text:
Human Genome Project
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Logo HGP; Vitruvian Man, Leonardo da Vinci
The Human Genome Project (HGP) was an international scientific research project with the goal of determining the sequence of nucleotide base pairs that make up human DNA, and of identifying and mapping all of the genes of the human genome from both a physical and a functional standpoint.[1] It remains the world's largest collaborative biological project.[2] After the idea was picked up in 1984 by the US government when the planning started, the project formally launched in 1990 and was declared complete in 2003[3]. Funding came from the US government through the National Institutes of Health (NIH) as well as numerous other groups from around the world. A parallel project was conducted outside government by the Celera Corporation, or Celera Genomics, which was formally launched in 1998. Most of the government-sponsored sequencing was performed in twenty universities and research centers in the United States, the United Kingdom, Japan, France, Germany, Spain and China.[4]
The Human Genome Project originally aimed to map the nucleotides contained in a human haploid reference genome (more than three billion). The "genome" of any given individual is unique; mapping the "human genome" involved sequencing a small number of individuals and then assembling these together to get a complete sequence for each chromosome. Therefore, the finished human genome is a mosaic, not representing any one individual.
Contents
[hide]
1 Human Genome Project
1.1 History
1.2 State of completion
2 Applications and proposed benefits
3 Techniques and analysis
3.1 Findings
3.2 Accomplishment
4 Public versus private approaches
5 Genome donors
6 Developments
7 Ethical, legal and social issues
8 See also
9 References
10 Further reading
11 External links
Human Genome Project[edit]
History[edit]
The Human Genome Project was a 15-year-long, publicly funded project initiated in 1990 with the objective of determining the DNA sequence of the entire euchromatic human genome within 15 years.[5] In May 1985, Robert Sinsheimer organized a workshop to discuss sequencing the human genome,[6] but for a number of reasons the NIH was uninterested in pursuing the proposal. The following March, the Santa Fe Workshop was organized by Charles DeLisi and David Smith of the Department of Energy's Office of Health and Environmental Research (OHER).[7] At the same time Renato Dulbecco proposed whole genome sequencing in an essay in Science.[8] James Watson followed two months later with a workshop held at the Cold Spring Harbor Laboratory.
The fact that the Santa Fe workshop was motivated and supported by a Federal Agency opened a path, albeit a difficult and tortuous one,[9] for converting the idea into a public policy in the United States. In a memo to the Assistant Secretary for Energy Research (Alvin Trivelpiece), Charles DeLisi, who was then Director of the OHER, outlined a broad plan for the project.[10] This started a long and complex chain of events which led to approved reprogramming of funds that enabled the OHER to launch the Project in 1986, and to recommend the first line item for the HGP, which was in President Reagan's 1988 budget submission,[9] and ultimately approved by the Congress. Of particular importance in Congressional approval was the advocacy of Senator Peter Domenici, whom DeLisi had befriended.[11] Domenici chaired the Senate Committee on Energy and Natural Resources, as well as the Budget Committee, both of which were key in the DOE budget process. Congress added a comparable amount to the NIH budget, thereby beginning official funding by both agencies.
Alvin Trivelpiece sought and obtained the approval of DeLisi's proposal by Deputy Secretary William Flynn Martin. This chart[12] was used in the spring of 1986 by Trivelpiece, then Director of the Office of Energy Research in the Department of Energy, to brief Martin and Under Secretary Joseph Salgado regarding his intention to reprogram $4 million to initiate the project with the approval of Secretary Herrington. This reprogramming was followed by a line item budget of $16 million in the Reagan Administration’s 1987 budget submission to Congress.[13] It subsequently passed both Houses. The Project was planned for 15 years.[14]
Candidate technologies were already being considered for the proposed undertaking at least as early as 1985.[15]
In 1990, the two major funding agencies, DOE and NIH, developed a memorandum of understanding in order to coordinate plans and set the clock for the initiation of the Project to 1990.[16] At that time, David Galas was Director of the renamed “Office of Biological and Environmental Research” in the U.S. Department of Energy’s Office of Science and James Watson headed the NIH Genome Program. In 1993, Aristides Patrinos succeeded Galas and Francis Collins succeeded James Watson, assuming the role of overall Project Head as Director of the U.S. National Institutes of Health (NIH) National Center for Human Genome Research (which would later become the National Human Genome Research Institute). A working draft of the genome was announced in 2000 and the papers describing it were published in February 2001. A more complete draft was published in 2003, and genome "finishing" work continued for more than a decade.
The $3-billion project was formally founded in 1990 by the US Department of Energy and the National Institutes of Health, and was expected to take 15 years.[17] In addition to the United States, the international consortium comprised geneticists in the United Kingdom, France, Australia, China and myriad other spontaneous relationships.[18]
Due to widespread international cooperation and advances in the field of genomics (especially in sequence analysis), as well as major advances in computing technology, a 'rough draft' of the genome was finished in 2000 (announced jointly by U.S. President Bill Clinton and the British Prime Minister Tony Blair on June 26, 2000).[19] This first available rough draft assembly of the genome was completed by the Genome Bioinformatics Group at the University of California, Santa Cruz, primarily led by then graduate student Jim Kent. Ongoing sequencing led to the announcement of the essentially complete genome on April 14, 2003, two years earlier than planned.[20][21] In May 2006, another milestone was passed on the way to completion of the project, when the sequence of the last chromosome was published in Nature.[22]
State of completion[edit]
The project was not able to sequence all the DNA found in human cells. It sequenced only "euchromatic" regions of the genome, which make up 92% of the human genome. The other regions, called "heterochromatic" are found in centromeres and telomeres, and were not sequenced under the project.[23]
