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Geologists have debated about what caused the slide debris to travel the distance it did . The " air cushion " theory , an early hypothesis , postulated that a layer of air was trapped between the mass of rock and the mountain , which caused the rock to move a greater distance than would otherwise be expected . " Acou...
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= = Legends = =
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Numerous legends and misconceptions were spawned in the aftermath of the slide . The entire town of Frank was claimed to have been buried , though much of the town itself was unscathed . The belief that a branch of the Union Bank of Canada had been buried with as much as $ 500 @,@ 000 persisted for many years . The ba...
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Several people , telling amazing stories to those who would listen , passed themselves off as the " sole survivor " in the years following the slide . The most common such tale is that of an infant girl said to have been the only survivor of the slide . Her real name unknown , the girl was called " Frankie Slide " . S...
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= = Legacy = =
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Curious sightseers flocked to the site of the slide within the day of the disaster . It has remained a popular tourist destination , in part due to its proximity to the Crowsnest Highway . The province built a roadside turnout in 1941 to accommodate the traffic . Town boosters unsuccessfully sought to have the site de...
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Though Frank recovered from the slide and achieved a peak population of 1 @,@ 000 shortly thereafter , the closure of the mine resulted in a longstanding decline in population . Frank ceased to be an independent community in 1979 when it was amalgamated into the Municipality of Crowsnest Pass along with the neighbouri...
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= Protein =
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Proteins ( / ˈproʊˌtiːnz / or / ˈproʊti.ᵻnz / ) are large biomolecules , or macromolecules , consisting of one or more long chains of amino acid residues . Proteins perform a vast array of functions within organisms , including catalysing metabolic reactions , DNA replication , responding to stimuli , and transporting...
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A linear chain of amino acid residues is called a polypeptide . A protein contains at least one long polypeptide . Short polypeptides , containing less than 20 @-@ 30 residues , are rarely considered to be proteins and are commonly called peptides , or sometimes oligopeptides . The individual amino acid residues are b...
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Once formed , proteins only exist for a certain period of time and are then degraded and recycled by the cell 's machinery through the process of protein turnover . A protein 's lifespan is measured in terms of its half @-@ life and covers a wide range . They can exist for minutes or years with an average lifespan of ...
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Like other biological macromolecules such as polysaccharides and nucleic acids , proteins are essential parts of organisms and participate in virtually every process within cells . Many proteins are enzymes that catalyse biochemical reactions and are vital to metabolism . Proteins also have structural or mechanical fu...
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Proteins may be purified from other cellular components using a variety of techniques such as ultracentrifugation , precipitation , electrophoresis , and chromatography ; the advent of genetic engineering has made possible a number of methods to facilitate purification . Methods commonly used to study protein structur...
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= = Biochemistry = =
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Most proteins consist of linear polymers built from series of up to 20 different L @-@ α @-@ amino acids . All proteinogenic amino acids possess common structural features , including an α @-@ carbon to which an amino group , a carboxyl group , and a variable side chain are bonded . Only proline differs from this basi...
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The peptide bond has two resonance forms that contribute some double @-@ bond character and inhibit rotation around its axis , so that the alpha carbons are roughly coplanar . The other two dihedral angles in the peptide bond determine the local shape assumed by the protein backbone . The end of the protein with a fre...
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= = = Abundance in cells = = =
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It has been estimated that average @-@ sized bacteria contain about 2 million proteins per cell ( e.g. E. coli and Staphylococcus aureus ) . Smaller bacteria , such as Mycoplasma or spirochetes contain fewer molecules , namely on the order of 50 @,@ 000 to 1 million . By contrast , eukaryotic cells are larger and thus...
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Importantly , the concentration of individual proteins ranges from a few molecules per cell to hundreds of thousands . In fact , about a third of all proteins is not produced in most cells or only induced under certain circumstances . For instance , of the 20 @,@ 000 or so proteins encoded by the human genome only 6 @...
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= = Synthesis = =
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= = = Biosynthesis = = =
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Proteins are assembled from amino acids using information encoded in genes . Each protein has its own unique amino acid sequence that is specified by the nucleotide sequence of the gene encoding this protein . The genetic code is a set of three @-@ nucleotide sets called codons and each three @-@ nucleotide combinatio...
