,000 Mcm. The fluorescence 4995:Pawlak A (2005). "Inspirierende Proteine". 1414:(Wash., USA), the place of GFP's discovery. 1148:Green fluorescent protein may be used as a 806:has shown high levels of brightness with a 5238: 5224: 5216: 2354:Cormack BP, Valdivia RH, Falkow S (1996). 2204:"A guide to choosing fluorescent proteins" 1289:. Correlation of viability as measured by 1202:(fluorescein isothiocyanate) are strongly 252: 143: 45: 5191:on FPbase, a fluorescent protein database 4922: 4793: 4736: 4574: 4457: 4329: 4280: 4270: 4221: 4180: 3998: 3941: 3931: 3753: 3743: 3588: 3578: 3459: 3367: 3240: 3191: 3041: 2842: 2832: 2783: 2773: 2673: 2581: 2371: 2178: 2168: 2100: 2072:Yang F, Moss LG, Phillips GN (Oct 1996). 1987: 1663: 678:often refers to a GFP with an N-terminal 3914:Lin W, Mehta S, Zhang J (October 2019). 2479:Journal of the American Chemical Society 2277:Heim R, Cubitt AB, Tsien RY (Feb 1995). 1181: 1085: 1053: 1041: 988: 855:conversion between red and green light. 572: 460: 450: 1450: 622:A palette of variants of GFP and DsRed. 5481:Photoactivated localization microscopy 5399:Protein–protein interaction prediction 3649:Korean Journal of Chemical Engineering 2473:Barondeau DP, Kassmann CJ, Tainer JA, 658:state of the cysteines determines the 366:(QY) of GFP is 0.79. The GFP from the 206: 36: 3481: 3479: 3011: 3009: 1644:Current Protein & Peptide Science 1440:Genetically encoded voltage indicator 1274:viruses and lentiviral viruses, etc. 302:when exposed to light in the blue to 7: 5139:Brief summary of landmark GFP papers 1599:Fundamentals of Fluorescence Imaging 520:samples to several labs. The lab of 5356:Freeze-fracture electron microscopy 5208:(Green fluorescent protein) at the 3920:The Journal of Biological Chemistry 2662:The Journal of Biological Chemistry 2397:Protein Expression and Purification 1281:GFP has been shown to be useful in 918:-derived GFP-like proteins require 1773:10.1016/B978-0-12-370458-0.50006-2 1207:tagged with fluorescent proteins. 1117:For a synthetic analogue see also 730:The purpose of both the (primary) 25: 5074:. Buffalo, NY: Prometheus Books. 710:) under a fluorescent microscope. 637:Förster resonance energy transfer 5336:Isothermal titration calorimetry 5316:Dual-polarization interferometry 5183:Molecule of the Month, June 2014 5177:Molecule of the Month, June 2003 5145:formation of the GFP chromophore 4738:10.1046/j.1365-313X.1999.00464.x 4655:10.1046/j.1469-8137.2002.00467.x 3266:Biochemical Society Transactions 3120:10.1146/annurev.biochem.67.1.509 1572:10.1146/annurev.biochem.67.1.509 1270:and the infection of individual 1124: 962:(180,000 M cm) and has a modest 902:without the need of an external 841:FMN-binding fluorescent proteins 483:that catalyzes the breakdown of 380:without requiring any accessory 5015:"Julian Voss-Andreae Sculpture" 1968:Inouye S, Tsuji FI (Mar 1994). 1553:"The green fluorescent protein" 1028:post-translational modification 682:that causes the GFP to bind to 499:, GFP fluorescence occurs when 5038:Pieribone V, Gruber D (2006). 3221:Trends in Biochemical Sciences 491:and its properties studied by 1: 5326:Chromatin immunoprecipitation 4083:10.1016/j.tibtech.2005.10.005 3842:10.1126/science.290.5496.1585 3108:Annual Review of Biochemistry 2279:"Improved green fluorescence" 2252:Wilhelmsson M, Tor Y (2016). 