The Human Genome Project was declared complete in April 2003. An initial rough draft of the human genome was available in June 2000 and by February 2001 a working draft had been completed and published followed by the final sequencing mapping of the human genome on April 14, 2003. Although this was reported to cover 99% of the euchromatic human genome with 99.99% accuracy, a major quality assessment of the human genome sequence was published on May 27, 2004 indicating over 92% of sampling exceeded 99.99% accuracy which was within the intended goal.[24] Further analyses and papers on the HGP continue to occur.[25]
Applications and proposed benefits[edit]
The sequencing of the human genome holds benefits for many fields, from molecular medicine to human evolution. The Human Genome Project, through its sequencing of the DNA, can help us understand diseases including: genotyping of specific viruses to direct appropriate treatment; identification of mutations linked to different forms of cancer; the design of medication and more accurate prediction of their effects; advancement in forensic applied sciences; biofuels and other energy applications; agriculture, animal husbandry, bioprocessing; risk assessment; bioarcheology, anthropology and evolution. Another proposed benefit is the commercial development of genomics research related to DNA based products, a multibillion-dollar industry.
The sequence of the DNA is stored in databases available to anyone on the Internet. The U.S. National Center for Biotechnology Information (and sister organizations in Europe and Japan) house the gene sequence in a database known as GenBank, along with sequences of known and hypothetical genes and proteins. Other organizations, such as the UCSC Genome Browser at the University of California, Santa Cruz,[26] and Ensembl[27] present additional data and annotation and powerful tools for visualizing and searching it. Computer programs have been developed to analyze the data, because the data itself is difficult to interpret without such programs. Generally speaking, advances in genome sequencing technology have followed Moore’s Law, a concept from computer science which states that integrated circuits can increase in complexity at an exponential rate.[28] This means that the speeds at which whole genomes can be sequenced can increase at a similar rate, as was seen during the development of the above-mentioned Human Genome Project.
Techniques and analysis[edit]
This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (April 2017) (Learn how and when to remove this template message)
The process of identifying the boundaries between genes and other features in a raw DNA sequence is called genome annotation and is in the domain of bioinformatics. While expert biologists make the best annotators, their work proceeds slowly, and computer programs are increasingly used to meet the high-throughput demands of genome sequencing projects. Beginning in 2008, a new technology known as RNA-seq was introduced that allowed scientists to directly sequence the messenger RNA in cells. This replaced previous methods of annotation, which relied on inherent properties of the DNA sequence, with direct measurement, which was much more accurate. Today, annotation of the human genome and other genomes relies primarily on deep sequencing of the transcripts in every human tissue using RNA-seq. These experiments have revealed that over 90% of genes contain at least one and usually several alternative splice variants, in which the exons are combined in different ways to produce 2 or more gene products from the same locus.[citation needed]
The genome published by the HGP does not represent the sequence of every individual's genome. It is the combined mosaic of a small number of anonymous donors, all of European origin. The HGP genome is a scaffold for future work in identifying differences among individuals. Subsequent projects sequenced the genomes of multiple distinct ethnic groups, though as of today there is still only one "reference genome."[citation needed]
Findings[edit]
Key findings of the draft (2001) and complete (2004) genome sequences include:
There are approximately 22,300[29] protein-coding genes in human beings, the same range as in other mammals.
The human genome has significantly more segmental duplications (nearly identical, repeated sections of DNA) than had been previously suspected.[30][31][32]
At the time when the draft sequence was published fewer than 7% of protein families appeared to be vertebrate specific.[33]
Accomplishment[edit]
The first printout of the human genome to be presented as a series of books, displayed at the Wellcome Collection, London
The Human Genome Project was started in 1990 with the goal of sequencing and identifying all three billion chemical units in the human genetic instruction set, finding the genetic roots of disease and then developing treatments. It is considered a megaproject because the human genome has approximately 3.3 billion base-pairs. With the sequence in hand, the next step was to identify the genetic variants that increase the risk for common diseases like cancer and diabetes.[16][34]
It was far too expensive at that time to think of sequencing patients’ whole genomes. So the National Institutes of Health embraced the idea for a "shortcut", which was to look just at sites on the genome where many people have a variant DNA unit. The theory behind the shortcut was that, since the major diseases are common, so too would be the genetic variants that caused them. Natural selection keeps the human genome free of variants that damage health before children are grown, the theory held, but fails against variants that strike later in life, allowing them to become quite common. (In 2002 the National Institutes of Health started a $138 million project called the HapMap to catalog the common variants in European, East Asian and African genomes.)[35]
The genome was broken into smaller pieces; approximately 150,000 base pairs in length.[34] These pieces were then ligated into a type of vector known as "bacterial artificial chromosomes", or BACs, which are derived from bacterial chromosomes which have been genetically engineered. The vectors containing the genes can be inserted into bacteria where they are copied by the bacterial DNA replication machinery. Each of these pieces was then sequenced separately as a small "shotgun" project and then assembled. The larger, 150,000 base pairs go together to create chromosomes. This is known as the "hierarchical shotgun" approach, because the genome is first broken into relatively large chunks, which are then mapped to chromosomes before being selected for sequencing.[36][37]
