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A gene is a molecular unit of heredity in a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains. Genes hold the information to build and maintain an organism's cells and pass genetic traits to offspring, although some organelles (e.g. mitochondria) are self-replicating and are not coded for by the organism's DNA. All organisms have many genes corresponding to various different biological traits, some of which are immediately visible, such as eye color or number of limbs, and some of which are not, such as blood type or increased risk for specific diseases, or the thousands of basic biochemical processes that comprise life.

A modern working definition of a gene is "''a locatable region of genomic sequence, corresponding to a unit of inheritance, which is associated with regulatory regions, transcribed regions, and or other functional sequence regions'' ". Colloquial usage of the term ''gene'' (e.g. "good genes", "hair color gene") may actually refer to an allele: a ''gene'' is the basic instruction, a sequence of nucleic acids (DNA or, in the case of certain viruses RNA), while an ''allele'' is one variant of that gene. Referring to having a gene for a trait, is no longer the scientifically accepted usage. In most cases, all people would have a gene for the trait in question, but certain people will have a specific allele of that gene, which results in the trait variant.

Physical definitions

RNA genes and genomes

When proteins are manufactured, the gene is first copied into RNA as an intermediate product. In other cases, the RNA molecules are the actual functional products. For example, RNAs known as ribozymes are capable of enzymatic function, and microRNA has a regulatory role. The DNA sequences from which such RNAs are transcribed are known as RNA genes.

Some viruses store their entire genomes in the form of RNA, and contain no DNA at all. Because they use RNA to store genes, their cellular hosts may synthesize their proteins as soon as they are infected and without the delay in waiting for transcription. On the other hand, RNA retroviruses, such as HIV, require the reverse transcription of their genome from RNA into DNA before their proteins can be synthesized. In 2006, French researchers came across a puzzling example of RNA-mediated inheritance in mice. Mice with a loss-of-function mutation in the gene Kit have white tails. Offspring of these mutants can have white tails despite having only normal Kit genes. The research team traced this effect back to mutated Kit RNA. While RNA is common as genetic storage material in viruses, in mammals in particular RNA inheritance has been observed very rarely.

Functional structure of a gene

The vast majority of living organisms encode their genes in long strands of DNA. DNA (deoxyribonucleic acid) consists of a chain made from four types of nucleotide subunits, each composed of: a five-carbon sugar (2'-deoxyribose), a phosphate group, and one of the four bases adenine, cytosine, guanine, and thymine. The most common form of DNA in a cell is in a double helix structure, in which two individual DNA strands twist around each other in a right-handed spiral. In this structure, the base pairing rules specify that guanine pairs with cytosine and adenine pairs with thymine. The base pairing between guanine and cytosine forms three hydrogen bonds, whereas the base pairing between adenine and thymine forms two hydrogen bonds. The two strands in a double helix must therefore be ''complementary'', that is, their bases must align such that the adenines of one strand are paired with the thymines of the other strand, and so on.

Due to the chemical composition of the pentose residues of the bases, DNA strands have directionality. One end of a DNA polymer contains an exposed hydroxyl group on the deoxyribose; this is known as the 3' end of the molecule. The other end contains an exposed phosphate group; this is the 5' end. The directionality of DNA is vitally important to many cellular processes, since double helices are necessarily directional (a strand running 5'-3' pairs with a complementary strand running 3'-5'), and processes such as DNA replication occur in only one direction. All nucleic acid synthesis in a cell occurs in the 5'-3' direction, because new monomers are added via a dehydration reaction that uses the exposed 3' hydroxyl as a nucleophile.

The expression of genes encoded in DNA begins by transcribing the gene into RNA, a second type of nucleic acid that is very similar to DNA, but whose monomers contain the sugar ribose rather than deoxyribose. RNA also contains the base uracil in place of thymine. RNA molecules are less stable than DNA and are typically single-stranded. Genes that encode proteins are composed of a series of three-nucleotide sequences called codons, which serve as the ''words'' in the genetic ''language''. The genetic code specifies the correspondence during protein translation between codons and amino acids. The genetic code is nearly the same for all known organisms.

All genes have regulatory regions in addition to regions that explicitly code for a protein or RNA product. A regulatory region shared by almost all genes is known as the promoter, which provides a position that is recognized by the transcription machinery when a gene is about to be transcribed and expressed. A gene can have more than one promoter, resulting in RNAs that differ in how far they extend in the 5' end. Although promoter regions have a consensus sequence that is the most common sequence at this position, some genes have "strong" promoters that bind the transcription machinery well, and others have "weak" promoters that bind poorly. These weak promoters usually permit a lower rate of transcription than the strong promoters, because the transcription machinery binds to them and initiates transcription less frequently. Other possible regulatory regions include enhancers, which can compensate for a weak promoter. Most regulatory regions are "upstream"—that is, before or toward the 5' end of the transcription initiation site. Eukaryotic promoter regions are much more complex and difficult to identify than prokaryotic promoters.

Many prokaryotic genes are organized into operons, or groups of genes whose products have related functions and which are transcribed as a unit. By contrast, eukaryotic genes are transcribed only one at a time, but may include long stretches of DNA called introns which are transcribed but never translated into protein (they are spliced out before translation). Splicing can also occur in prokaryotic genes, but is less common than in eukaryotes.

Chromosomes

The total complement of genes in an organism or cell is known as its genome, which may be stored on one or more chromosomes; the region of the chromosome at which a particular gene is located is called its locus. A chromosome consists of a single, very long DNA helix on which thousands of genes are encoded. Prokaryotes—bacteria and archaea—typically store their genomes on a single large, circular chromosome, sometimes supplemented by additional small circles of DNA called plasmids, which usually encode only a few genes and are easily transferable between individuals. For example, the genes for antibiotic resistance are usually encoded on bacterial plasmids and can be passed between individual cells, even those of different species, via horizontal gene transfer. Although some simple eukaryotes also possess plasmids with small numbers of genes, the majority of eukaryotic genes are stored on multiple linear chromosomes, which are packed within the nucleus in complex with storage proteins called histones. The manner in which DNA is stored on the histone, as well as chemical modifications of the histone itself, are regulatory mechanisms governing whether a particular region of DNA is accessible for gene expression. The ends of eukaryotic chromosomes are capped by long stretches of repetitive sequences called telomeres, which do not code for any gene product but are present to prevent degradation of coding and regulatory regions during DNA replication. The length of the telomeres tends to decrease each time the genome is replicated in preparation for cell division; the loss of telomeres has been proposed as an explanation for cellular senescence, or the loss of the ability to divide, and by extension for the aging process in organisms.

Whereas the chromosomes of prokaryotes are relatively gene-dense, those of eukaryotes often contain so-called "junk DNA", or regions of DNA that serve no obvious function. Simple single-celled eukaryotes have relatively small amounts of such DNA, whereas the genomes of complex multicellular organisms, including humans, contain an absolute majority of DNA without an identified function. However it now appears that, although protein-coding DNA makes up barely 2% of the human genome, about 80% of the bases in the genome may be expressed, so the term "junk DNA" may be a misnomer.

Gene expression

In all organisms, there are two major steps separating a protein-coding gene from its protein: First, the DNA on which the gene resides must be ''transcribed'' from DNA to messenger RNA (mRNA); and, second, it must be ''translated'' from mRNA to protein. RNA-coding genes must still go through the first step, but are not translated into protein. The process of producing a biologically functional molecule of either RNA or protein is called gene expression, and the resulting molecule itself is called a gene product.

Genetic code

The genetic code is the set of rules by which a gene is translated into a functional protein. Each gene consists of a specific sequence of nucleotides encoded in a DNA (or sometimes RNA) strand; a correspondence between nucleotides, the basic building blocks of genetic material, and amino acids, the basic building blocks of proteins, must be established for genes to be successfully translated into functional proteins. Sets of three nucleotides, known as codons, each correspond to a specific amino acid or to a signal; three codons are known as "stop codons" and, instead of specifying a new amino acid, alert the translation machinery that the end of the gene has been reached. There are 64 possible codons (four possible nucleotides at each of three positions, hence 4³ possible codons) and only 20 standard amino acids; hence the code is redundant and multiple codons can specify the same amino acid. The correspondence between codons and amino acids is nearly universal among all known living organisms.

Transcription

The process of genetic transcription produces a single-stranded RNA molecule known as messenger RNA, whose nucleotide sequence is complementary to the DNA from which it was transcribed. The DNA strand whose sequence matches that of the RNA is known as the coding strand and the strand from which the RNA was synthesized is the template strand. Transcription is performed by an enzyme called an RNA polymerase, which reads the template strand in the 3' to 5' direction and synthesizes the RNA from 5' to 3'. To initiate transcription, the polymerase first recognizes and binds a promoter region of the gene. Thus a major mechanism of gene regulation is the blocking or sequestering of the promoter region, either by tight binding by repressor molecules that physically block the polymerase, or by organizing the DNA so that the promoter region is not accessible.

In prokaryotes, transcription occurs in the cytoplasm; for very long transcripts, translation may begin at the 5' end of the RNA while the 3' end is still being transcribed. In eukaryotes, transcription necessarily occurs in the nucleus, where the cell's DNA is sequestered; the RNA molecule produced by the polymerase is known as the primary transcript and must undergo post-transcriptional modifications before being exported to the cytoplasm for translation. The splicing of introns present within the transcribed region is a modification unique to eukaryotes; alternative splicing mechanisms can result in mature transcripts from the same gene having different sequences and thus coding for different proteins. This is a major form of regulation in eukaryotic cells.

Translation

Translation is the process by which a mature mRNA molecule is used as a template for synthesizing a new protein. Translation is carried out by ribosomes, large complexes of RNA and protein responsible for carrying out the chemical reactions to add new amino acids to a growing polypeptide chain by the formation of peptide bonds. The genetic code is read three nucleotides at a time, in units called codons, via interactions with specialized RNA molecules called transfer RNA (tRNA). Each tRNA has three unpaired bases known as the anticodon that are complementary to the codon it reads; the tRNA is also covalently attached to the amino acid specified by the complementary codon. When the tRNA binds to its complementary codon in an mRNA strand, the ribosome ligates its amino acid cargo to the new polypeptide chain, which is synthesized from amino terminus to carboxyl terminus. During and after its synthesis, the new protein must fold to its active three-dimensional structure before it can carry out its cellular function.

DNA replication and inheritance

The growth, development, and reproduction of organisms relies on cell division, or the process by which a single cell divides into two usually identical daughter cells. This requires first making a duplicate copy of every gene in the genome in a process called DNA replication. The copies are made by specialized enzymes known as DNA polymerases, which "read" one strand of the double-helical DNA, known as the template strand, and synthesize a new complementary strand. Because the DNA double helix is held together by base pairing, the sequence of one strand completely specifies the sequence of its complement; hence only one strand needs to be read by the enzyme to produce a faithful copy. The process of DNA replication is semiconservative; that is, the copy of the genome inherited by each daughter cell contains one original and one newly synthesized strand of DNA.

After DNA replication is complete, the cell must physically separate the two copies of the genome and divide into two distinct membrane-bound cells. In prokaryotes - bacteria and archaea - this usually occurs via a relatively simple process called binary fission, in which each circular genome attaches to the cell membrane and is separated into the daughter cells as the membrane invaginates to split the cytoplasm into two membrane-bound portions. Binary fission is extremely fast compared to the rates of cell division in eukaryotes. Eukaryotic cell division is a more complex process known as the cell cycle; DNA replication occurs during a phase of this cycle known as S phase, whereas the process of segregating chromosomes and splitting the cytoplasm occurs during M phase. In many single-celled eukaryotes such as yeast, reproduction by budding is common, which results in asymmetrical portions of cytoplasm in the two daughter cells.