2038:10.1126/science.273.5280.1392 1701:10.1016/S0378-1097(01)00358-5 1560:Annual Review of Biochemistry 1480:10.1126/science.273.5280.1392 1266:receptors on cell membranes, 392:other than molecular oxygen. 147:Available protein structures: 5389:Protein structural alignment 5374:Protein structure prediction 5124:Resources in other libraries 5046:. Cambridge: Belknap Press. 4223:10.1016/j.neuron.2005.11.030 3540:10.1016/j.cplett.2015.04.061 2834:10.1371/journal.pone.0011517 2373:10.1016/0378-1119(95)00685-0 1989:10.1016/0014-5793(94)80472-9 1894:10.1016/0378-1119(92)90691-H 800:families. GFP isolated from 536:, publishing the results in 5473:Super-resolution microscopy 5379:Protein function prediction 5307:Peptide mass fingerprinting 5302:Protein immunoprecipitation 4857:"Glow-In-The Dark NeonMice" 4304:Joo KI, Wang P (Oct 2008). 3626:10.1016/j.vascn.2004.07.006 3075:10.1007/978-1-0716-2667-2_4 1320:epifluorescence microscopes 1026:formation. This process of 5553: 3405:10.1152/physrev.00038.2009 3233:10.1016/j.tibs.2016.09.010 3176:10.1038/s41467-023-39776-9 2143:green fluorescent protein" 1656:10.2174/138920308785132668 1435:Yellow fluorescent protein 1408:Friday Harbor Laboratories 1390:Friday Harbor Laboratories 1355:National Taiwan University 1175: 1116: 1051: 817:Other fluorescent proteins 612:yellow fluorescent protein 55:green fluorescent protein. 29: 5331:Surface plasmon resonance 5321:Microscale thermophoresis 5311:Protein mass spectrometry 5273:Green fluorescent protein 5189:Green Fluorescent Protein 5119:Resources in your library 5110:Green fluorescent protein 4974:10.1162/leon.2005.38.1.41 4690:10.1007/s00299-004-0755-5 4567:10.1016/j.ccr.2008.01.011 4368:10.1007/s11248-009-9352-2 4173:10.1007/s11068-008-9026-7 3991:10.1016/j.bpj.2010.03.032 3933:10.1074/jbc.REV119.006177 3661:10.1007/s11814-015-0253-1 2409:10.1016/j.pep.2004.12.030 1732:10.1385/1-59259-086-1:275 1689:FEMS Microbiology Letters 1186:Superresolution with two 288:green fluorescent protein 251: 210:Green fluorescent protein 142: 44: 40:Green fluorescent protein 18:Green Fluorescent Protein 5351:Cryo-electron microscopy 3524:Chemical Physics Letters 1386:University of Washington 880:, and named small ultra 869:Trichodesmium erythraeum 792:, particularly from the 437:Nobel Prize in Chemistry 407:is frequently used as a 319:and is sometimes called 5384:Protein–protein docking 5297:Protein electrophoresis 4503:10.1038/nphoton.2011.99 4272:10.1073/pnas.0832269100 4071:Trends in Biotechnology 3885:10.1126/science.1074952 3745:10.1073/pnas.0904061106 3720:"GFP: Lighting up life" 2719:10.1126/science.1068539 1939:10.1126/science.8303295 1402:'s GFP-based sculpture 1221:Drosophila melanogaster 1196:fluorescence microscopy 1178:Fluorescence microscope 1172:Fluorescence microscopy 1062:cut away to reveal the 882:red fluorescent protein 5283:Protein immunostaining 4450:10.1124/mol.109.062141 4438:Molecular Pharmacology 4411:10.1006/cryo.2000.2258 4040:10.1002/biot.200900005 3718:Chalfie M (Jun 2009). 