Funding came from the US government through the National Institutes of Health in the United States, and a UK charity organization, the Wellcome Trust, as well as numerous other groups from around the world. The funding supported a number of large sequencing centers including those at Whitehead Institute, the Wellcome Sanger Institute (then called The Sanger Centre) based at the Wellcome Genome Campus, Washington University in St. Louis, and Baylor College of Medicine.[17][38]
The United Nations Educational, Scientific and Cultural Organization (UNESCO) served as an important channel for the involvement of developing countries in the Human Genome Project.[39]
Public versus private approaches[edit]
In 1998, a similar, privately funded quest was launched by the American researcher Craig Venter, and his firm Celera Genomics. Venter was a scientist at the NIH during the early 1990s when the project was initiated. The $300,000,000 Celera effort was intended to proceed at a faster pace and at a fraction of the cost of the roughly $3 billion publicly funded project. The Celera approach was able to proceed at a much more rapid rate, and at a lower cost than the public project because it relied upon data made available by the publicly funded project.[40]
Celera used a technique called whole genome shotgun sequencing, employing pairwise end sequencing,[41] which had been used to sequence bacterial genomes of up to six million base pairs in length, but not for anything nearly as large as the three billion base pair human genome.
Celera initially announced that it would seek patent protection on "only 200–300" genes, but later amended this to seeking "intellectual property protection" on "fully-characterized important structures" amounting to 100–300 targets. The firm eventually filed preliminary ("place-holder") patent applications on 6,500 whole or partial genes. Celera also promised to publish their findings in accordance with the terms of the 1996 "Bermuda Statement", by releasing new data annually (the HGP released its new data daily), although, unlike the publicly funded project, they would not permit free redistribution or scientific use of the data. The publicly funded competitors were compelled to release the first draft of the human genome before Celera for this reason. On July 7, 2000, the UCSC Genome Bioinformatics Group released a first working draft on the web. The scientific community downloaded about 500 GB of information from the UCSC genome server in the first 24 hours of free and unrestricted access.[42]
In March 2000, President Clinton announced that the genome sequence could not be patented, and should be made freely available to all researchers. The statement sent Celera's stock plummeting and dragged down the biotechnology-heavy Nasdaq. The biotechnology sector lost about $50 billion in market capitalization in two days.
Although the working draft was announced in June 2000, it was not until February 2001 that Celera and the HGP scientists published details of their drafts. Special issues of Nature (which published the publicly funded project's scientific paper)[43] and Science (which published Celera's paper[44]) described the methods used to produce the draft sequence and offered analysis of the sequence. These drafts covered about 83% of the genome (90% of the euchromatic regions with 150,000 gaps and the order and orientation of many segments not yet established). In February 2001, at the time of the joint publications, press releases announced that the project had been completed by both groups. Improved drafts were announced in 2003 and 2005, filling in to approximately 92% of the sequence currently.
Genome donors[edit]
In the IHGSC international public-sector HGP, researchers collected blood (female) or sperm (male) samples from a large number of donors. Only a few of many collected samples were processed as DNA resources. Thus the donor identities were protected so neither donors nor scientists could know whose DNA was sequenced. DNA clones from many different libraries were used in the overall project, with most of those libraries being created by Pieter J. de Jong's. Much of the sequence (>70%) of the reference genome produced by the public HGP came from a single anonymous male donor from Buffalo, New York (code name RP11).[45][46]
HGP scientists used white blood cells from the blood of two male and two female donors (randomly selected from 20 of each) – each donor yielding a separate DNA library. One of these libraries (RP11) was used considerably more than others, due to quality considerations. One minor technical issue is that male samples contain just over half as much DNA from the sex chromosomes (one X chromosome and one Y chromosome) compared to female samples (which contain two X chromosomes). The other 22 chromosomes (the autosomes) are the same for both sexes.
Although the main sequencing phase of the HGP has been completed, studies of DNA variation continue in the International HapMap Project, whose goal is to identify patterns of single-nucleotide polymorphism (SNP) groups (called haplotypes, or “haps”). The DNA samples for the HapMap came from a total of 270 individuals: Yoruba people in Ibadan, Nigeria; Japanese people in Tokyo; Han Chinese in Beijing; and the French Centre d’Etude du Polymorphisme Humain (CEPH) resource, which consisted of residents of the United States having ancestry from Western and Northern Europe.
In the Celera Genomics private-sector project, DNA from five different individuals were used for sequencing. The lead scientist of Celera Genomics at that time, Craig Venter, later acknowledged (in a public letter to the journal Science) that his DNA was one of 21 samples in the pool, five of which were selected for use.[47][48]
In 2007, a team led by Jonathan Rothberg published James Watson's entire genome, unveiling the six-billion-nucleotide genome of a single individual for the first time.[49]
Developments[edit]
The work on interpretation and analysis of genome data is still in its initial stages. It is anticipated that detailed knowledge of the human genome will provide new avenues for advances in medicine and biotechnology. Clear practical results of the project emerged even before the work was finished. For example, a number of companies, such as Myriad Genetics, started offering easy ways to administer genetic tests that can show predisposition to a variety of illnesses, including breast cancer, hemostasis disorders, cystic fibrosis, liver diseases and many others. Also, the etiologies for cancers, Alzheimer's disease and other areas of clinical interest are considered likely to benefit from genome information and possibly may lead in the long term to significant advances in their management.[35][50]