Molecular inheritance

The duplication and transmission of genetic material from one generation of cells to the next is the basis for molecular inheritance, and the link between the classical and molecular pictures of genes. Organisms inherit the characteristics of their parents because the cells of the offspring contain copies of the genes in their parents' cells. In asexually reproducing organisms, the offspring will be a genetic copy or clone of the parent organism. In sexually reproducing organisms, a specialized form of cell division called meiosis produces cells called gametes or germ cells that are haploid, or contain only one copy of each gene. The gametes produced by females are called eggs or ova, and those produced by males are called sperm. Two gametes fuse to form a fertilized egg, a single cell that once again has a diploid number of genes—each with one copy from the mother and one copy from the father.

During the process of meiotic cell division, an event called genetic recombination or ''crossing-over'' can sometimes occur, in which a length of DNA on one chromatid is swapped with a length of DNA on the corresponding sister chromatid. This has no effect if the alleles on the chromatids are the same, but results in reassortment of otherwise linked alleles if they are different. The Mendelian principle of independent assortment asserts that each of a parent's two genes for each trait will sort independently into gametes; which allele an organism inherits for one trait is unrelated to which allele it inherits for another trait. This is in fact only true for genes that do not reside on the same chromosome, or are located very far from one another on the same chromosome. The closer two genes lie on the same chromosome, the more closely they will be associated in gametes and the more often they will appear together; genes that are very close are essentially never separated because it is extremely unlikely that a crossover point will occur between them. This is known as genetic linkage.

History

The notion of a gene is evolving with the science of genetics, which began when Gregor Mendel noticed that biological variations are inherited from parent organisms as specific, discrete traits. The biological entity responsible for defining traits was later termed a ''gene'', but the biological basis for inheritance remained unknown until DNA was identified as the genetic material in the 1940s. Prior to Mendel's work, the dominant theory of heredity was one of blending inheritance, which proposes that the traits of the parents blend or mix in a smooth, continuous gradient in the offspring. Although Mendel's work was largely unrecognized after its first publication in 1866, it was rediscovered in 1900 by three European scientists, Hugo de Vries, Carl Correns, and Erich von Tschermak, who had reached similar conclusions from their own research. However, these scientists were not yet aware of the identity of the 'discrete units' on which genetic material resides.

The existence of genes was first suggested by Gregor Mendel (1822–1884), who, in the 1860s, studied inheritance in peaplants (''Pisum sativum'') and hypothesized a factor that conveys traits from parent to offspring. He spent over 10 years of his life on one experiment. Although he did not use the term ''gene'', he explained his results in terms of inherited characteristics. Mendel was also the first to hypothesize independent assortment, the distinction between dominant and recessive traits, the distinction between a heterozygote and homozygote, and the difference between what would later be described as genotype (the genetic material of an organism) and phenotype (the visible traits of that organism).

Charles Darwin used the term Gemmule to describe a microscopic unit of inheritance, and what would later become known as Chromosomes had been observed separating out during cell division by Wilhelm Hofmeister as early as 1848. The idea that chromosomes are the carriers of inheritance was expressed in 1883 by Wilhelm Roux. Darwin also coined the word ''pangenesis'' by (1868). The word pangenesis is made from the Greek words ''pan'' (a prefix meaning "whole", "encompassing") and ''genesis'' ("birth") or ''genos'' ("origin").

Mendel's concept was given a name by Hugo de Vries in 1889, in his book ''Intracellular Pangenesis''; although probably unaware of Mendel's work at the time, he coined the term "pangen" for "the smallest particle [representing] one hereditary characteristic". Danish botanist Wilhelm Johannsen coined the word "gene" ("gen" in Danish and German) in 1909 to describe the fundamental physical and functional units of heredity, while the related word genetics was first used by William Bateson in 1905. He derived the word from de Vries' "pangen". In the early 1900s, Mendel's work received renewed attention from scientists. In 1910, Thomas Hunt Morgan showed that genes reside on specific chromosomes. He later showed that genes occupy specific locations on the chromosome. With this knowledge, Morgan and his students began the first chromosomal map of the fruit fly ''Drosophila''. In 1928, Frederick Griffith showed that genes could be transferred. In what is now known as Griffith's experiment, injections into a mouse of a deadly strain of bacteria that had been heat-killed transferred genetic information to a safe strain of the same bacteria, killing the mouse.

A series of subsequent discoveries led to the realization decades later that chromosomes within cells are the carriers of genetic material, and that they are made of DNA (deoxyribonucleic acid), a polymeric molecule found in all cells on which the 'discrete units' of Mendelian inheritance are encoded. In 1941, George Wells Beadle and Edward Lawrie Tatum showed that mutations in genes caused errors in specific steps in metabolic pathways. This showed that specific genes code for specific proteins, leading to the "one gene, one enzyme" hypothesis. Oswald Avery, Colin Munro MacLeod, and Maclyn McCarty showed in 1944 that DNA holds the gene's information. In 1953, James D. Watson and Francis Crick demonstrated the molecular structure of DNA. Together, these discoveries established the central dogma of molecular biology, which states that proteins are translated from RNA which is transcribed from DNA. This dogma has since been shown to have exceptions, such as reverse transcription in retroviruses.

In 1972, Walter Fiers and his team at the Laboratory of Molecular Biology of the University of Ghent (Ghent, Belgium) were the first to determine the sequence of a gene: the gene for Bacteriophage MS2 coat protein. Richard J. Roberts and Phillip Sharp discovered in 1977 that genes can be split into segments. This led to the idea that one gene can make several proteins. Recently (as of 2003–2006), biological results let the notion of gene appear more slippery. In particular, genes do not seem to sit side by side on DNA like discrete beads. Instead, regions of the DNA producing distinct proteins may overlap, so that the idea emerges that "genes are one long continuum". It was first hypothesized in 1986 by Walter Gilbert that neither DNA nor protein would be required in such a primitive system as that of a very early stage of the earth if RNA could perform as simply a catalyst and genetic information storage processor.

The modern study of genetics at the level of DNA is known as molecular genetics and the synthesis of molecular genetics with traditional Darwinian evolution is known as the modern evolutionary synthesis.

Mendelian inheritance and classical genetics

According to the theory of Mendelian inheritance, variations in phenotype—the observable physical and behavioral characteristics of an organism—are due to variations in genotype, or the organism's particular set of genes, each of which specifies a particular trait. Different forms of a gene, which may give rise to different phenotypes, are known as alleles. Organisms such as the pea plants Mendel worked on, along with many plants and animals, have two alleles for each trait, one inherited from each parent. Alleles may be dominant or recessive; dominant alleles give rise to their corresponding phenotypes when paired with any other allele for the same trait, whereas recessive alleles give rise to their corresponding phenotype only when paired with another copy of the same allele. For example, if the allele specifying tall stems in pea plants is dominant over the allele specifying short stems, then pea plants that inherit one tall allele from one parent and one short allele from the other parent will also have tall stems. Mendel's work found that alleles assort independently in the production of gametes, or germ cells, ensuring variation in the next generation.

Mutation

DNA replication is for the most part extremely accurate, with an error rate per site of around 10⁻⁶ to 10⁻¹⁰ in eukaryotes. Rare, spontaneous alterations in the base sequence of a particular gene arise from a number of sources, such as errors in DNA replication and the aftermath of DNA damage. These errors are called mutations. The cell contains many DNA repair mechanisms for preventing mutations and maintaining the integrity of the genome; however, in some cases—such as breaks in both DNA strands of a chromosome—repairing the physical damage to the molecule is a higher priority than producing an exact copy. Due to the degeneracy of the genetic code, some mutations in protein-coding genes are ''silent'', or produce no change in the amino acid sequence of the protein for which they code; for example, the codons UCU and UUC both code for serine, so the U↔C mutation has no effect on the protein. Mutations that do have phenotypic effects are most often neutral or deleterious to the organism, but sometimes they confer benefits to the organism's fitness.

Mutations propagated to the next generation lead to variations within a species' population. Variants of a single gene are known as alleles, and differences in alleles may give rise to differences in traits. Although it is rare for the variants in a single gene to have clearly distinguishable phenotypic effects, certain well-defined traits are in fact controlled by single genetic loci. A gene's most common allele is called the wild type allele, and rare alleles are called mutants. However, this does not imply that the wild-type allele is the ancestor from which the mutants are descended.

Genome

Chromosomal organization

The total complement of genes in an organism or cell is known as its genome. In prokaryotes, the vast majority of genes are located on a single chromosome of circular DNA, while eukaryotes usually possess multiple individual linear DNA helices packed into dense DNA-protein complexes called chromosomes. Genes that appear together on one chromosome of one species may appear on separate chromosomes in another species. Many species carry more than one copy of their genome within each of their somatic cells. Cells or organisms with only one copy of each chromosome are called haploid; those with two copies are called diploid; and those with more than two copies are called polyploid. The copies of genes on the chromosomes are not necessarily identical. In sexually reproducing organisms, one copy is normally inherited from each parent.

Number of genes

Early estimates of the number of human genes that used expressed sequence tag data put it at 50 000–100 000. Following the sequencing of the human genome and other genomes, it has been found that rather few genes (~20 000 in human, mouse and fly, ~13 000 in roundworm, >46 000 in rice) encode all the proteins in an organism. These protein-coding sequences make up 1–2% of the human genome. A large part of the genome is transcribed however, to introns, retrotransposons and seemingly a large array of noncoding RNAs. Total number of proteins (the Earth's proteome) is estimated to be 5 million sequences.

Genetic and genomic nomenclature

Gene nomenclature has been established by the HUGO Gene Nomenclature Committee (HGNC) for each known human gene in the form of an approved gene name and symbol (short-form abbreviation). All approved symbols are stored in the HGNC Database. Each symbol is unique and each gene is only given one approved gene symbol. This also facilitates electronic data retrieval from publications. In preference each symbol maintains parallel construction in different members of a gene family and can be used in other species, especially the mouse.

Evolutionary concept of a gene

George C. Williams first explicitly advocated the gene-centric view of evolution in his 1966 book ''Adaptation and Natural Selection''. He proposed an evolutionary concept of gene to be used when we are talking about natural selection favoring some genes. The definition is: "that which segregates and recombines with appreciable frequency." According to this definition, even an asexual genome could be considered a gene, insofar that it have an appreciable permanency through many generations.

The difference is: the molecular gene ''transcribes'' as a unit, and the evolutionary gene ''inherits'' as a unit.

Richard Dawkins' books ''The Selfish Gene'' (1976) and ''The Extended Phenotype'' (1982) defended the idea that the gene is the only replicator in living systems. This means that only genes transmit their structure largely intact and are potentially immortal in the form of copies. So, genes should be the unit of selection. In ''The Selfish Gene'' Dawkins attempts to redefine the word 'gene' to mean "an inheritable unit" instead of the generally accepted definition of "a section of DNA coding for a particular protein". In ''River Out of Eden'', Dawkins further refined the idea of gene-centric selection by describing life as a river of compatible genes flowing through geological time. Scoop up a bucket of genes from the river of genes, and we have an organism serving as temporary bodies or survival machines. A river of genes may fork into two branches representing two non-interbreeding species as a result of geographical separation.