3697:10.1002/smll.201201173 2941:10.1002/cbic.201600492 2675:10.1074/jbc.M312846200 2170:10.1073/pnas.94.6.2306 1820:10.1002/jcp.1030590302 1724:Gene Therapy of Cancer 1595:"Fluorescent Proteins" 1415: 1335: 1191: 1105: 1081: 1047: 994: 960:extinction coefficient 895:self-incorporates the 851: 826: 727: 718:In the marine copepod 711: 623: 597:extinction coefficient 578: 467: 458: 435:were awarded the 2008 409:reporter of expression 5341:X-ray crystallography 4786:10.1093/aobpla/pls003 4028:Biotechnology Journal 3791:10.1038/nmeth1205-902 3393:Physiological Reviews 3156:Nature Communications 2554:Nature Communications 2336:US patent 6172188 2256:. New Jersey: Wiley. 1607:10.1201/9781351129404 1398: 1333: 1245:(in a dish), or even 1185: 1089: 1057: 1045: 992: 849: 824: 717: 705: 650:Redox sensitive GFP ( 621: 576: 471:Wild-type GFP (wtGFP) 464: 454: 421:transgenic techniques 5527:Fluorescent proteins 5512:Recombinant proteins 5268:Protein purification 4863:on February 14, 2009 4600:Nature Biotechnology 3067:Fluorescent Proteins 2972:Nature Biotechnology 2886:Nature Biotechnology 2432:Nature Biotechnology 2366:(1 Spec No): 33–38. 2093:10.1038/nbt1096-1246 2081:Nature Biotechnology 1765:Medical Cell Biology 972:fluorescent proteins 563:guanidinium chloride 554:Nature Biotechnology 384:, gene products, or 298:that exhibits green 5293:Gel electrophoresis 4530:Scientific American 4495:2011NaPho...5..406G 4356:Transgenic Research 4263:2003PNAS..100.9280L 4126:10.1038/nature06293 4118:2007Natur.450...56L 3983:2010BpJ....98.2759K 3971:Biophysical Journal 3926:(40): 14814–14822. 3877:2002Sci...297.1873P 3834:2000Sci...290.1585T 3736:2009PNAS..10610073C 3730:(25): 10073–10080. 3532:2015CPL...631...42M 3321:10.1021/bi00262a003 3278:10.1042/BST20200444 3168:2023NatCo..14.4155M 2825:2010PLoSO...511517H 2766:2016NatSR...628350Y 2711:2002Sci...296..913Z 2619:1998Natur.394..192M 2566:2012NatCo...3..751G 2521:(42): 10038–10046. 2298:1995Natur.373..663H 2161:1997PNAS...94.2306B 2030:1996Sci...273.1392O 1931:1994Sci...263..802C 1857:10.1021/bi00709a028 1523:10.1021/bi00610a004 1472:1996Sci...273.1392O 1400:Julian Voss-Andreae 1378:Julian Voss-Andreae 1226:regulatory sequence 862:was evolved from a 860:fluorescent protein 803:Pontella mimocerami 362:. The fluorescence 5537:Cnidarian proteins 5436:Display techniques 5288:Protein sequencing 4915:10.1038/nmeth.1703 4822:planted.botany.org 4678:Plant Cell Reports 4322:10.1038/gt.2008.87 4161:Brain Cell Biology 3580:10.7717/peerj.2269 3526:. 631–632: 42–46. 3452:10.1002/prot.22089 3034:10.1038/nmeth.3935 2754:Scientific Reports 2574:10.1038/ncomms1738 1597:. In Cox G (ed.). 1416: 1336: 1260:membrane potential 1192: 1106: 1082: 1048: 1030:is referred to as 995: 852: 827: 728: 712: 624: 579: 468: 459: 373:Renilla reniformis 306:range. The label 5494: 5493: 5443:Bacterial display 5105:Library resources 5081:978-1-59102-253-4 5070:Zimmer M (2005). 5053:978-0-674-01921-8 4725:The Plant Journal 4524:Matson J (2011). 3977:(12): 2759–2769. 3501:10.1021/bi0361315 3495:(15): 4464–4472. 3360:10.1021/bi8007164 3315:(19): 4535–4540. 3084:978-1-0716-2666-5 2935:(24): 2361–2367. 2775:10.1038/srep28350 2527:10.1021/bi901093w 2491:10.1021/ja0176954 2485:(14): 3522–3524. 2263:978-1-118-17586-6 2141:Aequorea victoria 1782:978-0-12-370458-0 1551:Tsien RY (1998). 