There are also many tangible benefits for biologists. For example, a researcher investigating a certain form of cancer may have narrowed down their search to a particular gene. By visiting the human genome database on the World Wide Web, this researcher can examine what other scientists have written about this gene, including (potentially) the three-dimensional structure of its product, its function(s), its evolutionary relationships to other human genes, or to genes in mice or yeast or fruit flies, possible detrimental mutations, interactions with other genes, body tissues in which this gene is activated, and diseases associated with this gene or other datatypes. Further, deeper understanding of the disease processes at the level of molecular biology may determine new therapeutic procedures. Given the established importance of DNA in molecular biology and its central role in determining the fundamental operation of cellular processes, it is likely that expanded knowledge in this area will facilitate medical advances in numerous areas of clinical interest that may not have been possible without them.[51]
The analysis of similarities between DNA sequences from different organisms is also opening new avenues in the study of evolution. In many cases, evolutionary questions can now be framed in terms of molecular biology; indeed, many major evolutionary milestones (the emergence of the ribosome and organelles, the development of embryos with body plans, the vertebrate immune system) can be related to the molecular level. Many questions about the similarities and differences between humans and our closest relatives (the primates, and indeed the other mammals) are expected to be illuminated by the data in this project.[35][52]
The project inspired and paved the way for genomic work in other fields, such as agriculture. For example, by studying the genetic composition of Tritium aestivum, the world’s most commonly used bread wheat, great insight has been gained into the ways that domestication has impacted the evolution of the plant.[53] Which loci are most susceptible to manipulation, and how does this play out in evolutionary terms? Genetic sequencing has allowed these questions to be addressed for the first time, as specific loci can be compared in wild and domesticated strains of the plant. This will allow for advances in genetic modification in the future which could yield healthier, more disease-resistant wheat crops.
Ethical, legal and social issues[edit]
At the onset of the Human Genome Project several ethical, legal, and social concerns were raised in regards to how increased knowledge of the human genome could be used to discriminate against people. One of the main concerns of most individuals was the fear that both employers and health insurance companies would refuse to hire individuals or refuse to provide insurance to people because of a health concern indicated by someone's genes.[54] In 1996 the United States passed the Health Insurance Portability and Accountability Act (HIPAA) which protects against the unauthorized and non-consensual release of individually identifiable health information to any entity not actively engaged in the provision of healthcare services to a patient.[55]
Along with identifying all of the approximately 20,000–25,000 genes in the human genome, the Human Genome Project also sought to address the ethical, legal, and social issues that were created by the onset of the project. For that the Ethical, Legal, and Social Implications (ELSI) program was founded in 1990. Five percent of the annual budget was allocated to address the ELSI arising from the project.[17][56] This budget started at approximately $1.57 million in the year 1990, but increased to approximately $18 million in the year 2014.[57]
Whilst the project may offer significant benefits to medicine and scientific research, some authors have emphasized the need to address the potential social consequences of mapping the human genome. "Molecularising disease and their possible cure will have a profound impact on what patients expect from medical help and the new generation of doctors' perception of illness."[58]
See also[edit]
1000 Genomes Project
100,000 Genomes Project
Chimpanzee Genome Project
ENCODE
EuroPhysiome
Genome Compiler
HUGO Gene Nomenclature Committee
Human Brain Project
Human Connectome Project
Human Cytome Project
Human Microbiome Project
Human proteome project
Human Variome Project
List of biological databases
Neanderthal Genome Project
Sanger Institute
The Genographic Project
References[edit]
Jump up ^ Robert Krulwich (2001-04-17). Cracking the Code of Life (Television Show). PBS.
Jump up ^ "Economic Impact of the Human Genome Project – Battelle" (PDF). Retrieved 1 August 2013.
Jump up ^ https://www.genome.gov/11006943/human-genome-project-completion-frequently-asked-questions/. Missing or empty |title= (help)
Jump up ^ "Human Genome Project Completion: Frequently Asked Questions". genome.gov.
Jump up ^ "Human Genome Project: Sequencing the Human Genome | Learn Science at Scitable". www.nature.com. Retrieved 2016-01-25.
Jump up ^ Sinsheimer, Robert (1989). "The Santa Cruz Workshop, May 1985". Genomics. 5: 954–956. doi:10.1016/0888-7543(89)90142-0.
Jump up ^ DeLisi, Charles (October 2008). "Conferences That Changed the World". Nature. 455: 876–877. Bibcode:2008Natur.455..876D. doi:10.1038/455876a. PMID 18923499.
Jump up ^ Dulbecco, Renato (1986). "Turning Point in Cancer Research, Sequencing the Human Genome". Science. 231 (4742): 1055–1056. Bibcode:1986Sci...231.1055D. doi:10.1126/science.3945817. PMID 3945817.
^ Jump up to: a b Gene Wars, Op.Cit. p. 102.
Jump up ^ "Search". georgetown.edu.
Jump up ^ "President Clinton Awards the Presidential Citizens Medals". nara.gov.
Jump up ^ http://www.wpainc.com/Archive/Trivelpiece/HGP%20Presenation.jpg
Jump up ^ DeLisi, Charles (2008). "Meetings that changed the world: Santa Fe 1986: Human genome baby-steps". Nature. 455 (7215): 876–877. Bibcode:2008Natur.455..876D. doi:10.1038/455876a. PMID 18923499.
Jump up ^ DeLisi, Charles (1988). "The Human Genome Project". American Scientist. 76: 488. Bibcode:1988AmSci..76..488D.
Jump up ^ DeLisi, Charles (2001). "Genomes: 15 Years Later A Perspective by Charles Deli, HEP Pioneer". Human Genome News. 11: 3–4. Archived from the original on September 4, 2004. Retrieved 2005-02-03.
^ Jump up to: a b "About the Human Genome Project: What is the Human Genome Project". The Human Genome Management Information System (HGMIS). 2011-07-18. Retrieved 2011-09-02.
^ Jump up to: a b c Human Genome Information Archive. "About the Human Genome Project". U.S. Department of Energy & Human Genome Project program. Retrieved 1 August 2013.
Jump up ^ Collins F; Galas D (1993-10-01). "A New Five-Year Plan for the United States: Human Genome Program". National Human Genome Research Institute. Retrieved 1 August 2013.
Jump up ^ "White House Press Release". Retrieved 2006-07-22.
Jump up ^ Noble, Ivan (2003-04-14). "Human genome finally complete". BBC News. Retrieved 2006-07-22.