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The process of synthesizing a protein from an mRNA template is known as translation . The mRNA is loaded onto the ribosome and is read three nucleotides at a time by matching each codon to its base pairing anticodon located on a transfer RNA molecule , which carries the amino acid corresponding to the codon it recogni...
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The size of a synthesized protein can be measured by the number of amino acids it contains and by its total molecular mass , which is normally reported in units of daltons ( synonymous with atomic mass units ) , or the derivative unit kilodalton ( kDa ) . Yeast proteins are on average 466 amino acids long and 53 kDa i...
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= = = Chemical synthesis = = =
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Short proteins can also be synthesized chemically by a family of methods known as peptide synthesis , which rely on organic synthesis techniques such as chemical ligation to produce peptides in high yield . Chemical synthesis allows for the introduction of non @-@ natural amino acids into polypeptide chains , such as ...
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= = Structure = =
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Most proteins fold into unique 3 @-@ dimensional structures . The shape into which a protein naturally folds is known as its native conformation . Although many proteins can fold unassisted , simply through the chemical properties of their amino acids , others require the aid of molecular chaperones to fold into their...
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Primary structure : the amino acid sequence . A protein is a polyamide .
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Secondary structure : regularly repeating local structures stabilized by hydrogen bonds . The most common examples are the α @-@ helix , β @-@ sheet and turns . Because secondary structures are local , many regions of different secondary structure can be present in the same protein molecule .
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Tertiary structure : the overall shape of a single protein molecule ; the spatial relationship of the secondary structures to one another . Tertiary structure is generally stabilized by nonlocal interactions , most commonly the formation of a hydrophobic core , but also through salt bridges , hydrogen bonds , disulfid...
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Quaternary structure : the structure formed by several protein molecules ( polypeptide chains ) , usually called protein subunits in this context , which function as a single protein complex .
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Proteins are not entirely rigid molecules . In addition to these levels of structure , proteins may shift between several related structures while they perform their functions . In the context of these functional rearrangements , these tertiary or quaternary structures are usually referred to as " conformations " , an...
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Proteins can be informally divided into three main classes , which correlate with typical tertiary structures : globular proteins , fibrous proteins , and membrane proteins . Almost all globular proteins are soluble and many are enzymes . Fibrous proteins are often structural , such as collagen , the major component o...
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A special case of intramolecular hydrogen bonds within proteins , poorly shielded from water attack and hence promoting their own dehydration , are called dehydrons .
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= = = Structure determination = = =
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Discovering the tertiary structure of a protein , or the quaternary structure of its complexes , can provide important clues about how the protein performs its function . Common experimental methods of structure determination include X @-@ ray crystallography and NMR spectroscopy , both of which can produce informatio...
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Many more gene sequences are known than protein structures . Further , the set of solved structures is biased toward proteins that can be easily subjected to the conditions required in X @-@ ray crystallography , one of the major structure determination methods . In particular , globular proteins are comparatively eas...
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= = Cellular functions = =
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Proteins are the chief actors within the cell , said to be carrying out the duties specified by the information encoded in genes . With the exception of certain types of RNA , most other biological molecules are relatively inert elements upon which proteins act . Proteins make up half the dry weight of an Escherichia ...
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The chief characteristic of proteins that also allows their diverse set of functions is their ability to bind other molecules specifically and tightly . The region of the protein responsible for binding another molecule is known as the binding site and is often a depression or " pocket " on the molecular surface . Thi...
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Proteins can bind to other proteins as well as to small @-@ molecule substrates . When proteins bind specifically to other copies of the same molecule , they can oligomerize to form fibrils ; this process occurs often in structural proteins that consist of globular monomers that self @-@ associate to form rigid fibers...
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= = = Enzymes = = =
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The best @-@ known role of proteins in the cell is as enzymes , which catalyse chemical reactions . Enzymes are usually highly specific and accelerate only one or a few chemical reactions . Enzymes carry out most of the reactions involved in metabolism , as well as manipulating DNA in processes such as DNA replication...
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The molecules bound and acted upon by enzymes are called substrates . Although enzymes can consist of hundreds of amino acids , it is usually only a small fraction of the residues that come in contact with the substrate , and an even smaller fraction — three to four residues on average — that are directly involved in ...
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Dirigent proteins are members of a class of proteins that dictate the stereochemistry of a compound synthesized by other enzymes .