Gene targeting and implications

Gene targeting is commonly referred to techniques for altering or disrupting mouse genes and provides the mouse models for studying the roles of individual genes in embryonic development, human disorders, aging and diseases. The mouse models, where one or more of its genes are deactivated or made inoperable, are called knockout mice. Since the first reports in which homologous recombination in embryonic stem cells was used to generate gene-targeted mice, gene targeting has proven to be a powerful means of precisely manipulating the mammalian genome, producing at least ten thousand mutant mouse strains and it is now possible to introduce mutations that can be activated at specific time points, or in specific cells or organs, both during development and in the adult animal.

Gene targeting strategies have been expanded to all kinds of modifications, including point mutations, isoform deletions, mutant allele correction, large pieces of chromosomal DNA insertion and deletion, tissue specific disruption combined with spatial and temporal regulation and so on. It is predicted that the ability to generate mouse models with predictable phenotypes will have a major impact on studies of all phases of development, immunology, neurobiology, oncology, physiology, metabolism, and human diseases. Gene targeting is also in theory applicable to species from which totipotent embryonic stem cells can be established, and therefore may offer a potential to the improvement of domestic animals and plants.

Changing concept

The concept of the gene has changed considerably (see history section). From the original definition of a "unit of inheritance", the term evolved to mean a DNA-based unit that can exert its effects on the organism through RNA or protein products. It was also previously believed that one gene makes one protein; this concept was overthrown by the discovery of alternative splicing and trans-splicing.

The definition of a gene is still changing. The first cases of RNA-based inheritance have been discovered in mammals. Evidence is also accumulating that the control regions of a gene do not necessarily have to be close to the coding sequence on the linear molecule or even on the same chromosome. Spilianakis and colleagues discovered that the promoter region of the interferon-gamma gene on chromosome 10 and the regulatory regions of the T(H)2 cytokine locus on chromosome 11 come into close proximity in the nucleus possibly to be jointly regulated.

The concept that genes are clearly delimited is also being eroded. There is evidence for fused proteins stemming from two adjacent genes that can produce two separate protein products. While it is not clear whether these fusion proteins are functional, the phenomenon is more frequent than previously thought. Even more ground-breaking than the discovery of fused genes is the observation that some proteins can be composed of exons from far away regions and even different chromosomes. This new data has led to an updated, and probably tentative, definition of a gene as "a union of genomic sequences encoding a coherent set of potentially overlapping functional products." This new definition categorizes genes by functional products, whether they be proteins or RNA, rather than specific DNA loci; all regulatory elements of DNA are therefore classified as ''gene-associated'' regions.

See also

Big gene

Copy number variation

DNA

Epigenetics

Full genome sequencing

Gene-centric view of evolution

Gene dosage

Gene expression

Gene family

Gene patent

Gene pool

Gene redundancy

Gene therapy

Genetic algorithm

Genetic Engineering

Genetics

Genome

Genomics

List of gene prediction software

List of notable genes

Meme

Predictive medicine

Pseudogene

References

Further reading

Google Book Search; first published 1976.

External links

Comparative Toxicogenomics Database

DNA From The Beginning - a primer on genes and DNA

Genes And DNA - Introduction to genes and DNA aimed at non-biologist

Entrez Gene - a searchable database of genes

IDconverter - converts gene IDs between public databases

iHOP - Information Hyperlinked over Proteins

TranscriptomeBrowser - Gene expression profile analysis

The Protein Naming Utility, a database to identify and correct deficient gene names

''Genes'' - an Open Access journal

Category:Cloning Category:Genetics Category:Molecular biology Category:Units of morphological analysis

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Gene Gene the Dancing Machine, aka Eugene Patton (born April 25, 1932), was a member of the stage crew and occasional performer on ''The Gong Show''. Gene was the first African-American member of the International Alliance of Theatrical and Stage Employees, Local 33.

He was one of several amateur performers who would "warm up" and entertain the audience during commercial breaks. Host Chuck Barris found him so entertaining that he had him dance on the show on-air, and he proved so popular that he soon became a recurring act, an occasional judge, and eventually the regular closing act for the show, with the credits regularly rolling over his enthusiastic dancing, often joined by Chuck, the celebrity judges, the stage hands, and whoever else felt like joining in. Usually Gong paddles, flowers, plastic fake food, stuffed animals and various other items were thrown on stage for good measure. He appeared in ''The Gong Show Movie'' (1980) and had some dialog. His attire usually consisted of a green windbreaker jacket, bell-bottomed jeans, sneakers, a yellow polo shirt, and a black painter's hat.

The genial heavy-set man wore a green warm up outfit and often came onstage dancing to Milton Delugg's "Band with a Thug" while they played Count Basie's "Jumpin' at the Woodside" tune.

Gene had a cameo as himself in the film ''Confessions of a Dangerous Mind''.

After ''The Gong Show'' was cancelled, Patton remained a stagehand at NBC. He was recognized backstage at ''The Tonight Show'' in the 1990s.

Patton lost both legs to complications from diabetes in 2001, and wears prosthetics and walks with a cane.

References

Category:American amputees Category:Living people Category:1932 births

Coordinates	12°58′0″N77°34′0″N
name	Gene Simmons
birth name	Chaim Weitz (; ‏)
alias	"The Demon"
born	August 25, 1949Tirat HaCarmel, Israel
background	solo_singer
occupation	Musician, songwriter, producer, actor, businessman
genre	Heavy metal, hard rock, glam metal
instrument	Bass, vocals, guitar, piano
associated acts	Kiss, Wicked Lester
years active	1963–present
website
notable instruments	Cort Signature GS-1Gene Simmons Signature model }}

Gene Simmons (born Chaim Weitz ; August 25, 1949) is an Israeli-American rock bassist, singer, songwriter, musician, actor, and businessman. Known as "The Demon", he is the bassist/vocalist of Kiss, a hard rock band he co-founded in the early 1970s.

Simmons was ranked number 12 in Roadrunner's list of 50 Best Frontmen in Metal History.

Early life

Chaim Weitz (later Gene Simmons) was born at the Rambam Hospital in Haifa, Israel in 1949. Mother and children emigrated to Jackson Heights, Queens in New York City when he was eight years old. His mother Flóra Klein was born in Jánd, Hungary. The German name Klein (means: small) translates to Hungarian as ''Kiss'', that gave the rock band's name. Florence and her brother, Larry Klein, were the only members of the family to survive the Holocaust. Simmons' father, Feri Witz, also Hungarian-born, remained in Israel. Simmons says the family was "dirt poor," scraping by on bread and milk. In the United States, Simmons changed his name to Eugene Klein (later Gene Klein), adopting his mother's maiden name. He attended Yeshiva Torah Vodaath in Williamsburg, Brooklyn as a child, from 7 a.m. to 9:30 p.m.

Music career

Simmons became involved with his first band, Lynx, then renamed The Missing Links, when he was a teenager. Eventually, he disbanded The Missing Links to form The Long Island Sounds, the name being a play on words relating to the estuary separating Long Island from Westchester County, New York, Connecticut and Rhode Island. While he played in these bands, he kept up odd jobs on the side to make more money, including trading used comic books. Simmons attended Sullivan County Community College in Loch Sheldrake, New York. He then joined a new band, Bullfrog Bheer, and the band recorded a demo, "Leeta"; this was later included on the Kiss box set.

thumb|left|120px|The Demon Simmons formed the rock band Wicked Lester in the early 1970s with Stanley Harvey Eisen (now known as Paul Stanley) and recorded one album, which was never released. Dissatisfied with Wicked Lester's sound and look, Simmons and Stanley attempted to fire their band members; they were met with resistance, and they quit Wicked Lester, walking away from their record deal with Epic Records. They decided to form the ultimate rock band, and started looking for a drummer. Simmons and Stanley found an ad placed by Peter Criscoula, known as Peter Criss, who was playing clubs in Brooklyn at the time; they joined and started out as a trio. Paul Frehley, better known as Ace Frehley, responded to an ad they put in ''The Village Voice'' for a lead guitar player, and soon joined them. Kiss released its self-titled debut album in February 1974. Stanley quickly took on the role of lead performer on stage, while Simmons became the driving force behind what became an extensive Kiss merchandising franchise. The eye section of his "Demon" makeup with KISS came from the wing design of comic book character Black Bolt. In 1983, while Kiss' fame was waning, the members took off their trademark make-up and enjoyed a resurgence in popularity that continued into the 1990s. The band hosted their own fan conventions throughout 1995, and fan feedback about the original Kiss members reunion influenced the highly successful 1996–1997 Alive Worldwide reunion tour. In 1998, the band released ''Psycho Circus'',. Since then, the original line-up has once again dissolved, with Tommy Thayer replacing Ace Frehley on lead guitar and Eric Singer (who performed with Kiss from 1992 up through 1996) replacing Peter Criss on drums.

Instruments

Gene Simmons has used various bass guitars during his career including:

Axe Bass (Kramer, jackson, b.c. rich and now Cort)

Punisher Bass (B.C.Rich, Now made by Cort)

Spector Gene Simmons Signature

Pedulla Basses (mainly in the 1980s and early 1990s)

Gibson Grabber (mainly in Kiss' early years)

Gibson Ripper (mainly in Kiss' early years)

Custom lebau bass (wicked lester, very early kiss circa 73-74)

Cort Guitars Signature GS-1 Bass

He uses Ampeg SVT CL Series amps with 8x10 cabinets. He uses Silver Herco Flex .75 Nylon Flat Guitar Picks. In 2010 Gene Simmons collaborated with Cort Guitars to make his own signature bass line called the GS-AXE-2.

Film and television career

Simmons has been the creative force behind such television projects as:

''My Dad the Rock Star'', a cartoon by the Canadian animation company Nelvana, about the mild mannered son of a Gene Simmons-like rock star;

''Mr. Romance'', a show created and hosted by Simmons on the Oxygen cable television channel;

''Rock School'', a reality show in which Simmons tries to make a rock band out of a group of students of Christ's Hospital School in the first season, and in the second, a group of kids from a comprehensive school in Lowestoft;

''Gene Simmons Family Jewels'', a reality show documenting the personal lives of Simmons, his girlfriend, his son and daughter

On March 9, 2011, Simmons and Kiss co-founder Paul Stanley and E! Entertainment announced that they have finalized a production and development deal to create an as-yet-untitled comedic half-hour kids' television series.

Simmons appeared as a psychic working at the Mystic Journey Bookstore in Venice, California on the American hidden camera prank TV series ''I Get That a Lot''.

Political views

While a self-described social liberal, Simmons was a supporter of the foreign policy of the George W. Bush administration. He supported the 2003 invasion of Iraq, writing on his website: "I'm ashamed to be surrounded by people calling themselves liberal who are, in my opinion, spitting on the graves of brave American soldiers who gave their life to fight a war that wasn't theirs...in a country they've never been to... simply to liberate the people therein". In a follow-up, Simmons explained his position and wrote about his love and support for the United States: "I wasn't born here. But I have a love for this country and its people that knows no bounds. I will forever be grateful to America for going into World War II, when it had nothing to gain, in a country that was far away... and rescued my mother from the Nazi German concentration camps. She is alive and I am alive because of America. And, if you have a problem with America, you have a problem with me".

During the 2006 Lebanon War between Israel and Lebanon, Simmons sent a televised message of support (in both English and Hebrew) to an Israeli soldier seriously wounded in fighting in Lebanon, calling him his "hero".