Jump up ^ Kolata, Gina (15 April 2013). "Human Genome, Then and Now". The New York Times. Retrieved 24 April 2014.
Jump up ^ "Guardian Unlimited |UK Latest | Human Genome Project finalised". The Guardian. London. Retrieved 2006-07-22. [dead link]
Jump up ^ "The Human Genome Project FAQ". Genoscope. Centre National de Séquençage. Retrieved 12 February 2015.
Jump up ^ Schmutz, Jeremy; Wheeler, Jeremy; Grimwood, Jane; Dickson, Mark; Yang, Joan; Caoile, Chenier; Bajorek, Eva; Black, Stacey; Chan, Yee Man; Denys, Mirian; Escobar, Julio; Flowers, Dave; Fotopulos, Dea; Garcia, Carmen; Gomez, Maria; Gonzales, Eidelyn; Haydu, Lauren; Lopez, Frederick; Ramirez, Lucia; Retterer, James; Rodriguez, Alex; Rogers, Stephanie; Salazar, Angelica; Tsai, Ming; Myers, Richard M. (2004). "Quality assessment of the human genome sequence". Nature. 429 (6990): 365–368. Bibcode:2004Natur.429..365S. doi:10.1038/nature02390. PMID 15164052.
Jump up ^ "Landmark Human Genome Project Papers". ornl.gov.
Jump up ^ "An Overview of the Human Genome Project".
Jump up ^ "Ensembl Genome Browser". ensembl.org.
Jump up ^ Mardis, E. (2008). "The impact of next-generation sequencing technology on genetics". Trends in Genetics. 24 (3): 133–41. doi:10.1016/j.tig.2007.12.007. PMID 18262675.
Jump up ^ Mihaela Pertea & Steven Salzberg (2010). "Between a chicken and a grape: estimating the number of human genes". Genome Biology. 11: 206. doi:10.1186/gb-2010-11-5-206. PMC 2898077  . PMID 20441615.
Jump up ^ Venter, JC; et al. (2001). "The sequence of the human genome". Science. 291: 1304–1351. Bibcode:2001Sci...291.1304V. doi:10.1126/science.1058040. PMID 11181995. CS1 maint: Explicit use of et al. (link)
Jump up ^ International Human Genome Sequencing Consortium (IHGSC) (2004). "Finishing the euchromatic sequence of the human genome". Nature. 431 (7011): 931–945. Bibcode:2004Natur.431..931H. doi:10.1038/nature03001. PMID 15496913.
Jump up ^ Spencer, Geoff (20 December 2004). "International Human Genome Sequencing Consortium Describes Finished Human Genome Sequence". NIH Nes Release. National Institutes of Health.
Jump up ^ Bryant, J. A (2007). Design and information in biology: From molecules to systems. p. 108. ISBN 9781853128530. ...brought to light about 1200 protein families. Only 94 protein families, or 7%, appear to be vertebrate specific
^ Jump up to: a b Wellcome Sanger Institute. "The Human Genome Project: a new reality". Wellcome Trust Sanger Institute, Genome Research Limited. Archived from the original on 2013-08-01. Retrieved 1 August 2013.
^ Jump up to: a b c Naidoo N; Pawitan Y; Soong R; Cooper DN; Ku CS (2011). "Human genetics and genomics a decade after the release of the draft sequence of the human genome". Hum Genomics. 5 (6): 577–622. doi:10.1186/1479-7364-5-6-577. PMC 3525251  . PMID 22155605.
Jump up ^ "Celera: A Unique Approach to Genome Sequencing". ocf.berkeley.edu. Biocomputing. 2006. Retrieved 1 August 2013.
Jump up ^ Davidson College (2002). "Sequencing Whole Genomes: Hierarchical Shotgun Sequencing v. Shotgun Sequencing". bio.davidson.edu. Department of Biology, Davidson College. Retrieved 1 August 2013.
Jump up ^ Human Genome Project Information Archive (2013). "U.S. & International HGP Research Sites". U.S. Department of Energy & Human Genome Project. Retrieved 1 August 2013.
Jump up ^ Vizzini, Casimiro (March 19, 2015). "The Human Variome Project: Global Coordination in Data Sharing". Science & Diplomacy. 4 (1).
Jump up ^ Venter, J. C.; Adams, M. D.; Myers, E. W.; Li, P. W.; Mural, R. J.; Sutton, G. G.; Smith, H. O.; Yandell, M.; Evans, C. A. (2001-02-16). "The sequence of the human genome". Science. 291 (5507): 1304–1351. Bibcode:2001Sci...291.1304V. doi:10.1126/science.1058040. ISSN 0036-8075. PMID 11181995.
Jump up ^ Roach JC; Boysen C; Wang K; Hood L (1995). "Pairwise end sequencing: a unified approach to genomic mapping and sequencing". Genomics. 26 (2): 345–353. doi:10.1016/0888-7543(95)80219-C. PMID 7601461.
Jump up ^ Center for Biomolecular Science & Engineering. "The Human Genome Project Race". Center for Biomolecular Science and Engineering. Retrieved 1 August 2013.
Jump up ^ International Human Genome Sequencing Consortium (2001). "Initial sequencing and analysis of the human genome" (PDF). Nature. 409 (6822): 860–921. Bibcode:2001Natur.409..860L. doi:10.1038/35057062. PMID 11237011.
Jump up ^ Venter, JC; et al. (2001). "The sequence of the human genome" (PDF). Science. 291 (5507): 1304–1351. Bibcode:2001Sci...291.1304V. doi:10.1126/science.1058040. PMID 11181995.
Jump up ^ Osoegawa, Kazutoyo; Mammoser, AG; Wu, C; Frengen, E; Zeng, C; Catanese, JJ; De Jong, PJ (2001). "A Bacterial Artificial Chromosome Library for Sequencing the Complete Human Genome". Genome Research. 11 (3): 483–96. doi:10.1101/gr.169601. PMC 311044  . PMID 11230172.
Jump up ^ Tuzun, E; et al. (2005). "Fine-scale structural variation of the human genome". Nature Genetics. 37 (7): 727–737. doi:10.1038/ng1562. PMID 15895083.
Jump up ^ Kennedy D (2002). "Not wicked, perhaps, but tacky". Science. 297 (5585): 1237. doi:10.1126/science.297.5585.1237. PMID 12193755.
Jump up ^ Venter D (2003). "A Part of the Human Genome Sequence". Science. 299 (5610): 1183–4. doi:10.1126/science.299.5610.1183. PMID 12595674.
Jump up ^ Wadman, Meredith (2008-04-16). "James Watson's genome sequenced at high speed". Nature News. 452 (7189): 788–788. Bibcode:2008Natur.452R....W. doi:10.1038/452788b.
Jump up ^ Gonzaga-Jauregui C; Lupski JR; Gibbs RA (2012). "Human genome sequencing in health and disease". Annu Rev Med. 63 (1): 35–61. doi:10.1146/annurev-med-051010-162644. PMC 3656720  . PMID 22248320.
Jump up ^ Snyder M, Du J; Gerstein M (2012). "Personal genome sequencing: current approaches and challenges". Genes Dev. 24 (5): 423–431. doi:10.1101/gad.1864110. PMC 2827837  . PMID 20194435.
Jump up ^ Lander ES (2011). "Initial impact of the sequencing of the human genome". Nature. 470 (7333): 187–197. Bibcode:2011Natur.470..187L. doi:10.1038/nature09792. PMID 21307931.
Jump up ^ Peng, J; Sun, E; Nevo, D (2011). "Domestication Evolution, Genetics And Genomics In Wheat". Molecular Breeding. 28 (3): 281–301. doi:10.1007/s11032-011-9608-4.
Jump up ^ Greely, Henry (1992). The Code of Codes: Scientific and Social Issues in the Human Genome Project. Cambridge, Massachusetts: Harvard University Press. pp. 264–65. ISBN 0-674-13646-2.
Jump up ^ US Department of Health and Human Services. "Understanding Health Information Privacy".
Jump up ^ Genetics Home Reference (2013). "What were some of the ethical, legal, and social implications addressed by the Human Genome Project?". ghr.nlm.nih.gov. Retrieved 1 August 2013.
Jump up ^ "ELSI Research Program Fact Sheet - National Human Genome Research Institute (NHGRI)". www.genome.gov. Retrieved 2016-09-27.