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= = = Cell signaling and ligand binding = = =
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Many proteins are involved in the process of cell signaling and signal transduction . Some proteins , such as insulin , are extracellular proteins that transmit a signal from the cell in which they were synthesized to other cells in distant tissues . Others are membrane proteins that act as receptors whose main functi...
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Antibodies are protein components of an adaptive immune system whose main function is to bind antigens , or foreign substances in the body , and target them for destruction . Antibodies can be secreted into the extracellular environment or anchored in the membranes of specialized B cells known as plasma cells . Wherea...
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Many ligand transport proteins bind particular small biomolecules and transport them to other locations in the body of a multicellular organism . These proteins must have a high binding affinity when their ligand is present in high concentrations , but must also release the ligand when it is present at low concentrati...
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Transmembrane proteins can also serve as ligand transport proteins that alter the permeability of the cell membrane to small molecules and ions . The membrane alone has a hydrophobic core through which polar or charged molecules cannot diffuse . Membrane proteins contain internal channels that allow such molecules to ...
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= = = Structural proteins = = =
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Structural proteins confer stiffness and rigidity to otherwise @-@ fluid biological components . Most structural proteins are fibrous proteins ; for example , collagen and elastin are critical components of connective tissue such as cartilage , and keratin is found in hard or filamentous structures such as hair , nail...
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Other proteins that serve structural functions are motor proteins such as myosin , kinesin , and dynein , which are capable of generating mechanical forces . These proteins are crucial for cellular motility of single celled organisms and the sperm of many multicellular organisms which reproduce sexually . They also ge...
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= = Methods of study = =
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The activities and structures of proteins may be examined in vitro , in vivo , and in silico . In vitro studies of purified proteins in controlled environments are useful for learning how a protein carries out its function : for example , enzyme kinetics studies explore the chemical mechanism of an enzyme 's catalytic...
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= = = Protein purification = = =
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To perform in vitro analysis , a protein must be purified away from other cellular components . This process usually begins with cell lysis , in which a cell 's membrane is disrupted and its internal contents released into a solution known as a crude lysate . The resulting mixture can be purified using ultracentrifuga...
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For natural proteins , a series of purification steps may be necessary to obtain protein sufficiently pure for laboratory applications . To simplify this process , genetic engineering is often used to add chemical features to proteins that make them easier to purify without affecting their structure or activity . Here...
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= = = Cellular localization = = =
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The study of proteins in vivo is often concerned with the synthesis and localization of the protein within the cell . Although many intracellular proteins are synthesized in the cytoplasm and membrane @-@ bound or secreted proteins in the endoplasmic reticulum , the specifics of how proteins are targeted to specific o...
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Other methods for elucidating the cellular location of proteins requires the use of known compartmental markers for regions such as the ER , the Golgi , lysosomes or vacuoles , mitochondria , chloroplasts , plasma membrane , etc . With the use of fluorescently tagged versions of these markers or of antibodies to known...
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Other possibilities exist , as well . For example , immunohistochemistry usually utilizes an antibody to one or more proteins of interest that are conjugated to enzymes yielding either luminescent or chromogenic signals that can be compared between samples , allowing for localization information . Another applicable t...
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Finally , the gold @-@ standard method of cellular localization is immunoelectron microscopy . This technique also uses an antibody to the protein of interest , along with classical electron microscopy techniques . The sample is prepared for normal electron microscopic examination , and then treated with an antibody t...
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Through another genetic engineering application known as site @-@ directed mutagenesis , researchers can alter the protein sequence and hence its structure , cellular localization , and susceptibility to regulation . This technique even allows the incorporation of unnatural amino acids into proteins , using modified t...
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= = = Proteomics = = =
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The total complement of proteins present at a time in a cell or cell type is known as its proteome , and the study of such large @-@ scale data sets defines the field of proteomics , named by analogy to the related field of genomics . Key experimental techniques in proteomics include 2D electrophoresis , which allows ...
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= = = Bioinformatics = = =
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A vast array of computational methods have been developed to analyze the structure , function , and evolution of proteins .
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The development of such tools has been driven by the large amount of genomic and proteomic data available for a variety of organisms , including the human genome . It is simply impossible to study all proteins experimentally , hence only a few are subjected to laboratory experiments while computational tools are used ...