In 2010, Simmons said he regretted voting for Barack Obama and criticized the 2009 health care reforms. Following Obama's 2011 Mideast speech, in which the President called on Israel and the Palestinians to negotiate a settlement "based on the 1967 lines with mutually agreed swaps," Simmons told ''CNBC'' that Obama was gravely misguided. "If you have never been to the moon, you can't issue policy about the moon. For the president to be sitting in Washington D.C. and saying, 'Go back to your '67 borders in Israel' – how about you live there and try to defend an indefensible border – nine miles wide?" Simmons also accused the United Nations of being "the most pathetic body on the face of the earth."

During his visit to Israel in 2011, he stated that the artists refusing to perform in Israel for political reasons are "stupid," referring to artists who canceled planned concerts in Israel.

Personal life

He became a science fiction fan and published several science fiction fanzines, among them ''Sirruish'', ''Id'', ''Cosmos'' (which eventually merged with ''Stilletto'' to become ''Cosmos-Stilletto'' and then ''Faun''), ''Tinderbox'', ''Sci-Fi Showcase'', ''Mantis'' and ''Adventure''. He also contributed to other fanzines, among them ''BeABohema''.

In the late 1960s, he changed his name to Gene Simmons, after legendary rockabilly performer Jumpin' Gene Simmons. Simmons' legal name after arriving from Israel is Gene Klein.

Simmons lives in Beverly Hills, California along with his fiancée, former Playboy Playmate and actress Shannon Tweed. They have two children: a son, Nick (born January 22, 1989), and a daughter, Sophie, (born July 7, 1992). He formerly had live-in relationships with Cher and Diana Ross. Simmons can speak English, German, Hungarian and Hebrew.e

Homecoming visit to Israel

In March 2011, Simmons had a homecoming visit to Israel where he was born. He described the trip as a "life changing experience". He talked about how he still feels that he is an Israeli – "I'm Israeli. I'm a stranger in America. I'm an outsider". While there, Gene met his half brother Kobi, and half sisters Drora, Sharon and Ogenia, the latter two being twins. Simmons has plans to bring his band, KISS, to Israel. He has always said that he is an ardent supporter of Israel. At a press conference in Israel, he spoke both Hebrew and English.

Controversy

In February 2002, Simmons was interviewed on the NPR radio show ''Fresh Air'' and asked about his claim of having had sex with 4,600 women. He told Terry Gross: "If you want to welcome me with open arms, I'm afraid you're also going to have to welcome me with open legs," paraphrasing a lyric from The Who's 1981 hit song "You Better You Bet". Gross replied: "That's a really obnoxious thing to say." At the time, Simmons refused to grant permission to NPR to make the interview available online. However, it appears in print in Gross's book ''All I Did Was Ask'' (ISBN 1-4013-0010-3) and unauthorized transcripts are available. NPR re-broadcast part of the interview in August 2007. Simmons' bandmate Paul Stanley has frequently used the phrase "... welcomed us with open arms and open legs" in onstage patter during Kiss concerts. In 2004, during an interview in Melbourne, Australia, while talking about Islamic extremists, Simmons described Islam as a "vile culture", saying that Muslim women had to walk behind their husbands, weren't allowed to be educated or to own houses. He said: "They want to come and live right where you live and they think that you're evil." Amongst ensuing criticism, Australian Muslim of the Year Susan Carland argued that Simmons' stereotyping of Muslims was inaccurate. Simmons later clarified his comments on his website, saying he had been talking specifically about Muslim extremists. In 2005, Simmons was sued by a former girlfriend, Georgeann Walsh Ward, who said she had been "defamed" in the VH1 documentary ''When Kiss Ruled the World'' and portrayed as an "unchaste woman". A settlement was reached in June 2006. In 2011, Kiss was removed from the lineup for Michael Jackson's tribute concert because of Simmons' allegations and negative comments regarding Jackson's personal life and legal issues.

Awards and recognition

On January 28 in 2011, Simmons was presented the key to the city in Dallas, Texas and a street, Gene Simmons Boulevard, was named for him. On June 15 of the same year he was given the key to the city by Winnipeg, Manitoba.

Filmography

! Year	! Film	! Role	! Bandmates	! Miscellaneous
1978	''Kiss Meets the Phantom of the Park''	The Demon	Peter Criss, Ace Frehley, Paul Stanley	TV movie
1984		Dr. Charles Luther
1986		Nuke (radio DJ)
1986	''Never Too Young to Die''	Carruthers / Velvet Von Ragner
1987		Malak Al Rahim
1988	''The Decline of Western Civilization Part II: The Metal Years''	Himself	Paul Stanley	Documentary
1989	''Red Surf''	Doc
1999		Himself	Peter Criss, Ace Frehley, Paul Stanley	Simmons also produced
2002	''The New Guy''	Reverend
2002	''Wish You Were Dead''	Vinny
2008	''Detroit Metal City''	Jack lll Dark		Japanese movie
2009		Joe Adler
2010	''Expecting Mary''	Taylor
2010	''Rush: Beyond the Lighted Stage	Himself		Documentary

Television appearances

! Year	! Show	! Episode	! Role	! Bandmates	! Miscellaneous
1974	''The Mike Douglas Show''	June 11, 1974	Himself	Peter Criss, Ace Frehley, Paul Stanley	Kiss' first national television appearance, performed "Firehouse"
1976	''The Paul Lynde Halloween Special''	October 29, 1976	Himself	Peter Criss, Ace Frehley, Paul Stanley	The band performed "Detroit Rock City (song)
1985	''[[Miami Vice''	"Prodigal Son"	Newton Blade
1997	''Action League Now!''	"Rock-A-Big-Baby"	Toy version of himself	Peter Criss, Ace Frehley, Paul Stanley	Performed "Rock and Roll All Nite"
1998	''MADtv''	October 31, 1998 (#406)	Himself	Peter Criss, Ace Frehley, Paul Stanley.	Halloween special; performed in five sketches
2001	''Family Guy''	"A Very Special Family Guy Freakin' Christmas"	Animated version of himself	Peter Criss, Ace Frehley, Paul Stanley
2001		May 27, 2001	Himself		Won $500,000 for a charity
2001	''The Daily Show with Jon Stewart''	June 12, 2001	Himself		Almost licked Jon Stewart's face in response to Jon being unable to explain the difference between pleasure and joy to a man with a 'twelve-inch tongue'
2002	''Family Guy''	"Road to Europe"	Animated version of himself	Peter Criss, Ace Frehley, Paul Stanley
2002		September 6, 2002	Himself
2003	''King of the Hill''	"Reborn to Be Wild"	Jessie
2004	''Third Watch''	"Higher Calling"	Donald Mann
2004	''Third Watch''	"Monsters"	Donald Mann
2004	''Third Watch''	"More Monsters"	Donald Mann
2005	''American Idol''	"Auditions: New Orleans"	Himself- guest judge
2005	''Family Guy''		Animated version of himself/ Prisoner #3
2005	''Mind of Mencia''	Episode #1.7	Himself
2006		August 1, 2006	Himself		Gene Simmons Family Jewels
2007	''SpongeBob SquarePants''	"20,000 Patties Under the Sea"	Sea Monster
2007		"Gene Simmons"	Himself		UK's More4 show
2008	''Entertainment Tonight''	January 2, 2008	Himself
2008	''Jimmy Kimmel Live!''	January 18, 2008	Himself
2008		March 11, 2008	Himself
2008	''Criss Angel Mindfreak''	"Mindfreaking with the Stars"	Himself
2008	''Ugly Betty''	"The Kids Are Alright"	Himself
2008	''Ugly Betty''	"A Thousand Words by Friday"	Himself
2008		Episode #3.9	Himself		Won $500,000 for the Elizabeth Glaser Pediatric AIDS Foundation charity
2008		First three episodes	Himself		Fired in the third episode
2008	''Jingles''		Celebrity judge		Mark Burnett reality show
2009	''Glenn Martin DDS''	Glenn Gary, Glenn Martin	himself
2009	''American Idol''	Season Finale	Himself	Eric Singer, Tommy Thayer, Paul Stanley	Performed medley of 3 songs with contestant Adam Lambert
2009	''The Fairly OddParents''		Animated version of himself	Eric Singer, Tommy Thayer, Paul Stanley
2009	''The Fairly OddParents''		Animated version of himself	Eric Singer, Tommy Thayer, Paul Stanley
2010	''I Get That a Lot''	Episode 2	As Himself		Simmons appeared as a psychic working at the Mystic Journey Bookstore in Venice, California
2010	''Extreme Makeover: Home Edition''	Episode 160	As Himself		The band Kiss made a personal appearance during the Wagstaff family's vacation in Disneyland and honored them as special guests at one of their concerts. Kiss also made a personal appearance at a local school where a donation of new musical instruments was made in the Wagstaff family's name.
2010	''I'm in a Rock 'n' Roll Band!''	Episode 1 And 5	As Himself		Discussing the requirements of being in a rock band.
2011		To Love and Die in LA	As Himself

Video appearances

In 2007, he appeared alongside other celebrities, as well as regular people, in the music video for "Rockstar" by Nickelback.

Video game appearances

Gene Simmons is a playable character in ''Tony Hawk's Underground'', unlocked when completing the story mode on Normal difficulty, and also appears with his Kiss bandmates in the Hotter Than Hell level to play one of three songs upon collecting the four K-I-S-S letters.

Gene Simmons' Kiss character, The Demon, is a playable character in ''Kiss: Psycho Circus: The Nightmare Child''.

Simmons also has a large role in the 2010 music video game ''Guitar Hero: Warriors of Rock''. In addition to narrating the main storyline and doing advertising for the game, the Kiss song "Love Gun" is playable.

Discography

Kiss

Solo

''Gene Simmons'' (1978)

''Sex Money Kiss'' (audiobook CD), 2003)

''Asshole'' (2004)

''Speaking in Tongues'' (spoken word CD, 2004)

Publishing career

In 2002, Simmons launched ''Gene Simmons' Tongue'', a men's lifestyle magazine. The magazine lasted five issues before being discontinued.

Published works

''Kiss and Make-Up'', Gene Simmons (ISBN 0-609-81002-2)

''Sex Money Kiss'', Gene Simmons (ISBN 1-893224-86-4)

''Kiss: The Early Years'', Gene Simmons and Paul Stanley (ISBN 0-609-81028-6)

''Kiss: Behind the Mask'', David Leaf and Ken Sharp (ISBN 0-446-96524-6)

''Ladies of the Night: A Historical and Personal Perspective on the Oldest Profession in the World'', Gene Simmons (ISBN 1-59777-501-0)

References

External links

Category:1949 births Category:American entertainment industry businesspeople Category:American baritones Category:American bloggers Category:American heavy metal bass guitarists Category:American heavy metal singers Category:American people of Israeli descent Category:American male singers Category:American record producers Category:American rock singers Category:American voice actors Category:Children of Holocaust survivors Category:Israeli bloggers Category:Israeli heavy metal bass guitarists Category:Israeli heavy metal singers Category:Israeli emigrants to the United States Category:Israeli Jews Category:Israeli male singers Category:Israeli record producers Category:Israeli voice actors Category:Jewish American musicians Category:Jewish singers Category:Kiss (band) members Category:Living people Category:Naturalized citizens of the United States Category:Participants in American reality television series Category:People from Haifa Category:People from Staten Island Category:Science fiction fans Category:Singers from New York City Category:The Apprentice (U.S. TV series) contestants Category:American musicians of Hungarian descent

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Coordinates	12°58′0″N77°34′0″N
name	Gene Kelly
birth name	Eugene Curran Kelly
birth date	August 23, 1912
birth place	Pittsburgh, Pennsylvania, United States
death date	February 02, 1996
death place	Beverly Hills, California, United States
occupation	Actor, dancer, singer, director, producer, choreographer
years active	1938–94
spouse	}}

Eugene Curran "Gene" Kelly (August 23, 1912 February 2, 1996) was an American dancer, actor, singer, film director and producer, and choreographer. Kelly was known for his energetic and athletic dancing style, his good looks and the likeable characters that he played on screen.