Jump up ^ Rheinberger, H.J. (2000). Living and Working with the New Medical Technologies. Cambridge: Cambridge University Press. p. 20.
Further reading[edit]
McElheny, Victor K. (2010). Drawing the Map of Life: Inside the Human Genome Project. Basic Books. ISBN 978-0-465-03260-0. 361 pages. Examines the intellectual origins, history, and motivations of the project to map the human genome; draws on interviews with key figures.
Collins, F. (2006). The Language of God: A Scientist Presents Evidence for Belief. Free Press. ISBN 0-7432-8639-1. OCLC 65978711.
Venter, J. Craig (October 18, 2007). A Life Decoded: My Genome: My Life. New York, New York: Viking Adult. ISBN 0-670-06358-4. OCLC 165048736.
Cook-Deegan, Robert (1994). The Gene Wars: Science, Politics, and the Human Genome. New York: W W Norton.
External links[edit]
Human Genome Project — official information page
Wikinews has related news: Mexico presents first population-wide genome map for a Latin country
Wikibooks has a book on the topic of: Genes, Technology and Policy
National Human Genome Research Institute (NHGRI). NHGRI led the National Institutes of Health's contribution to the International Human Genome Project. This project, which had as its primary goal the sequencing of the three thousand million base pairs that make up human genome, was successfully completed in April 2003.
Human Genome News. Published from 1989 to 2002 by the US Department of Energy, this newsletter was a major communications method for coordination of the Human Genome Project. Complete online archives are available.
The HGP information pages Department of Energy's portal to the international Human Genome Project, Microbial Genome Program, and Genomics:GTL systems biology for energy and environment
yourgenome.org: The Sanger Institute public information pages has general and detailed primers on DNA, genes and genomes, the Human Genome Project and science spotlights.
Ensembl project, an automated annotation system and browser for the human genome
UCSC genome browser, This site contains the reference sequence and working draft assemblies for a large collection of genomes. It also provides a portal to the ENCODE project.
Nature magazine's human genome gateway, including the HGP's paper on the draft genome sequence
Wellcome Trust Human Genome website A free resource allowing you to explore the human genome, your health and your future.
Learning about the Human Genome. Part 1: Challenge to Science Educators. ERIC Digest.
Learning about the Human Genome. Part 2: Resources for Science Educators. ERIC Digest.
Patenting Life by Merrill Goozner
Prepared Statement of Craig Venter of Celera Venter discusses Celera's progress in deciphering the human genome sequence and its relationship to healthcare and to the federally funded Human Genome Project.
Cracking the Code of Life Companion website to 2-hour NOVA program documenting the race to decode the genome, including the entire program hosted in 16 parts in either QuickTime or RealPlayer format.
Lone Dog L (1999). "Whose genes are they? The Human Genome Diversity Project". J Health Soc Policy. 10 (4): 51–66. doi:10.1300/J045v10n04_04. PMID 10538186.
Bioethics Research Library Numerous original documents at Georgetown University.
Works by archive
Works by Human Genome Project at Project Gutenberg
Project Gutenberg hosts e-texts for Human Genome Project, titled Human Genome Project, Chromosome Number # (# denotes 01-22, X and Y). This information is raw sequence, released in November 2002; access to entry pages with download links is available through https://www.gutenberg.org/etext/3501 for Chromosome 1 sequentially to https://www.gutenberg.org/etext/3524 for the Y Chromosome. Note that this sequence might not be considered definitive due to ongoing revisions and refinements. In addition to the chromosome files, there is a supplementary information file dated March 2004 which contains additional sequence information.
Works by or about Human Genome Project at Internet Archive
[show]
v
t
e
Omics
Genomics
Cognitive genomics
Computational genomics
Comparative genomics
Functional genomics
Genome project
Human Genome Project
Metagenomics
Personal genomics
Social genomics
Structural genomics
Bioinformatics
Biochip
Cheminformatics
Chemogenomics
Connectomics
Glycomics
Immunomics
Lipidomics
Metabolomics
Microbiomics
Nutrigenomics
Paleopolyploidy
Pharmacogenetics
Pharmacogenomics
Systems biology
Toxicogenomics
Transcriptomics
Structural biology
Proteomics
Human Proteome Project
Call-map proteomics
Structure-based drug design
Expression proteomics
Research tools
2-D electrophoresis
Mass spectrometer
Electrospray ionization
Matrix-assisted laser desorption ionization
Matrix-assisted laser desorption ionization-time of flight mass spectrometer
Microfluidic-based tools
Isotope affinity tags
Organizations
National Institutes of Health (USA)
DNA Data Bank of Japan (JP)
European Molecular Biology Laboratory (EU)
Sanger Centre (EN)
[show]
v
t
e
Personal genomics
Data collection
Biobank
Biological database
Field concepts
Biological specimen
De-identification
Human genetic variation
Genetic linkage
Single-nucleotide polymorphisms
Identity by descent
Genetic disorder
Applications
Personalized medicine
Predictive medicine
Genetic epidemiology
Analysis techniques
Whole genome sequencing
Genome-wide association study
SNP array
Major projects
Human Genome Project
International HapMap Project
1000 Genomes Project
Human Genome Diversity Project
[show]
v
t
e
Ape-related articles
Extant ape species
Human (Homo sapiens)
Chimpanzee (Pan spp.)
Bonobo (Pan paniscus)
Common chimpanzee (Pan troglodytes)
Gorilla (Gorilla spp.)
Western gorilla (Gorilla gorilla)
Eastern gorilla (Gorilla beringei)
Orangutan (Pongo spp.)
Bornean orangutan (Pongo abelii)
Sumatran orangutan (Pongo pygmaeus)
Tapanuli orangutan (Pongo tapanuliensis)
Gibbon (family: Hylobatidae)
Study of apes
Ape language
Dian Fossey
Birutė Galdikas
Jane Goodall
Chimpanzee genome project
Human genome project
Neanderthal genome project
Willie Smits
Lone Drøscher Nielsen
Ian Redmond
Elgin Center
Iowa Primate Learning Sanctuary
Borneo Orangutan Survival
Legal and social status
Personhood
Research ban
Kinshasa Declaration
Great Ape Project
Survival Project
International Primate Day
See also
List of individual apes (non-human)
Apes in space (non-human)
Bigfoot
Bushmeat
Chimpanzee–human last common ancestor
List of fictional primates (non-human)
Great apes
Human evolution
Monkey Day
Mythic humanoids
Yeren
Yeti
[show]
v
t
e
Wellcome Trust
Centres and institutes
Current
Francis Crick Institute
Gurdon Institute
Sainsbury Wellcome Centre for Neural Circuits and Behaviour
Science Learning Centres
WTC for Cell-Matrix Research
WTC for Gene Regulation and Expression
WTC for Human Genetics
WTC for Mitochondrial Research
WTC for Molecular Parasitology
WTC for Neuroimaging
WTC for Stem Cell Research
Wellcome Sanger Institute