1313:Macro-photography 1110: 1109: 944:hydrogen peroxide 938:does not require 928:hydrogen peroxide 893:autocatalytically 831:Aequorea victoria 489:Aequorea victoria 456:Aequorea victoria 401:molecular biology 316:Aequorea victoria 284: 283: 280: 279: 225:Aequorea victoria 205: 204: 201: 200: 196:structure summary 53:Aequorea victoria 51:Structure of the 16:(Redirected from 5544: 5458:Ribosome display 5394:Protein ontology 5240: 5233: 5226: 5217: 5093: 5065: 5045: 5025: 5024: 5022: 5021: 5011: 5005: 5004: 4992: 4986: 4985: 4957: 4951: 4950: 4943: 4937: 4936: 4926: 4890: 4884: 4879: 4873: 4872: 4870: 4868: 4859:. Archived from 4853: 4847: 4846: 4838: 4832: 4831: 4829: 4828: 4814: 4808: 4807: 4797: 4765: 4759: 4758: 4740: 4716: 4710: 4709: 4673: 4667: 4666: 4638: 4632: 4631: 4595: 4589: 4588: 4578: 4546: 4540: 4539: 4537: 4536: 4521: 4515: 4514: 4483:Nature Photonics 4478: 4472: 4471: 4461: 4429: 4423: 4422: 4394: 4388: 4387: 4350: 4344: 4343: 4333: 4301: 4295: 4294: 4284: 4274: 4242: 4236: 4235: 4225: 4201: 4195: 4194: 4184: 4152: 4146: 4145: 4101: 4095: 4094: 4066: 4060: 4059: 4019: 4013: 4012: 4002: 3962: 3956: 3955: 3945: 3935: 3911: 3905: 3904: 3871:(5588): 1873–7. 3860: 3854: 3853: 3828:(5496): 1585–8. 3817: 3811: 3810: 3774: 3768: 3767: 3757: 3747: 3715: 3709: 3708: 3679: 3673: 3672: 3644: 3638: 3637: 3609: 3603: 3602: 3592: 3582: 3558: 3552: 3551: 3519: 3513: 3512: 3483: 3474: 3473: 3463: 3431: 3425: 3424: 3388: 3382: 3381: 3371: 3354:(38): 10111–22. 3339: 3333: 3332: 3304: 3298: 3297: 3272:(6): 2657–2667. 3261: 3255: 3254: 3244: 3212: 3206: 3205: 3195: 3146: 3140: 3139: 3103: 3097: 3096: 3062: 3056: 3055: 3045: 3013: 3004: 3003: 2967: 2961: 2960: 2924: 2918: 2917: 2881: 2875: 2874: 2863: 2857: 2856: 2846: 2836: 2804: 2798: 2797: 2787: 2777: 2745: 2739: 2738: 2705:(5569): 913–16. 2694: 2688: 2687: 2677: 2668:(13): 13044–53. 2653: 2647: 2646: 2602: 2596: 2595: 2585: 2545: 2539: 2538: 2509: 2503: 2502: 2470: 2464: 2463: 2427: 2421: 2420: 2392: 2386: 2385: 2375: 2351: 2345: 2344: 2343: 2339: 2332: 2326: 2325: 2306:10.1038/373663b0 2283: 2274: 2268: 2267: 2249: 2243: 2242: 2223:10.1038/nmeth819 2208: 2199: 2193: 2192: 2182: 2172: 2155:(6): 2306–2311. 2134: 2123: 2122: 2104: 2078: 2069: 2058: 2057: 2024:(5280): 1392–5. 2013: 2002: 2001: 1991: 1965: 1959: 1958: 1912: 1906: 1905: 1875: 1869: 1868: 1838: 1832: 1831: 1801: 1795: 1794: 1760: 1754: 1753: 1719: 1713: 1712: 1684: 1678: 1677: 1667: 1635: 1629: 1628: 1593:Salih A (2019). 1590: 1584: 1583: 1557: 1548: 1535: 1534: 1506: 1500: 1499: 1466:(5280): 1392–5. 1455: 1128: 1103: 1079: 1066:(highlighted as 1050: 1049: 874:phycobiliprotein 833:. These include 760:Most species of 417:proof of concept 360:visible spectrum 253: 227: 207: 144: 49: 37: 21: 5552: 5551: 5547: 5546: 5545: 5543: 5542: 5541: 5532:Bioluminescence 5522:Protein imaging 5507:Protein methods 5497: 5496: 5495: 5490: 5467: 5431: 5427:Secretion assay 5403: 5360: 5254: 5244: 5164:Osamu Shimomura 5130: 5129: 5128: 5113: 5112: 5108: 5101: 5096: 5082: 5069: 5054: 5037: 5033: 5031:Further reading 5028: 5019: 5017: 5013: 5012: 5008: 4994: 4993: 4989: 4959: 4958: 4954: 4945: 4944: 4940: 4892: 4891: 4887: 4880: 4876: 4866: 4864: 4855: 4854: 4850: 4840: 4839: 4835: 4826: 4824: 4816: 4815: 4811: 4767: 4766: 4762: 4718: 4717: 4713: 4675: 4674: 4670: 4643:New Phytologist 4640: 4639: 4635: 4612:10.1038/nbt1044 4597: 4596: 4592: 4548: 4547: 4543: 4534: 4532: 4523: 4522: 4518: 4480: 4479: 4475: 4431: 4430: 4426: 4396: 4395: 4391: 4352: 4351: 4347: 4316:(20): 1384–96. 