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= = = = Structure prediction and simulation = = = =
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Complementary to the field of structural genomics , protein structure prediction seeks to develop efficient ways to provide plausible models for proteins whose structures have not yet been determined experimentally . The most successful type of structure prediction , known as homology modeling , relies on the existenc...
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The processes of protein folding and binding can be simulated using such technique as molecular mechanics , in particular , molecular dynamics and Monte Carlo , which increasingly take advantage of parallel and distributed computing ( Folding @ home project ; molecular modeling on GPU ) . The folding of small α @-@ he...
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= = = = Protein disorder and unstructure prediction = = = =
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Many proteins ( in Eucaryota ~ 33 % ) contain large unstructured but biologically functional segments and can be classified as intrinsically disordered proteins . Predicting and analysing protein disorder is , therefore , an important part of protein structure characterisation .
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= = Nutrition = =
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Most microorganisms and plants can biosynthesize all 20 standard amino acids , while animals ( including humans ) must obtain some of the amino acids from the diet . The amino acids that an organism cannot synthesize on its own are referred to as essential amino acids . Key enzymes that synthesize certain amino acids ...
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In animals , amino acids are obtained through the consumption of foods containing protein . Ingested proteins are then broken down into amino acids through digestion , which typically involves denaturation of the protein through exposure to acid and hydrolysis by enzymes called proteases . Some ingested amino acids ar...
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= = History and etymology = =
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Proteins were recognized as a distinct class of biological molecules in the eighteenth century by Antoine Fourcroy and others , distinguished by the molecules ' ability to coagulate or flocculate under treatments with heat or acid . Noted examples at the time included albumin from egg whites , blood serum albumin , fi...
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Proteins were first described by the Dutch chemist Gerardus Johannes Mulder and named by the Swedish chemist Jöns Jacob Berzelius in 1838 . Mulder carried out elemental analysis of common proteins and found that nearly all proteins had the same empirical formula , C400H620N100O120P1S1 . He came to the erroneous conclu...
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Early nutritional scientists such as the German Carl von Voit believed that protein was the most important nutrient for maintaining the structure of the body , because it was generally believed that " flesh makes flesh . " Karl Heinrich Ritthausen extended known protein forms with the identification of glutamic acid ....
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The difficulty in purifying proteins in large quantities made them very difficult for early protein biochemists to study . Hence , early studies focused on proteins that could be purified in large quantities , e.g. , those of blood , egg white , various toxins , and digestive / metabolic enzymes obtained from slaughte...
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Linus Pauling is credited with the successful prediction of regular protein secondary structures based on hydrogen bonding , an idea first put forth by William Astbury in 1933 . Later work by Walter Kauzmann on denaturation , based partly on previous studies by Kaj Linderstrøm @-@ Lang , contributed an understanding o...
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The first protein to be sequenced was insulin , by Frederick Sanger , in 1949 . Sanger correctly determined the amino acid sequence of insulin , thus conclusively demonstrating that proteins consisted of linear polymers of amino acids rather than branched chains , colloids , or cyclols . He won the Nobel Prize for thi...
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The first protein structures to be solved were hemoglobin and myoglobin , by Max Perutz and Sir John Cowdery Kendrew , respectively , in 1958 . As of 2016 , the Protein Data Bank has over 115 @,@ 000 atomic @-@ resolution structures of proteins . In more recent times , cryo @-@ electron microscopy of large macromolecu...
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= = Textbooks = =
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= = = Databases and projects = = =
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The Protein Naming Utility
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Human Protein Atlas
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NCBI Entrez Protein database
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NCBI Protein Structure database
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Human Protein Reference Database
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Human Proteinpedia
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Folding @ Home ( Stanford University )
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Comparative Toxicogenomics Database curates protein – chemical interactions , as well as gene / protein – disease relationships and chemical @-@ disease relationships .
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Bioinformatic Harvester A Meta search engine ( 29 databases ) for gene and protein information .
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Protein Databank in Europe ( see also PDBeQuips , short articles and tutorials on interesting PDB structures )
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Research Collaboratory for Structural Bioinformatics ( see also Molecule of the Month , presenting short accounts on selected proteins from the PDB )
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Proteopedia – Life in 3D : rotatable , zoomable 3D model with wiki annotations for every known protein molecular structure .
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