Former
Wellcome Trust Centre for the History of Medicine
Wellcome Research Laboratories
Projects and facilities
1000 Genomes Project
Big Picture
Cambridge Biomedical Campus
Cancer Genome Project
ChEMBL
COSMIC cancer database
DECIPHER
Diamond Light Source
eLife
Ensembl
Farmcare
Genome Reference Consortium
Human Genome Project
Malawi Liverpool Wellcome Trust
MEROPS
Pfam
Rfam
UK Biobank
Wellcome Collection
Wellcome Genome Campus
Wellcome Library
WormBase
Board of Governors
Tobias Bonhoeffer
Alan Brown
Damon Buffini
William Burns
Kay Davies
Michael Ferguson
Bryan Grenfell
Richard Hynes
Anne Johnson
Eliza Manningham-Buller
Peter Rigby
Executive Board
Jeremy Farrar
Stephen Caddick
Simon Chaplin
Tim Livett
Clare Matterson
Ted Smith
Danny Truell
Former directors
Peter Williams (1965–1991)
Bridget Ogilvie (1991–1998)
Michael Dexter (1998–2008)
Mark Walport (2003–2013)
Other key people
William Castell
Dominic Cadbury
Harold Cook
Oliver Franks
Roger Gibbs
Henry Dale
Roy Porter
David Steel
David Stuart
John Sulston
Henry Wellcome
Awards and fellowships
Capital Awards
Collaborative Awards in Science
Investigator Awards in Science
Institutional Strategic Support Fund
Science Strategic Award
Sir Henry Dale Fellowship
Sir Henry Wellcome Postdoctoral Fellowship
Wellcome Book Prize
Wellcome Image Awards
Wellcome Trust Centre
Wellcome Trust Principal Research Fellow
Wellcome Trust Senior Research Fellow
Category
[show]
v
t
e
History of biology
Fields,
disciplines
Agricultural science
Anatomy
Biochemistry
Biotechnology
Botany
Ecology
Evolutionary thought
Genetics
Geology
Immunology
Medicine
Model organisms
Molecular biology
Molecular evolution
Paleontology
Phycology
Plant systematics
RNA biology
Zoology (since 1859)
Zoology (through 1859)
Institutions
Cold Spring Harbor Laboratory
Laboratory of Molecular Biology
Marine Biological Laboratory
Max Planck Society
Pasteur Institute
Rockefeller University
Rothamsted Experimental Station
Stazione Zoologica
Woods Hole Oceanographic Institute
Theories,
concepts
Germ theory of disease
Central dogma of molecular biology
Darwinism
Great chain of being
Hierarchy of life
Lamarckism
One gene–one enzyme hypothesis
Protocell
RNA world hypothesis
Sequence hypothesis
Spontaneous generation
History
Classical
antiquity
Aristotle
Aristotle's biology
On Generation and Corruption
History of Animals
Theophrastus
Historia Plantarum
Dioscorides
De Materia Medica
Galen
Renaissance,
Early Modern
Conrad Gessner
Historia animalium
Andreas Vesalius
De humani corporis fabrica
William Harvey
De Motu Cordis
Frederik Ruysch
Jan Swammerdam
Antonie van Leeuwenhoek
Robert Hooke
Micrographia
Francesco Redi
Evolution
19th
century
Linnaeus
Systema Naturae
Buffon
Histoire Naturelle
Lamarck
Philosophie Zoologique
Humboldt
Charles Lyell
Principles of Geology
Charles Darwin
On the Origin of Species
The Descent of Man
Gregor Mendel
Alfred Russel Wallace
Martinus Beijerinck
Henry Walter Bates
Modern
synthesis
William Bateson
Theodosius Dobzhansky
Genetics and the Origin of Species
R. A. Fisher
E. B. Ford
J. B. S. Haldane
Ernst Mayr
Thomas Hunt Morgan
George Gaylord Simpson
Hugo de Vries
Sewall Wright
Recent
Stephen Jay Gould
W. D. Hamilton
Lynn Margulis
Aleksandr Oparin
George C. Williams
Carl Woese
Microbiology
Ferdinand Cohn
Alexander Fleming
Felix d'Herelle
Robert Koch
Louis Pasteur
Lazzaro Spallanzani
Sergei Winogradsky
Develop. biol.,
Evo-devo
Karl Ernst von Baer
Gavin de Beer
Sean B. Carroll
Scott F. Gilbert
Walter Gehring
Ernst Haeckel
François Jacob
Edward B. Lewis
Jacques Monod
Christiane Nüsslein-Volhard
Eric Wieschaus
E. B. Wilson
Genetics,
Molecular
biology
Experiments
Griffith's (1928)
Luria–Delbrück (1943)
Avery–MacLeod–McCarty (1944)
Miller–Urey (1952)
Hershey–Chase (1952)
Meselson–Stahl (1958)
Crick, Brenner et al. (1961)
Nirenberg–Matthaei (1961)
Nirenberg–Leder (1964)
People
Barbara McClintock
George Beadle
Seymour Benzer
Rosalind Franklin
Photo 51
James D. Watson and Francis Crick
"Molecular structure of Nucleic Acids"
Linus Pauling
"Sickle Cell Anemia, a Molecular Disease"
Fred Sanger
Max Perutz
John Kendrew
Sydney Brenner
Joshua Lederberg
Walter Gilbert
Kary Mullis
Emmanuelle Charpentier
Jennifer Doudna
Ecology
Rachel Carson
Frederic Clements
Charles Elton
Henry Gleason
Arthur Tansley
Eugenius Warming
Ethology
Niko Tinbergen
Karl von Frisch
Konrad Lorenz
Frans de Waal
Jane Goodall
Ivan Pavlov
Related
History of science
Philosophy of biology
Teleology
Ethnobotany
Eugenics
History of the creation-evolution controversy
Human Genome Project
Humboldtian science
Natural history
Natural philosophy
Natural theology
Relationship between religion and science
Timeline of biology and organic chemistry
Category
Commons
Portal
Biotechnology portal
Molecular and cellular biology portal
Biology portal
Medicine portal
Retrieved from "https://en.wikipedia.org/w/index.php?title=Human_Genome_Project&oldid=834010272"
Categories:
Human genome projects
Human Genome Project scientists
Biotechnology
Genetics
Life sciences industry
Wellcome Trust
Projects established in 1990
1990 in biotechnology
1990 in biology
1990 in science
1990 establishments in the United States
2003 in biotechnology
Hidden categories:
Pages with citations lacking titles
Pages with citations having bare URLs
All articles with dead external links
Articles with dead external links from August 2014
CS1 maint: Explicit use of et al.
Articles needing additional references from April 2017
All articles needing additional references
All articles with unsourced statements
Articles with unsourced statements from January 2015
Articles with Project Gutenberg links
Articles with Internet Archive links
Navigation menu
Personal tools
Not logged in
Talk
Contributions
Create account
Log in
Namespaces
Article
Talk
Variants
Views
Read
Edit
View history
More
Search
Navigation
Main page
Contents
Featured content
Current events
Random article
Donate to Wikipedia
Wikipedia store
Interaction
Help
About Wikipedia
Community portal
Recent changes
Contact page
Tools
What links here
Related changes
Upload file
Special pages
Permanent link
Page information
Wikidata item
Cite this page
Print/export
Create a book
Download as PDF
Printable version
In other projects
Wikimedia Commons
Languages
العربية
বাংলা
Bân-lâm-gú
Bosanski
Català
Čeština
Dansk
Deutsch
Eesti
Español
فارسی
Français
Galego
한국어
Hrvatski
Bahasa Indonesia
Italiano
עברית
Lietuvių
Magyar
Македонски
മലയാളം
Bahasa Melayu
Nederlands
日本語
Norsk
Polski
Português
Română
Русский
සිංහල
Slovenčina
Slovenščina
کوردی
Српски / srpski
Srpskohrvatski / српскохрватски
Suomi
Svenska
தமிழ்
ไทย
Türkçe
Українська
Tiếng Việt
中文
35 more
Edit links
This page was last edited on 3 April 2018, at 13:24.
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.
Privacy policy
About Wikipedia
Disclaimers
Contact Wikipedia
Developers
Cookie statement
Mobile view
Enable previews