4303: 4302: 4298: 4244: 4243: 4239: 4203: 4202: 4198: 4154: 4153: 4149: 4112:(7166): 56–62. 4103: 4102: 4098: 4068: 4067: 4063: 4021: 4020: 4016: 3964: 3963: 3959: 3913: 3912: 3908: 3862: 3861: 3857: 3819: 3818: 3814: 3776: 3775: 3771: 3717: 3716: 3712: 3681: 3680: 3676: 3646: 3645: 3641: 3611: 3610: 3606: 3560: 3559: 3555: 3521: 3520: 3516: 3485: 3484: 3477: 3433: 3432: 3428: 3390: 3389: 3385: 3341: 3340: 3336: 3306: 3305: 3301: 3263: 3262: 3258: 3214: 3213: 3209: 3148: 3147: 3143: 3105: 3104: 3100: 3085: 3064: 3063: 3059: 3015: 3014: 3007: 2984:10.1038/nbt1293 2969: 2968: 2964: 2926: 2925: 2921: 2883: 2882: 2878: 2865: 2864: 2860: 2806: 2805: 2801: 2747: 2746: 2742: 2696: 2695: 2691: 2655: 2654: 2650: 2613:(6689): 192–5. 2604: 2603: 2599: 2547: 2546: 2542: 2511: 2510: 2506: 2472: 2471: 2467: 2444:10.1038/nbt1172 2429: 2428: 2424: 2394: 2393: 2389: 2353: 2352: 2348: 2341: 2334: 2333: 2329: 2292:(6516): 663–4. 2281: 2276: 2275: 2271: 2264: 2251: 2250: 2246: 2206: 2201: 2200: 2196: 2136: 2135: 2126: 2087:(10): 1246–51. 2076: 2071: 2070: 2061: 2015: 2014: 2005: 1982:(2–3): 277–80. 1967: 1966: 1962: 1925:(5148): 802–5. 1914: 1913: 1909: 1877: 1876: 1872: 1851:(12): 2656–62. 1840: 1839: 1835: 1803: 1802: 1798: 1783: 1762: 1761: 1757: 1742: 1721: 1720: 1716: 1686: 1685: 1681: 1637: 1636: 1632: 1617: 1592: 1591: 1587: 1555: 1550: 1549: 1538: 1517:(17): 3448–53. 1508: 1507: 1503: 1457: 1456: 1452: 1448: 1421: 1412:San Juan Island 1404:Steel Jellyfish 1375: 1328: 1326:Transgenic pets 1315: 1306: 1287:viability assay 1188:fusion proteins 1180: 1174: 1165: 1146: 1144:Reporter assays 1141: 1122: 1115: 1095: 1071: 1003: 878:allophycocyanin 858:A new class of 819: 780:photoprotective 732:bioluminescence 706:Live lancelet ( 700: 672: 571: 569:GFP derivatives 510:Douglas Prasher 503:interacts with 493:Osamu Shimomura 473: 449: 429:Osamu Shimomura 352:excitation peak 223: 56: 35: 32:Pembrey Airport 28: 23: 22: 15: 12: 11: 5: 5550: 5548: 5540: 5539: 5534: 5529: 5524: 5519: 5514: 5509: 5499: 5498: 5492: 5491: 5489: 5488: 5483: 5477: 5475: 5469: 5468: 5466: 5465: 5460: 5455: 5450: 5445: 5439: 5437: 5433: 5432: 5430: 5429: 5424: 5419: 5413: 5411: 5405: 5404: 5402: 5401: 5396: 5391: 5386: 5381: 5376: 5370: 5368: 5366:Bioinformatics 5362: 5361: 5359: 5358: 5353: 5348: 5343: 5338: 5333: 5328: 5323: 5318: 5313: 5304: 5299: 5290: 5285: 5280: 5275: 5270: 5264: 5262: 5256: 5255: 5245: 5243: 5242: 5235: 5228: 5220: 5214: 5213: 5192: 5186: 5180: 5174: 5172:Roger Y. Tsien 5168:Martin Chalfie 5157: 5152: 5147: 5141: 5136: 5127: 5126: 5121: 5115: 5114: 5103: 5102: 5100: 5099:External links 5097: 5095: 5094: 5080: 5067: 5052: 5034: 5032: 5029: 5027: 5026: 5006: 4997:Physik Journal 4987: 4952: 4938: 4903:Nature Methods 4885: 4874: 4848: 4833: 4809: 4760: 4711: 4668: 4649:(3): 461–468. 4633: 4590: 4541: 4516: 4489:(7): 406–410. 4473: 4424: 4405:(4): 360–369. 4389: 4345: 4296: 4257:(16): 9280–5. 4237: 4196: 4167:(1–4): 53–67. 4147: 4096: 4077:(12): 605–13. 4061: 4014: 3957: 3906: 3855: 3812: 3779:Nature Methods 3769: 3710: 3674: 3639: 3604: 3553: 3514: 3475: 3446:(3): 539–551. 