Although he is known today for his performances in ''Singin' in the Rain'' and ''An American in Paris'', he was a dominant force in Hollywood musical films from the mid 1940s until this art form fell out of fashion in the late 1950s. His many innovations transformed the Hollywood musical film, and he is credited with almost single-handedly making the ballet form commercially acceptable to film audiences.

Kelly was the recipient of an Academy Honorary Award in 1952 for his career achievements. He later received lifetime achievement awards in the Kennedy Center Honors, and from the Screen Actors Guild and American Film Institute; in 1999, the American Film Institute also numbered him 15th in their Greatest Male Stars of All Time list.

Early life

He was the third son of James Kelly, a phonograph salesman, and Harriet Curran, who were both children of Irish Roman Catholic immigrants. He was born in the Highland Park neighborhood of Pittsburgh, Pennsylvania and, at the age of eight, was enrolled by his mother in dance classes, along with his elder brother James. They both rebelled, and, according to Kelly: "We didn't like it much and were continually involved in fistfights with the neighborhood boys who called us sissies...I didn't dance again until I was fifteen." He thought it would be a good way to get girls. Kelly returned to dance on his own initiative and by then was an accomplished sportsman and well able to take care of himself. He attended St. Raphael Elementary School in the Morningside neighborhood of Pittsburgh, PA. He graduated from Peabody High School in 1929 at the age of sixteen. He enrolled in Pennsylvania State College to study journalism but the economic crash obliged him to seek employment to help with the family's finances. At this time, he worked up dance routines with his younger brother Fred in order to earn prize money in local talent contests, and they also performed in local nightclubs.

In 1931, Kelly enrolled at the University of Pittsburgh to study economics where he joined the Phi Kappa Theta fraternity. While at Pitt, Kelly became involved in the university's Cap and Gown Club, which staged original, comedic musical productions. Earning a Bachelor of Arts in Economics with his graduation from Pitt in 1933, he remained active with the Cap and Gown Club, serving as its director from 1934 to 1938, while at the same time enrolling in the University of Pittsburgh Law School. Also during this period, the Kelly's family started a dance studio on Munhall Road in the Squirrel Hill neighborhood of Pittsburgh in 1930. In 1932, the dance studio was renamed The Gene Kelly Studio of the Dance. A second location was opened in Johnstown, Pennsylvania in 1933. Kelly served as a teacher at the dance studio during both his undergraduate and law student years at Pitt. In 1931, he was approached by the Rodef Shalom synagogue in Pittsburgh to teach dance and stage the annual Kermess and was so successful that his services were retained for seven years until his departure for New York. Eventually, though, he decided to pursue his career as a dance teacher and entertainer full-time and so dropped out of law school after two months. He began to focus increasingly on performing, later claiming: "With time I became disenchanted with teaching because the ratio of girls to boys was more than ten to one, and once the girls reached sixteen the dropout rate was very high." In 1937, having successfully managed and developed the family's dance school business, he moved to New York City in search of work as a choreographer.

Stage career

After a fruitless search, Kelly returned to Pittsburgh, to his first position as a choreographer with the Charles Gaynor musical revue ''Hold Your Hats'' at the Pittsburgh Playhouse in April, 1938. Kelly appeared in six of the sketches, one of which, "La Cumparsita", became the basis of an extended Spanish number in ''Anchors Aweigh'' eight years later.

His first Broadway assignment, in November 1938, was as a dancer in Cole Porter's ''Leave It to Me!'' as the American ambassador's secretary who supports Mary Martin while she sings "My Heart Belongs to Daddy". He had been hired by Robert Alton who had staged a show at the Pittsburgh Playhouse and been impressed by Kelly's teaching skills. When Alton moved on to choreograph ''One for the Money'' he hired Kelly to act, sing and dance in a total of eight routines. In 1939, he was selected to be part of a musical revue "One for the Money" produced by the actress Katharine Cornell, who was known for finding and hiring talented young actors.

Kelly's first career breakthrough was in the Pulitzer Prize-winning ''The Time of Your Life'', which opened on October 25, 1939, where for the first time on Broadway he danced to his own choreography. In the same year he received his first assignment as a Broadway choreographer, for ''Billy Rose's Diamond Horseshoe''. His future wife, Betsy Blair, was a member of the cast. They began dating and married on October 16, 1941.

In 1940, he was given the leading role in Rodgers and Hart's ''Pal Joey'', again choreographed by Robert Alton, and this role propelled him to stardom. During its run he told reporters: "I don't believe in conformity to any school of dancing. I create what the drama and the music demand. While I am a hundred percent for ballet technique, I use only what I can adapt to my own use. I never let technique get in the way of mood or continuity." It was at this time also, that his phenomenal commitment to rehearsal and hard work was noticed by his colleagues. Van Johnson who also appeared in ''Pal Joey'' recalled: "I watched him rehearsing, and it seemed to me that there was no possible room for improvement. Yet he wasn't satisfied. It was midnight and we had been rehearsing since eight in the morning. I was making my way sleepily down the long flight of stairs when I heard staccato steps coming from the stage...I could see just a single lamp burning. Under it, a figure was dancing...Gene."

Offers from Hollywood began to arrive but Kelly was in no particular hurry to leave New York. Eventually, he signed with David O. Selznick, agreeing to go to Hollywood at the end of his commitment to ''Pal Joey'', in October 1941. Prior to his contract, he also managed to fit in choreographing the stage production of ''Best Foot Forward''.

Film career

1941–1944: Becoming established in Hollywood

Selznick sold half of Kelly's contract to MGM and loaned him out to MGM for his first motion picture: ''For Me and My Gal'' (1942) with Judy Garland. Kelly was "appalled at the sight of myself blown up twenty times. I had an awful feeling that I was a tremendous flop" but the picture did well and, in the face of much internal resistance, Arthur Freed of MGM picked up the other half of Kelly's contract. After appearing in the B-movie drama ''Pilot #5'' he took the male lead in Cole Porter's ''Du Barry Was a Lady'' opposite Lucille Ball. His first opportunity to dance to his own choreography came in his next picture ''Thousands Cheer'', where he performed a mock-love dance with a mop.

He achieved his breakthrough as a dancer on film, when MGM loaned him out to Columbia to work with Rita Hayworth in ''Cover Girl'' (1944), where he created a memorable routine dancing to his own reflection. In his next film ''Anchors Aweigh'' (1945), MGM virtually gave him a free hand to devise a range of dance routines, including the celebrated and much imitated animated dances with Jerry Mouse, and his duets with co-star Frank Sinatra. Anchors Aweigh became one of the most successful films of 1945 and it garnered Kelly his first and only Academy Award nomination for Best Actor. In ''Ziegfeld Follies'' (1946) – which was produced in 1944 but not released until 1946 – Kelly collaborated with Fred Astaire – for whom he had the greatest admiration – in the famous "The Babbitt and the Bromide" challenge dance routine before leaving the studio for wartime service. Throughout this period Kelly was obliged to appear in straight acting roles in a series of cheap B-movies, now largely forgotten.

At the end of 1944, Kelly enlisted in the U.S. Naval Air Service and was commissioned as lieutenant junior grade. He was stationed in the Photographic Section, Washington D.C., where he was involved in writing and directing a range of documentaries, and this stimulated his interest in the production side of film-making.

1946–1952: MGM

On his return to Hollywood in the spring of 1946, MGM had nothing lined up and used him in yet another B-movie: ''Living in a Big Way''. The film was considered so weak that Kelly was asked to design and insert a series of dance routines, and his ability to carry off such assignments was noticed. This led to his next picture with Judy Garland and director Vincente Minnelli, the film version of Cole Porter's ''The Pirate'', in which Kelly plays the eponymous swashbuckler. Now regarded as a classic, the film was ahead of its time and was not well received. ''The Pirate'' gave full rein to Kelly's athleticism and is probably best remembered for Kelly's work with The Nicholas Brothers – the leading African-American dancers of their day – in a virtuoso dance routine. Although MGM wanted Kelly to return to safer and more commercial vehicles, he ceaselessly fought for an opportunity to direct his own musical film. In the interim, he capitalised on his swashbuckling image as d'Artagnan in ''The Three Musketeers''. and also appeared with Vera-Ellen in the ''Slaughter on Tenth Avenue'' ballet in ''Words and Music'' (1948). He was due to play the male lead opposite Garland in ''Easter Parade'' (1948), but broke his ankle playing volleyball. He withdrew from the film and encouraged Fred Astaire to come out of retirement to replace him. There followed ''Take Me Out to the Ball Game'' (1949), his second film with Sinatra, where Kelly paid tribute to his Irish heritage in ''The Hat My Father Wore on St. Patrick's Day'' routine. It was this musical film which persuaded Arthur Freed to allow Kelly to make ''On the Town'', where he partnered with Frank Sinatra for the third and final time, creating a breakthrough in the musical film genre which has been described as "the most inventive and effervescent musical thus far produced in Hollywood."

Stanley Donen, brought to Hollywood by Kelly to be his assistant choreographer, received co-director credit for ''On the Town''. According to Kelly: "...when you are involved in doing choreography for film you must have expert assistants. I needed one to watch my performance, and one to work with the cameraman on the timing..without such people as Stanley, Carol Haney and Jeanne Coyne I could never have done these things. When we came to do ''On the Town'', I knew it was time for Stanley to get screen credit because we weren't boss-assistant anymore but co-creators." Together, they opened up the musical form, taking the film musical out of the studio and into real locations, with Donen taking responsibility for the staging and Kelly handling the choreography. Kelly went much further than before in introducing modern ballet into his dance sequences, going so far in the "Day in New York" routine as to substitute four leading ballet specialists for Sinatra, Munshin, Garrett and Miller.

It was now Kelly's turn to ask the studio for a straight acting role and he took the lead role in the early mafia melodrama: ''The Black Hand'' (1949). This expose of organized crime is set in New York's "Little Italy" the late 19th century, and focuses on the Black Hand, a group which extorts money upon threat of death. In the real-life incidents upon which this film is based, it was the Mafia, not the Black Hand, who functioned as the villain. Even in 1950, however, Hollywood had to tread gingerly whenever dealing with big-time crime; it was easier (and safer) to go after a "dead" criminal organization than a "live" one.

There followed ''Summer Stock'' (1950) – Judy Garland's last musical film for MGM – in which Kelly performed the celebrated "You, You Wonderful You" solo routine with a newspaper and a squeaky floorboard. In his book "Easy the Hard Way", Joe Pasternak, head of one of the other musical units within MGM, singled out Kelly for his patience and willingness to spend as much time as necessary to enable the ailing Garland to complete her part.

There followed in quick succession two musicals which have secured Kelly's reputation as a major figure in the American musical film, ''An American in Paris'' (1951) and – probably the most popular and admired of all film musicals – ''Singin' in the Rain'' (1952). As co-director, lead star and choreographer, Kelly was the central driving force. Johnny Green, head of music at MGM at the time, described him as follows:

"Gene is easygoing as long as you know exactly what you are doing when you're working with him. He's a hard taskmaster and he loves hard work. If you want to play on his team you'd better like hard work, too. He isn't cruel but he is tough, and if Gene believed in something he didn't care who he was talking to, whether it was Louis B. Mayer or the gatekeeper. He wasn't awed by anybody, and he had a good record of getting what he wanted".

''An American in Paris'' won six Academy Awards, including Best Picture and, in the same year, Kelly was presented with an honorary Academy Award for his contribution to film musicals and the art of choreography. The film also marked the debut of Leslie Caron, whom Kelly had spotted in Paris and brought to Hollywood. Its dream ballet sequence, lasting an unprecedented seventeen minutes, was the most expensive production number ever filmed up to that point and was described by Bosley Crowther as, "''whoop-de-doo'' ... one of the finest ever put on the screen." ''Singin' in the Rain'' featured Kelly's celebrated and much imitated solo dance routine to the title song, along with the "Moses Supposes" routine with Donald O'Connor and the "Broadway Melody" finale with Cyd Charisse, and while it did not initially generate the same enthusiasm as ''An American in Paris'', it subsequently overtook the earlier film to occupy its current pre-eminent place among critics and filmgoers alike.

1953–57: The Decline of the Hollywood Musical

Kelly, at the very peak of his creative powers, now made what in retrospect is seen as a serious mistake. In December 1951 he signed a contract with MGM which sent him to Europe for nineteen months so that Kelly could use MGM funds frozen in Europe to make three pictures while personally benefiting from tax exemptions. Only one of these pictures was a musical, ''Invitation to the Dance'', a pet project of Kelly's to bring modern ballet to mainstream film audiences. It was beset with delays and technical problems, and flopped when finally released in 1956. When Kelly returned to Hollywood in 1953, the film musical was already beginning to feel the pressures from television, and MGM cut the budget for his next picture ''Brigadoon'' (1954), with Cyd Charisse, forcing the film to be made on studio backlots instead of on location in Scotland. This year also saw him appear as guest star with his brother Fred in the celebrated "I Love to Go Swimmin' with Wimmen" routine in ''Deep in My Heart''. MGM's refusal to loan him out for ''Guys and Dolls'' and ''Pal Joey'' put further strains on his relationship with the studio. He negotiated an exit to his contract which involved making three further pictures for MGM.

The first of these, ''It's Always Fair Weather'' (1956) co-directed with Donen, was a musical satire on television and advertising, and includes his famous roller skate dance routine to "I Like Myself", and a dance trio with Michael Kidd and Dan Dailey which allowed Kelly to experiment with the widescreen possibilities of Cinemascope. A modest success, it was followed by Kelly's last musical film for MGM, ''Les Girls'' (1957), in which he partnered a trio of leading ladies, Mitzi Gaynor, Kay Kendall and Taina Elg, fittingly ending, as he had begun, with a Cole Porter musical. The third picture he completed was a co-production between MGM and himself, the B-movie ''The Happy Road'', set in his beloved France, his first foray in his new role as producer-director-actor.

1958–1996: Years of perseverance

Kelly did not return to stage work until his MGM contract ended in 1957, when in 1958 he directed Rodgers and Hammerstein's musical play ''Flower Drum Song''. Early in 1960 Kelly, an ardent Francophile and fluent French speaker, was invited by A. M. Julien, the general administrator of the Paris Opéra and Opéra-Comique, to select his own material and create a modern ballet for the company, the first time an American received such an assignment. The result was ''Pas de Dieux'', based on Greek mythology combined with the music of George Gershwin's ''Concerto in F''. It was a major success, and led to his being honored with the Chevalier of the Legion d'Honneur by the French Government.

Kelly continued to make some film appearances, such as Hornbeck in the 1960 Hollywood production of ''Inherit the Wind''. However, most of his efforts were now concentrated on film production and directing. He directed Jackie Gleason in ''Gigot'' in Paris, but the film was subsequently drastically recut by Seven Arts Productions and flopped. Another French effort, Jacques Demy's homage to the MGM musical: ''Les Demoiselles de Rochefort'' (1967) in which Kelly appeared, also performed poorly. He appeared as himself in George Cukor's ''Let's Make Love'' (1960).

His first foray into television was a documentary for NBC's ''Omnibus'', ''Dancing is a Man's Game'' (1958) where he assembled a group of America's greatest sportsmen – including Mickey Mantle, Sugar Ray Robinson and Bob Cousy – and reinterpreted their moves choreographically, as part of his lifelong quest to remove the effeminate stereotype of the art of dance, while articulating the philosophy behind his dance style. It gained an Emmy nomination for choreography and now stands as the key document explaining Kelly's approach to modern dance.

Kelly also frequently appeared on television shows during the 1960s, but his one effort at television series, as Father Chuck O'Malley in ''Going My Way'' (1962–63), based on the Best Picture of 1944 starring Bing Crosby, was dropped after thirty episodes, although it enjoyed great popularity in Roman Catholic countries outside of the United States. He also appeared in three major TV specials: ''New York, New York'' (1966), ''The Julie Andrews Show'' (1965), and ''Jack and the Beanstalk'' (1967) a show he produced and directed which returned to a combination of cartoon animation with live dance, winning him an Emmy Award for Outstanding Children's Program.

In 1963, Kelly joined Universal Pictures for a two-year stint which proved to be the most unproductive of his career so far. He joined 20th Century Fox in 1965, but had little to do – partly due to his decision to decline assignments away from Los Angeles for family reasons. His perseverance finally paid off with the major box-office hit ''A Guide for the Married Man'' (1967) where he directed Walter Matthau and a major opportunity arose when Fox – buoyed by the returns from ''The Sound of Music'' (1965) – commissioned Kelly to direct ''Hello, Dolly!'' (1969), again directing Matthau along with Barbra Streisand, but which unfortunately failed to recoup the enormous production expenses.

In 1970, he made another TV special: ''Gene Kelly and 50 Girls'' and was invited to bring the show to Las Vegas, which he duly did for an eight-week stint – on condition he be paid more than any artist had hitherto been paid there. He directed veteran actors James Stewart and Henry Fonda in the comedy western ''The Cheyenne Social Club'' (1970) which performed very well at the box-office. In 1973 he would work again with Frank Sinatra as part of Sinatra's Emmy nominated TV special ''Ol' Blue Eyes Is Back''. Then, in 1974, he appeared as one of many special narrators in the surprise hit of the year ''That's Entertainment!'' and subsequently directed and co-starred with his friend Fred Astaire in the sequel ''That's Entertainment, Part II'' (1976). It was a measure of his powers of persuasion that he managed to coax the 77-year-old Astaire – who had insisted that his contract rule out any dancing, having long since retired – into performing a series of song and dance duets, evoking a powerful nostalgia for the glory days of the American musical film. Kelly continued to make frequent TV appearances and in 1980, appeared in an acting and dancing role opposite Olivia Newton-John in ''Xanadu'' (1980), an expensive theatrical flop which has since attained a cult following. In Kelly's opinion "The concept was marvelous but it just didn't come off." In the same year, he was invited by Francis Ford Coppola to recruit a production staff for American Zoetrope's ''One from the Heart'' (1982). Although Coppola's ambition was for him to establish a production unit to rival the Freed Unit at MGM, the film's failure put an end to this idea. In 1985, Kelly served as executive producer and co-host of ''That's Dancing!'' – a celebration of the history of dance in the American musical. After his final on-screen appearance introducing ''That's Entertainment! III'' in 1994, his final film project was the animated movie ''Cats Don't Dance'', released in 1997 and dedicated to him, on which Kelly acted as uncredited choreographic consultant.

Working methods and influence on filmed dance

When he began his collaborative film work, he was influenced by Robert Alton and John Murray Anderson, striving to create moods and character insight with his dances. He choreographed his own movement, along with that of the ensemble, with the assistance of Jeanne Coyne, Stanley Donen, Carol Haney, and Alex Romero. He experimented with lighting, camera techniques and special effects in order to achieve true integration of dance with film, and was one of the first to use split screens, double images, live action with animation and is credited as the person who made the ballet form commercially acceptable to film audiences.

There was a clear progression in his development, from an early concentration on tap and musical comedy style to greater complexity using ballet and modern dance forms. Kelly himself refused to categorize his style: "I don't have a name for my style of dancing...It's certainly hybrid...I've borrowed from the modern dance, from the classical, and certainly from the American folk dance – tap-dancing, jitterbugging...But I have tried to develop a style which is indigenous to the environment in which I was reared." He especially acknowledged the influence of George M. Cohan: "I have a lot of Cohan in me. It's an Irish quality, a jaw-jutting, up-on-the-toes cockiness – which is a good quality for a male dancer to have." He was also heavily influenced by an African-American dancer Dancing Dotson, whom he saw at Loew's Penn. Theatre around 1929, and was briefly taught by Frank Harrington, an African-American tap specialist from New York. However, his main interest was in ballet, which he studied under Kotchetovsky in the early Thirties. As biographer Clive Hirschhorn explains: "As a child he used to run for miles through parks and streets and woods – anywhere, just as long as he could feel the wind against his body and through his hair. Ballet gave him the same feeling of exhilaration, and in 1933 he was convinced it was the most satisfying form of self-expression." He also studied Spanish dancing under Angel Cansino, Rita Hayworth's uncle. Generally speaking, he tended to use tap and other popular dance idioms to express joy and exuberance – as in the title song from ''Singin' in the Rain'' or "I Got Rhythm" from ''An American in Paris'', whereas pensive or romantic feelings were more often expressed via ballet or modern dance, as in "Heather on the Hill" from ''Brigadoon'' or "Our Love Is Here to Stay" from ''An American in Paris''.

According to Delamater, Kelly's work "seems to represent the fulfillment of dance-film integration in the 1940s and 1950s". While Fred Astaire had revolutionized the filming of dance in the 1930s by insisting on full-figure photography of dancers while allowing only a modest degree of camera movement, Kelly freed up the camera, making greater use of space, camera movement, camera angles and editing, creating a partnership between dance movement and camera movement without sacrificing full-figure framing. Kelly's reasoning behind this was that he felt the kinetic force of live dance often evaporated when brought to film, and he sought to partially overcome this by involving the camera in movement and giving the dancer a greater number of directions in which to move. Examples of this abound in Kelly's work and are well illustrated in the "Prehistoric Man" sequence from ''On the Town'' and "The Hat My Father Wore on St. Patrick's Day" from ''Take Me Out to the Ball Game''. In 1951, he summed up his vision as follows: "If the camera is to make a contribution at all to dance, this must be the focal point of its contribution; the fluid background, giving each spectator an undistorted and altogether similar view of dancer and background. To accomplish this, the camera is made fluid, moving with the dancer, so that the lens becomes the eye of the spectator, ''your eye''".

Kelly's athleticism gave his moves a distinctive broad, muscular quality, and this was a very deliberate choice on his part, as he explained: "There's a strong link between sports and dancing, and my own dancing springs from my early days as an athlete...I think dancing is a man's game and if he does it well he does it better than a woman." He railed against what he saw as the widespread effeminacy in male dancing which, in his opinion, "tragically" stigmatized the genre, alienating boys from entering the field: "Dancing ''does'' attract effeminate young men. I don't object to that as long as they don't dance effeminately. I just say that if a man dances effeminately he dances badly — just as if a woman comes out on stage and starts to sing bass. Unfortunately people confuse gracefulness with softness. John Wayne is a graceful man and so are some of the great ball players...but, of course, they don't run the risk of being called sissies." In his view, "one of our problems is that so much dancing is taught by women. You can spot many male dancers who have this tuition by their arm movements — they are soft, limp and feminine." He acknowledged that, in spite of his efforts — in TV programs such as ''Dancing: A Man's Game'' (1958) for example — the situation changed little over the years.