3426: 3399:(3): 1103–63. 3383: 3334: 3299: 3256: 3227:(2): 111–129. 3207: 3141: 3098: 3083: 3057: 3022:Nature Methods 3005: 2978:(4): 443–445. 2962: 2919: 2898:10.1038/nbt778 2876: 2858: 2799: 2740: 2689: 2648: 2597: 2540: 2504: 2465: 2422: 2403:(1): 121–127. 2387: 2346: 2327: 2269: 2262: 2244: 2211:Nature Methods 2194: 2124: 2059: 2003: 1960: 1907: 1870: 1833: 1796: 1781: 1755: 1740: 1714: 1679: 1630: 1615: 1585: 1536: 1501: 1449: 1447: 1444: 1443: 1442: 1437: 1432: 1427: 1420: 1417: 1374: 1371: 1327: 1324: 1314: 1311: 1305: 1302: 1262:, tracking of 1176:Main article: 1173: 1170: 1164: 1161: 1145: 1142: 1140: 1137: 1114: 1111: 1108: 1107: 1092:ribbon diagram 1083: 1068:ball-and-stick 1002: 999: 924:stoichiometric 922:and produce a 864:cyanobacterial 818: 815: 699: 696: 684:cell membranes 680:palmitoylation 671: 668: 662:properties of 570: 567: 522:Martin Chalfie 472: 469: 448: 445: 433:Martin Chalfie 425:Roger Y. Tsien 282: 281: 278: 277: 272: 268: 267: 262: 258: 257: 249: 248: 243: 237: 236: 233: 229: 228: 221: 217: 216: 212: 211: 203: 202: 199: 198: 193: 187: 186: 173: 167: 166: 156: 149: 148: 140: 139: 126: 120: 119: 114: 108: 107: 102: 96: 95: 90: 83: 82: 77: 71: 70: 67: 63: 62: 58: 57: 50: 42: 41: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 5549: 5538: 5535: 5533: 5530: 5528: 5525: 5523: 5520: 5518: 5515: 5513: 5510: 5508: 5505: 5504: 5502: 5487: 5484: 5482: 5479: 5478: 5476: 5474: 5470: 5464: 5463:Yeast display 5461: 5459: 5456: 5454: 5453:Phage display 5451: 5449: 5446: 5444: 5441: 5440: 5438: 5434: 5428: 5425: 5423: 5422:Protein assay 5420: 5418: 5415: 5414: 5412: 5410: 5406: 5400: 5397: 5395: 5392: 5390: 5387: 5385: 5382: 5380: 5377: 5375: 5372: 5371: 5369: 5367: 5363: 5357: 5354: 5352: 5349: 5347: 5344: 5342: 5339: 5337: 5334: 5332: 5329: 5327: 5324: 5322: 5319: 5317: 5314: 5312: 5308: 5305: 5303: 5300: 5298: 5294: 5291: 5289: 5286: 5284: 5281: 5279: 5276: 5274: 5271: 5269: 5266: 5265: 5263: 5261: 5257: 5252: 5248: 5241: 5236: 5234: 5229: 5227: 5222: 5221: 5218: 5211: 5207: 5206: 5201: 5197: 5193: 5190: 5187: 5184: 5181: 5178: 5175: 5173: 5169: 5165: 5161: 5158: 5156: 5153: 5151: 5148: 5146: 5142: 5140: 5137: 5135: 5132: 5131: 5125: 5122: 5120: 5117: 5116: 5111: 5106: 5098: 5091: 5087: 5083: 5077: 5073: 5068: 5063: 5059: 5055: 5049: 5044: 5043: 5036: 5035: 5030: 5016: 5010: 5007: 5002: 4998: 4991: 4988: 4983: 4979: 4975: 4971: 4967: 4963: 4956: 4953: 4948: 4942: 4939: 4934: 4930: 4925: 4920: 4916: 4912: 4909:(10): 853–9. 4908: 4904: 4900: 4896: 4889: 4886: 4883: 4878: 4875: 4862: 4858: 4852: 4849: 4844: 4837: 4834: 4823: 4819: 4813: 4810: 4805: 4801: 4796: 4791: 4787: 4783: 4779: 4775: 4771: 4764: 4761: 4756: 4752: 4748: 4744: 4739: 4734: 4731:(4): 455–63. 4730: 4726: 4722: 4715: 4712: 4707: 4703: 4699: 4695: 4691: 4687: 4683: 4679: 4672: 4669: 4664: 4660: 4656: 4652: 4648: 4644: 4637: 4634: 4629: 4625: 4621: 4617: 4613: 4609: 4605: 4601: 4594: 4591: 4586: 4582: 4577: 4572: 4568: 4564: 4561:(2): 141–52. 4560: 4556: 4552: 4545: 4542: 4531: 4527: 4520: 4517: 4512: 4508: 4504: 4500: 4496: 4492: 4488: 4484: 4477: 4474: 4469: 4465: 4460: 4455: 4451: 4447: 4444:(4): 559–66. 4443: 4439: 4435: 4428: 4425: 4420: 4416: 4412: 4408: 4404: 4400: 4393: 4390: 4385: 4381: 4377: 4373: 4369: 4365: 4362:(5): 745–63. 4361: 4357: 4349: 4346: 4341: 4337: 4332: 4327: 4323: 4319: 4315: 4311: 4307: 4300: 4297: 4292: 4288: 4283: 4278: 4273: 4268: 4264: 4260: 4256: 4252: 4248: 4241: 4238: 4233: 4229: 4224: 4219: 4216:(6): 977–85. 4215: 4211: 4207: 4200: 4197: 4192: 4188: 4183: 4178: 4174: 4170: 4166: 4162: 4158: 4151: 4148: 4143: 4139: 4135: 4131: 4127: 4123: 4119: 4115: 4111: 4107: 4100: 4097: 4092: 4088: 4084: 4080: 4076: 4072: 4065: 4062: 4057: 4053: 4049: 4045: 4041: 4037: 4034:(6): 927–38. 4033: 4029: 4025: 4018: 4015: 4010: 4006: 4001: 3996: 3992: 3988: 3984: 3980: 3976: 3972: 3968: 3961: 3958: 3953: 3949: 3944: 3939: 3934: 3929: 3925: 3921: 3917: 3910: 3907: 3902: 3898: 3894: 3890: 3886: 3882: 3878: 3874: 3870: 3866: 3859: 3856: 3851: 3847: 3843: 3839: 3835: 3831: 3827: 3823: 3816: 3813: 3808: 3804: 3800: 3796: 3792: 3788: 3785:(12): 902–4. 3784: 3780: 3773: 3770: 3765: 3761: 3756: 3751: 3746: 3741: 3737: 3733: 3729: 3725: 3721: 3714: 3711: 3706: 3702: 3698: 3694: 3690: 3686: 3678: 3675: 3670: 3666: 3662: 3658: 3655:(4): 1331–6. 3654: 3650: 3643: 3640: 3635: 3631: 3627: 3623: 3619: 3615: 3608: 3605: 3600: 3596: 3591: 3586: 3581: 3576: 3572: 3568: 3564: 3557: 3554: 3549: 3545: 3541: 3537: 3533: 3529: 3525: 3518: 3515: 3510: 3506: 3502: 3498: 3494: 3490: 3482: 3480: 3476: 3471: 3467: 3462: 3457: 3453: 3449: 3445: 3441: 3437: 3430: 3427: 3422: 3418: 3414: 3410: 3406: 3402: 3398: 3394: 3387: 3384: 3379: 3375: 3370: 3365: 3361: 3357: 3353: 3349: 3345: 3338: 3335: 3330: 3326: 3322: 3318: 3314: 3310: 3303: 3300: 3295: 3291: 3287: 3283: 3279: 3275: 3271: 3267: 3260: 3257: 3252: 3248: 3243: 3238: 3234: 3230: 3226: 3222: 3218: 3211: 3208: 3203: 3199: 3194: 3189: 3185: 3181: 3177: 3173: 3169: 3165: 3161: 3157: 3153: 3145: 3142: 3137: 3133: 3129: 3125: 3121: 3117: 3114:(1): 509–44. 3113: 3109: 3102: 3099: 3094: 3090: 3086: 3080: 3076: 3072: 3068: 3061: 3058: 3053: 3049: 3044: 3039: 3035: 3031: 3027: 3023: 3019: 3012: 3010: 3006: 3001: 2997: 2993: 2989: 2985: 2981: 2977: 2973: 2966: 2963: 2958: 2954: 2950: 2946: 2942: 2938: 2934: 2930: 2923: 2920: 2915: 2911: 2907: 2903: 2899: 2895: 2891: 2887: 2880: 2877: 2872: 2868: 2862: 2859: 2854: 2850: 2845: 2840: 2835: 2830: 2826: 2822: 2819:(7): e11517. 2818: 2814: 2810: 2803: 2800: 2795: 2791: 2786: 2781: 2776: 2771: 2767: 2763: 2759: 2755: 2751: 2744: 2741: 2736: 2732: 2728: 2724: 2720: 2716: 2712: 2708: 2704: 2700: 2693: 2690: 2685: 2681: 2676: 2671: 2667: 2663: 2659: 2652: 2649: 2644: 2640: 2636: 2632: 2628: 2627:10.1038/28190 2624: 2620: 2616: 2612: 2608: 2601: 2598: 2593: 2589: 2584: 2579: 2575: 2571: 2567: 2563: 2559: 2555: 2551: 2544: 2541: 2536: 2532: 2528: 2524: 2520: 2516: 2508: 2505: 2500: 2496: 2492: 2488: 2484: 2480: 2476: 2469: 2466: 2461: 2457: 2453: 2449: 2445: 2441: 2437: 2433: 2426: 2423: 2418: 2414: 2410: 2406: 2402: 2398: 2391: 2388: 2383: 2379: 2374: 2369: 2365: 2361: 2357: 2350: 2347: 2337: 2331: 2328: 2323: 2319: 2315: 2311: 2307: 2303: 2299: 2295: 2291: 2287: 2280: 2273: 2270: 2265: 2259: 2255: 2248: 2245: 2240: 2236: 2232: 2228: 2224: 2220: 2217:(12): 905–9. 