He also sought to break from the class-conscious conventions of the 1930s and early 40s, when top hat and tails or tuxedos were the norm, by dancing in casual or everyday work clothes, so as to make his dancing more relevant to the cinema-going public. As his first wife, actress and dancer Betsy Blair explained: "A sailor suit or his white socks and loafers, or the T-shirts on his muscular torso, gave everyone the feeling that he was a regular guy, and perhaps they too could express love and joy by dancing in the street or stomping through puddles...he democratized the dance in movies." In particular, he wanted to create a completely different image from that associated with Fred Astaire, not least because he believed his physique didn't suit such refined elegance: "I used to envy his cool aristocratic style, so intimate and contained. Fred wears top hat and tails to the manor born — I put them on and look like a truck driver."

Personal life and death

Kelly was married to Betsy Blair for 15 years (1941–1957) and they had one child, Kerry. Kelly divorced Blair in 1957. In 1960, Kelly married his choreographic assistant Jeanne Coyne, who had divorced Stanley Donen in 1949 after a brief marriage. He remained married to Coyne from 1960 until her death in 1973 and they had two children, Bridget and Tim. He was married to Patricia Ward from 1990 until his death in 1996.

Kelly was a lifelong supporter of the Democratic Party with strong progressive convictions, which occasionally created difficulty for him as his period of greatest prominence coincided with the McCarthy era in the U.S. In 1947, he was part of the Committee for the First Amendment, the Hollywood delegation which flew to Washington to protest at the first official hearings by the House Committee on Un-American Activities. His first wife, Betsy Blair, was suspected of being a Communist sympathizer and when MGM, who had offered Blair a part in ''Marty'' (1955), were considering withdrawing her under pressure from the American Legion, Kelly successfully threatened MGM with a pullout from ''It's Always Fair Weather'' unless his wife was restored to the part. He used his position on the board of directors of the Writers Guild of America, West on a number of occasions to mediate disputes between unions and the Hollywood studios, and although he was frequently accused by some on the right of championing the unions, he was valued by the studios as an effective mediator.

Kelly was raised as a Roman Catholic, but in September 1939, Kelly, having already become disgusted by the Roman Catholic Church's support for Francisco Franco against the Spanish Republic, officially severed his ties with the church after a trip to Mexico convinced him that the church had failed to look out for the poor. For the rest of his life he was an agnostic.

Kelly was a major financial supporter of the Irish Republican Army (IRA), and he left thousands of pounds to NORAID in his will.

He retained a lifelong passion for sports and relished competition. He was known as a big fan of the Pittsburgh Steelers and New York Yankees. From the mid-40s through the early 50s, he and Blair organized weekly parties at their Beverly Hills home which were renowned for an intensely competitive and physical version of charades, known as "The Game".

Kelly died in his sleep on February 2, 1996, in Beverly Hills, California after a stroke – he had also suffered a stroke the year before. His body was cremated the same day and he had left instructions that there was to be no funeral and no memorial services. Kelly's papers are currently housed at the Howard Gotlieb Archival Research Center at Boston University.

Awards and honors

1946 – Academy Award nomination for Best Actor in ''Anchors Aweigh'' (1945).

1952 – Honorary Academy Award "in appreciation of his versatility as an actor, singer, director and dancer, and specifically for his brilliant achievements in the art of choreography on film." This Oscar was lost in a fire in 1983 and replaced at the 1984 Academy Awards.

1956 – Golden Bear at the 6th Berlin International Film Festival for ''Invitation to the Dance''.

1958 – Nomination for Golden Laurel Award for Best Male Musical Performance in ''Les Girls''.

1958 – ''Dance Magazine'''s annual TV Award for ''Dancing: A Man's Game'' from the ''Omnibus'' television series. It was also nominated for an Emmy for best choreography.

1960 – In France, Kelly was made a Chevalier of the Legion of Honor.

1962 – Gene Kelly Dance Film Festival staged by the Museum of Modern Art.

1964 – Silver Sail Best Actor for ''What a Way to Go!'' (1964) at the Locarno International Film Festival.

1967 – Emmy for Outstanding Children's Program for ''Jack and the Beanstalk''.

1970 – Nomination for Golden Globe, Best Director for ''Hello, Dolly!'', 1969.

1981 – Cecil B. DeMille Award at Golden Globes.

1981 – Gene Kelly was the subject of a two-week film festival in France.

1982 – Lifetime Achievement Award in the fifth annual Kennedy Center Honors.

1985 – Lifetime Achievement Award from the American Film Institute.

1989 – Life Achievement Award from Screen Actors Guild.

1991 – University of Pittsburgh inaugurates The Gene Kelly Awards, given annually to high school musicals in Allegheny County, Pennsylvania.

1992 – Induction into the Theater Hall of Fame.

1994 – National Medal of Arts awarded by United States President Bill Clinton.

1994 – The Three Tenors performed ''Singin' in the Rain'' in his presence during a concert at Dodger Stadium in Los Angeles.

1996 – Honorary César Award The César is the main national film award in France.

1996 – At the Academy Awards ceremony, director Quincy Jones organised a tribute to the just-deceased Kelly, in which Savion Glover performed the dance to "Singin' in the Rain".

1997 – Ranked #26 in Empire (UK) magazine's "The Top 100 Movie Stars of All Time" list.

1999 – Ranked #15 in the American Film Institute's "Greatest Legends" list.

Work

Filmography

Musical films

Gene Kelly appeared as actor and dancer in the following musical films. He always choreographed his own dance routines, and often the dance routines of others, and often used assistants. As was the practice at the time, he was rarely formally credited in the film titles:

{|class="wikitable" style="font-size: 90%;" border="2" cellpadding="4" background: #f9f9f9; |- align="center" ! style="background:#B0C4DE;" | Year ! style="background:#B0C4DE;" | Film ! style="background:#B0C4DE;" | Role ! style="background:#B0C4DE;" | Notes |- |1942 |''For Me and My Gal'' |Harry Palmer | |- |rowspan=2|1943 |''Du Barry Was a Lady'' |Alec Howe/Black Arrow | |- |''Thousands Cheer'' |Private Eddie Marsh | |- |1944 |''Cover Girl'' |Danny McGuire | |- |rowspan=2|1945 |''Anchors Aweigh'' |Joseph Brady |Nominated — Academy Award for Best Actor |- |''Ziegfeld Follies'' |Gentleman in 'The Babbit and the Bromide' | |- |1947 |''Living in a Big Way'' |Leo Gogarty | |- |rowspan=2|1948 |''The Pirate'' |Serafin | |- |''Words and Music'' |Himself | |- |rowspan=2|1949 |''Take Me Out to the Ball Game'' |Eddie O'Brien | |- |''On the Town'' |Gabey | |- |1950 |''Summer Stock'' |Joe D. Ross | |- |1951 |''An American in Paris'' |Jerry Mulligan |Nominated — Golden Globe Award for Best Actor – Motion Picture Musical or Comedy |- |1952 |''Singin' in the Rain'' |Don Lockwood | |- |rowspan=2|1954 |''Brigadoon'' |Tommy Albright | |- |''Deep in My Heart'' |Specialty in 'Dancing Around' | |- |1955 |''It's Always Fair Weather'' |Ted Riley | |- |1956 |''Invitation to the Dance'' |Host/Pierrot/The Marine/Sinbad | |- |1957 |''Les Girls'' |Barry Nichols | |- |1958 |''Marjorie Morningstar'' |Noel Airman | |- |1960 |''Inherit the Wind'' |E.K. Hornbeck | |- |1960 |''Let's Make Love'' |Himself | |- |1964 |''What a Way to Go!'' |Pinky Benson | |- |1966 |''Les Demoiselles de Rochefort'' |Andy Miller | |- |1974 |''That's Entertainment!'' |Himself |(also archive footage) |- |1976 |''That's Entertainment, Part II'' |Himself |(also archive footage) |- |1980 |''Xanadu'' |Danny McGuire |- |1994 |''That's Entertainment, Part III'' |Himself |(also archive footage) |- |}

Stage

{|class="wikitable" style="font-size: 90%;" border="2" cellpadding="4" background: #f9f9f9; |- align="center" ! style="background:#B0C4DE;" | Dates ! style="background:#B0C4DE;" | Title ! style="background:#B0C4DE;" | Role ! style="background:#B0C4DE;" | Notes |- |November 9, 1938 – July 15, 1939 |''Leave It to Me!'' |Secretary to Mr. Goodhue | |- |February 4, 1939 – May 27, 1939 |''One for the Money'' |various roles | |- |October 25, 1939 – April 6, 1940 |''The Time of Your Life'' |Harry | |- |September 23, 1940 – October 19, 1940 |''The Time of Your Life'' |Harry | |- |December 25, 1940 – November 29, 1941 |''Pal Joey'' |Joey Evans | |- |October 1, 1941 – July 4, 1942 |''Best Foot Forward'' | |Choreography |- |December 1, 1958 – May 7, 1960 |''Flower Drum Song'' | | Director |- |February 22, 1979 – April 1, 1979 |''Coquelico'' | |Producer |- |July 2, 1985 – May 18, 1986 |''Singin' in the Rain'' | |Original film choreographyNominated — Drama Desk Award for Outstanding Choreography |}

Television

{|class="wikitable" style="font-size: 90%;" border="2" cellpadding="4" background: #f9f9f9; |- align="center" ! style="background:#B0C4DE;" | Year ! style="background:#B0C4DE;" | Title ! style="background:#B0C4DE;" | Role ! style="background:#B0C4DE;" | Notes |- |1948 |''The Three Musketeers'' |D'Artagnan |- |1958 |''Dancing: A Man's Game'' |Himself |''Omnibus'' |- |1962–1963 |''Going My Way'' |Father Chuck O'Malley |(30 episodes) |- |rowspan=2|1965 |''Gene Kelly: New York, New York'' |Himself | |- |''The Julie Andrews Show'' |Himself | |- |1967 |''Jack and the Beanstalk'' |Jeremy Keen, Proprietor (Peddler) |Emmy Award for Best Children's Program |- |1971 |''The Funny Side'' |Himself |Series host |- |1973 |''Frank Sinatra: Ol' Blue Eyes is Back'' |Himself | |- |1978 |''Gene Kelly: An American in Pasadena'' |Himself | |- |1980 |''Muppet Show'' |Himself | |- |1985 |''North and South'' |Senator Charles Edwards | |- |1986 |''Sins'' |Eric Hovland | |}

Notes

External links

The Gene Kelly Awards – University of Pittsburgh

Obituary, NY Times, February 3, 1996

Naval Intelligence File on Gene Kelly

Gene Kelly – An American Life – PBS

Category:1912 births Category:1996 deaths Category:Academy Honorary Award recipients Category:Actors from Pittsburgh, Pennsylvania Category:American choreographers Category:American dancers Category:American film actors Category:20th-century actors Category:Traditional pop music singers Category:American film directors Category:American male singers Category:American tap dancers Category:Burials at Westwood Village Memorial Park Cemetery Category:César Award winners Category:Chevaliers of the Légion d'honneur Category:Deaths from stroke Category:Emmy Award winners Category:American musicians of Irish descent Category:American people of Irish descent Category:Kennedy Center honorees Category:United States National Medal of Arts recipients Category:University of Pittsburgh alumni Category:Film choreographers Category:American agnostics Category:Former Roman Catholics Category:Pennsylvania Democrats Category:California Democrats

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Coordinates	12°58′0″N77°34′0″N
name	Gene Krupa
background	non_vocal_instrumentalist
birth name	Eugene Bertram Krupa
born	January 15, 1909Chicago, Illinois, U.S.
died	October 16, 1973Yonkers, New York, U.S.
instrument	Drums
genre	JazzSwingDixielandBig band
occupation	DrummerComposerBandleader
years active	1920s – mid 1960s, with intermittent performances in the 1970s
associated acts	Eddie CondonBenny GoodmanLouie BellsonAnita O'Day }}

Gene Krupa (January 15, 1909 – October 16, 1973) was an influential American jazz and big band drummer and composer, known for his highly energetic and flamboyant style.