2216: 2212: 2205: 2198: 2195: 2190: 2186: 2181: 2176: 2171: 2166: 2162: 2158: 2154: 2150: 2149: 2144: 2142: 2133: 2131: 2129: 2125: 2120: 2116: 2112: 2108: 2103: 2098: 2094: 2090: 2086: 2082: 2075: 2068: 2066: 2064: 2060: 2055: 2051: 2047: 2043: 2039: 2035: 2031: 2027: 2023: 2019: 2012: 2010: 2008: 2004: 1999: 1995: 1990: 1985: 1981: 1977: 1976: 1971: 1964: 1961: 1956: 1952: 1948: 1944: 1940: 1936: 1932: 1928: 1924: 1920: 1919: 1911: 1908: 1903: 1899: 1895: 1891: 1888:(2): 229–33. 1887: 1883: 1882: 1874: 1871: 1866: 1862: 1858: 1854: 1850: 1846: 1845: 1837: 1834: 1829: 1825: 1821: 1817: 1814:(3): 223–39. 1813: 1809: 1808: 1800: 1797: 1792: 1788: 1784: 1778: 1774: 1770: 1766: 1759: 1756: 1751: 1747: 1743: 1741:1-59259-086-1 1737: 1733: 1729: 1725: 1718: 1715: 1710: 1706: 1702: 1698: 1694: 1690: 1683: 1680: 1675: 1671: 1666: 1661: 1657: 1653: 1650:(4): 338–69. 1649: 1645: 1641: 1634: 1631: 1626: 1622: 1618: 1616:9781351129404 1612: 1608: 1604: 1600: 1596: 1589: 1586: 1581: 1577: 1573: 1569: 1565: 1561: 1554: 1547: 1545: 1543: 1541: 1537: 1532: 1528: 1524: 1520: 1516: 1512: 1505: 1502: 1497: 1493: 1489: 1485: 1481: 1477: 1473: 1469: 1465: 1461: 1454: 1451: 1445: 1441: 1438: 1436: 1433: 1431: 1428: 1426: 1423: 1422: 1418: 1413: 1409: 1405: 1401: 1397: 1393: 1391: 1387: 1384:in 1962, the 1383: 1379: 1372: 1370: 1368: 1364: 1360: 1356: 1352: 1348: 1344: 1340: 1332: 1325: 1323: 1321: 1312: 1310: 1303: 1301: 1298: 1294: 1292: 1288: 1284: 1279: 1275: 1273: 1269: 1265: 1261: 1258: 1254: 1250: 1249: 1244: 1243: 1237: 1234: 1229: 1227: 1223: 1222: 1217: 1212: 1208: 1205: 1201: 1197: 1189: 1184: 1179: 1171: 1169: 1162: 1160: 1158: 1153: 1151: 1150:reporter gene 1143: 1138: 1136: 1133: 1129: 1127: 1120: 1112: 1102: 1098: 1093: 1088: 1084: 1078: 1074: 1069: 1065: 1061: 1056: 1052: 1044: 1040: 1037: 1033: 1029: 1025: 1020: 1016: 1012: 1008: 1000: 998: 991: 987: 985: 981: 977: 974:derived from 973: 969: 965: 964:quantum yield 961: 957: 953: 949: 946:and uses the 945: 941: 937: 933: 929: 925: 921: 917: 913: 909: 906:, known as a 905: 901: 898: 894: 891: 887: 883: 879: 875: 871: 870: 865: 861: 856: 848: 844: 842: 838: 836: 832: 823: 816: 814: 813: 809: 808:quantum yield 805: 804: 799: 795: 791: 788: 783: 781: 777: 773: 769: 768: 763: 758: 756: 755:Friday Harbor 751: 746: 741: 737: 733: 726: 725: 722: 716: 709: 704: 697: 695: 693: 689: 685: 681: 677: 669: 667: 665: 661: 657: 653: 648: 646: 645:synaptobrevin 640: 638: 632: 629: 620: 616: 613: 607: 604: 602: 601:quantum yield 598: 593: 589: 585: 575: 568: 566: 564: 559: 555: 551: 550: 544: 541: 540: 535: 534: 529: 528: 523: 519: 515: 511: 506: 502: 498: 494: 490: 486: 482: 478: 470: 463: 457: 453: 446: 444: 440: 438: 434: 430: 426: 422: 418: 414: 410: 406: 402: 398: 393: 391: 387: 383: 379: 375: 374: 369: 365: 364:quantum yield 361: 357: 353: 349: 346:The GFP from 344: 342: 338: 334: 330: 326: 322: 318: 317: 313: 309: 305: 301: 297: 293: 289: 276: 273: 269: 266: 263: 259: 254: 250: 247: 244: 242: 238: 234: 230: 226: 222: 218: 213: 208: 197: 194: 192: 188: 185: 181: 177: 174: 172: 168: 164: 160: 157: 154: 150: 145: 141: 138: 134: 130: 127: 125: 121: 118: 115: 113: 109: 106: 103: 101: 97: 94: 91: 88: 84: 81: 78: 76: 72: 68: 64: 59: 54: 48: 43: 38: 33: 19: 5517:Cell imaging 5448:mRNA display 5417:Enzyme assay 5278:Western blot 5272: 5260:Experimental 5203: 5109: 5071: 5041: 5018:. 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