Biography

Eugene Bertram Krupa was born in Chicago, Illinois, the youngest of Anna (Oslowski) and Bartlomiej Krupa's nine children. Bartlomiej was an immigrant from Poland, and Anna was born in Shamokin, Northumberland County, Pennsylvania. His siblings were Clarence, Eleanor, Casimir, Leo, Peter and Julius.

Krupa studied with Sanford A. Moeller and began playing professionally in the mid 1920s with bands in Wisconsin. He broke into the Chicago scene in 1927, when he was picked by MCA to become a member of "Thelma Terry and Her Playboys," the first notable American Jazz band (outside of all-girl bands) to be led by a female musician. The Playboys were the house band at ''The Golden Pumpkin'' nightclub in Chicago and also toured extensively throughout the eastern and central United States.

Krupa made his first recordings in 1927, with a band under the leadership of banjoist Eddie Condon and Red McKenzie: along with other recordings beginning in 1924 by musicians known in the "Chicago" scene such as Bix Beiderbecke, these sides are examples of "Chicago Style" jazz. The numbers recorded at that session were: "China Boy", "Sugar", "Nobody's Sweetheart" and "Liza". The McKenzie - Condon sides are also notable for being some of the early examples of the use of a full drum kit on recordings. Eddie Condon describes what happened in the Okeh Records studio on that day (in 'We Called It Music' - pub: Peter Davis, 1948):

Krupa's big influences during this time were Tubby Hall and Zutty Singleton. The drummer who probably had the greatest influence on Gene in this period was the great Baby Dodds. Dodds' use of press rolls was highly reflected in Gene's playing.

Quietly we waited for the playback. When it came, pounding out through the big speaker, we listened stiffly for a moment. We had never been an audience for ourselves...Rockwell came out of the control-room smiling. 'We'll have to get some more of this... (Rockwell nodded towards Krupa): didn't bother the equipment at all,' he said. 'I think we've got something,'.

Krupa also appeared on six recordings made by the Thelma Terry band in 1928 In 1929, he was part of the Mound City Blue Blowers sessions, that also included Red McKenzie, Glenn Miller, and Coleman Hawkins, which produced "Hello Lola" and "One Hour", which Krupa was credited with co-writing.

In 1929 he moved to New York City and worked with the band of Red Nichols. In 1933, Krupa first played with Benny Goodman. He became part of the Benny Goodman trio, the first popular integrated musical group in the United States. In 1934 he joined Benny Goodman's band, where his featured drum work made him a national celebrity. His tom-tom interludes on their hit "Sing, Sing, Sing" were the first extended drum solos to be recorded commercially. In 1938, Krupa performed with the Goodman Orchestra in the famous Carnegie Hall Jazz Concert.

After a public fight with Goodman at the Earl Theater in Philadelphia, Krupa left Goodman to launch his own band and had several hits with singer Anita O'Day and trumpeter Roy Eldridge.

In 1939, Gene Krupa and his Orchestra appeared in the Paramount movie ''Some Like It Hot,'' which starred [Bob Hope & Shirley Ross], performing the title song, "Blue Rhythm Fantasy," and "The Lady's in Love with You." Krupa made a memorable cameo appearance in a pivotal scene of the 1941 film ''Ball of Fire,'' in which he and his band performed an extended version of the hit "Drum Boogie," which he composed with trumpeter Roy Eldridge.

thumb|300px|Gene Krupa Drive in Yonkers, NYIn 1943, Krupa was arrested for possession of two marijuana cigarettes and was given a three-month jail sentence. Krupa was not a wealthy man and spent most of his savings defending himself of this charge and fell into a depression for several months, believing his career to be over. Then, Goodman invited him to perform with his orchestra. Audiences welcomed Krupa's performances, and while the reunion would never last, Krupa was performing again, thanks to this nudge.

Krupa soon formed his second orchestra. This one was notable for its large string section, and also featured Charlie Ventura on sax. It was one of the largest dance bands of the era, sometimes containing up to forty musicians. He also invited another drummer into the band so that he could take breaks and lead the orchestra from the front. However, audiences were not paying to see him conduct, and he gradually accepted this.

The end of the swing era

As the 1940s closed, large orchestras fell by the wayside: Count Basie closed his large band and Woody Herman reduced his to an octet. Krupa also gradually cut down the size of the band in the late 1940s, and from 1951 on led a trio or quartet, often featuring the multi-instrumentalist Eddie Shu on tenor sax, clarinet and harmonica. He appeared regularly with the ''Jazz At the Philharmonic'' shows.

Krupa also had a fleeting brush with Hollywood. Along with his musical sequence in the 1941 film comedy hit Ball of Fire, he also delivered a cameo appearance in the 1946 screen classic ''The Best Years Of Our Lives''. His athletic drumming style, timing methods and cymbal technique evolved during this decade to fit in with changed fashions and tastes, but he never quite adjusted to the Be-Bop period. In 1954, Krupa returned to Hollywood, performing, along with Louis Armstrong, "Basin Street Blues" in Jimmy Stewart's bio-pic ''The Glenn Miller Story''. He also joined fellow Benny Goodman alumni Harry James, Teddy Wilson, and Lionel Hampton in ''The Benny Goodman Story'', starring Steve Allen. In 1959, the movie biography ''The Gene Krupa Story'' was released, with Sal Mineo portraying Krupa and a cameo appearance by Red Nichols. Rainn Wilson writes for Turner Classic Movies about this film, which was most unusual for a biopic in its era. Wilson says, "More fact than fiction, The Gene Krupa Story avoids sugarcoating Krupa's life and takes a warts-and-all approach which gives the film an emotional honesty that other screen biographies often lack. In fact, Mineo's portrayal of Krupa is so needy, egocentric, manic and ruthlessly ambitious that you may find yourself rooting for his comeuppance which he receives in spades, starting with a drug bust for marijuana. Dave Frishberg, a pianist who played with Krupa, was particularly struck by the accuracy of one key moment in the film. "The scene where the Krupa character drops his sticks during the big solo, and the audience realizes that he's "back on the stuff." I remember at least a couple of occasions in real life when Gene dropped a stick, and people in the audience began whispering among themselves and pointing at Gene."

Krupa continued to perform even in famous clubs in the 1960s like the Metropole, near Times Square in New York City, often playing duets with African American drummer Cozy Cole. Increasingly troubled by back pain, he retired in the late 1960s and opened a music school. One of his pupils was Kiss drummer Peter Criss. He occasionally played in public in the early 1970s until shortly before his death. Krupa married Ethel Maguire—twice, in fact; the first marriage lasted from 1934–1942; the second one dates from 1946 to her death in 1955. Their relationship was dramatized in the biopic about him. Krupa remarried in 1959 (to Patty Bowler). He died of leukemia and heart failure in Yonkers, New York at the age of sixty-four. He was buried in Holy Cross Cemetery in Calumet City, Illinois.

Legacy

Many consider Krupa to be one of the most influential drummers of the 20th century, particularly regarding the development of the drum kit. Many jazz historians believe he made history in 1927 as the first kit drummer ever to record using a bass drum pedal. Others, however, believe this was done earlier by Baby Dodds. His drum method was published in 1938 and immediately became the standard text. He is also credited with inventing the rim shot on the snare drum.

Krupa in the 1930s prominently featured Slingerland drums. At Krupa's urging, Slingerland developed tom-toms with tuneable top and bottom heads, which immediately became important elements of virtually every drummer's set-up. Krupa also developed and popularised many of the cymbal techniques that became standards. His collaboration with Armand Zildjian of the Avedis Zildjian Company developed the modern hi-hat cymbals and standardized the names and uses of the ride cymbal, the crash cymbal, the splash cymbal, the pang cymbal and the swish cymbal.

Krupa was featured in the 1946 Warner Bros. cartoon ''Book Revue'' in which a rotoscoped version of Krupa's drumming is used in an impromptu jam session.

The 1937 recording of Louis Prima's "Sing, Sing, Sing (With a Swing)" by Benny Goodman and His Orchestra featuring Gene Krupa on drums was inducted into the Grammy Hall of Fame.

In 1978, Gene Krupa became the first drummer inducted into the Modern Drummer Hall of Fame.

Krupa was mentioned in the Simpsons episode "Hurricane Neddy", when Ned Flanders parents are being told they must control Ned, Ned's father responds "We can't do it man! That's discipline! That's like tellin' Gene Krupa not to go "Boom boom bah bah bah, boom boom bah bah bah, boom boom boom bah bah bah bah, boom boom tss!"".

Rhythm, the UK's best selling drum magazine voted Gene Krupa the third most influential drummer ever, in a poll conducted for its February 2009 issue. Voters included over 50 top-name drummers.

Discography

Benny Goodman: ''The Famous Carnegie Hall Concert 1938'' (Columbia)

''Drummin´ Man'' (Charly, 1938–41) with Roy Eldridge, Anita O'Day, Benny Carter, Charlie Ventura

''Drum Boogie'' (Columbia, 1940–41)

''Uptown'' (Columbia, 1941–1949)

''Lionel Hampton/Gene Krupa'' (Forlane, 1949) with Don Fagerquist, Frank Rehak, Frank Rosolino, Roy Eldridge

''The Exciting Gene Krupa'' (Enoch's Music, 1953) with Charlie Shavers, Bill Harris, Willie Smith, Ben Webster, Teddy Wilson, Herb Ellis, Ray Brown, Israel Crosby

''Krupa and Rich'' (Verve, 1955) with Roy Eldridge, Dizzy Gillespie, Illinois Jacquet, Flip Phillips, Oscar Peterson, Herb Ellis, Ray Brown, Buddy Rich

''Gene Krupa Big Band: Drummer Man featuring Anita O'Day & Roy Eldridge'' (Verve, 1956)

References

External links

Gene Krupa at traditional-jazz.com

The Gene Krupa Reference Page

Gene Krupa Drummerworld Page

''Let Me Off Uptown: A Gene Krupa Biography'' by John Twomey

Category:1909 births Category:1973 deaths Category:American jazz drummers Category:American jazz bandleaders Category:American people of Polish descent Category:American musicians of Polish descent Category:Big band bandleaders Category:Big band drummers Category:Cancer deaths in New York Category:Deaths from leukemia Category:Dixieland drummers Category:Musicians from Chicago, Illinois Category:RCA Victor artists Category:Verve Records artists Category:Columbia Records artists Category:Swing drummers

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