values: mw-page-base noprint

values: mw-head-base noprint

values: firstHeading firstHeading en

values: siteSub noprint

values: jump-to-nav mw-jump

attrs: alt src width height class srcset data-file-width data-file-height

attrs: href class title

attrs: class style

value: b

values:
/wiki/Scientific_research
mw-redirect
Scientific research

values:
/wiki/Base_pairs
mw-redirect
Base pairs

values:
/wiki/Peter_Domenici
mw-redirect
Peter Domenici

values:
/wiki/Reagan_Administration
mw-redirect
Reagan Administration

values: United States National Institutes of Health

values: National Human Genome Research Institute

values:
/wiki/Genome_Project
mw-redirect
Genome Project

values:
/wiki/Chromosome_1_(human)
mw-redirect
Chromosome 1 (human)

values: List of distinct cell types in the adult human body

values: Edit section: Applications and proposed benefits

values: National Center for Biotechnology Information

values: Help:Introduction to referencing with Wiki Markup/1

values: plainlinks metadata ambox ambox-content ambox-Refimprove

values:
/wiki/Genome_annotation
mw-redirect
Genome annotation

values: This claim needs references to reliable sources. (January 2015)

values:
noprint Inline-Template Template-Fact
white-space:nowrap;

values: Edit section: Public versus private approaches

values:
/wiki/Research_funding
mw-redirect
Research funding

values:
/wiki/News_release
mw-redirect
News release

values:
/wiki/DNA_library
mw-redirect
DNA library

values: Pieter J. de Jong's (page does not exist)

values: Center for the Study of Human Polymorphisms

values:
/wiki/Blood_clotting
mw-redirect
Blood clotting

values:
/wiki/Etiology_(medicine)
mw-redirect
Etiology (medicine)

values: Edit section: Ethical, legal and social issues

values: Health Insurance Portability and Accountability Act

values:
/wiki/Chimpanzee_Genome_Project
mw-redirect
Chimpanzee Genome Project

values:
/wiki/Genome_Compiler
mw-redirect
Genome Compiler

values:
/wiki/Human_Cytome_Project
mw-redirect
Human Cytome Project

values:
/wiki/Neanderthal_Genome_Project
mw-redirect
Neanderthal Genome Project

values:
/wiki/Sanger_Institute
mw-redirect
Sanger Institute

values:
/wiki/The_Genographic_Project
mw-redirect
The Genographic Project

values:
nofollow
https://www.pbs.org/wgbh/nova/genome/
external text

values:
/wiki/Public_Broadcasting_Service
mw-redirect
Public Broadcasting Service

values:
nofollow
external text
http://www.genome.gov/11006943

values:
nofollow
external text
//doi.org/10.1038%2F455876a

values:
/wiki/PubMed_Identifier
mw-redirect
PubMed Identifier

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18923499

values:
nofollow
external text
//doi.org/10.1126%2Fscience.3945817

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/3945817

values:
nofollow
external text
http://www.genome.gov/10001476

values:
nofollow
external text
//doi.org/10.1038%2Fnature02390

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15164052

values:
nofollow
external text
http://genome.ucsc.edu

values:
nofollow
external text
http://www.ensembl.org

values:
nofollow
external text
//doi.org/10.1016%2Fj.tig.2007.12.007

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18262675

values:
nofollow
external text
//doi.org/10.1186%2Fgb-2010-11-5-206

attrs: alt src title width height srcset data-file-width data-file-height

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/20441615

values:
nofollow
external text
//doi.org/10.1126%2Fscience.1058040

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11181995

values: Category:CS1 maint: Explicit use of et al.

values:
nofollow
external text
//doi.org/10.1038%2Fnature03001

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15496913

values:
nofollow
external text
http://www.genome.gov/12513430

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/22155605

values:
nofollow
external text
//www.worldcat.org/issn/0036-8075

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/7601461

values:
nofollow
external text
//doi.org/10.1038%2F35057062

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11237011

values:
nofollow
external text
//doi.org/10.1101%2Fgr.169601

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11230172

values:
nofollow
external text
//doi.org/10.1038%2Fng1562

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15895083

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/12193755

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/12595674

values:
nofollow
external text
//doi.org/10.1038%2F452788b

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/22248320

values:
nofollow
external text
//doi.org/10.1101%2Fgad.1864110

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/20194435

values:
nofollow
external text
//doi.org/10.1038%2Fnature09792

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/21307931

values: -moz-column-width: 30em; -webkit-column-width: 30em; column-width: 30em; list-style-type: decimal;

values:
nofollow
external text
//www.worldcat.org/oclc/65978711

values:
/wiki/Viking_Adult
mw-redirect
Viking Adult

values:
nofollow
external text
//www.worldcat.org/oclc/165048736

values: wikinews:Mexico presents first population-wide genome map for a Latin country

values:
nofollow
external text
http://www.genome.gov

values:
nofollow
http://www.doegenomes.org/
external text

values:
nofollow
http://www.yourgenome.org/
external text

values:
nofollow
http://www.ensembl.org/
external text

values:
nofollow
http://www.nature.com/genomics/human/
external text

values:
nofollow
external text
//doi.org/10.1300%2FJ045v10n04_04

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10538186

values:
nofollow
external free
https://www.gutenberg.org/etext/3501

values:
nofollow
external free
https://www.gutenberg.org/etext/3524

values:
nofollow
external text
https://www.gutenberg.org/etext/11799

values:
/wiki/Human_Proteome_Project
mw-redirect
Human Proteome Project

values:
/wiki/Call-map_proteomics
mw-redirect
Call-map proteomics

values:
/wiki/Structure-based_drug_design
mw-redirect
Structure-based drug design

values:
/wiki/Expression_proteomics
mw-redirect
Expression proteomics

values:
/wiki/2-D_electrophoresis
mw-redirect
2-D electrophoresis

values:
/wiki/Mass_spectrometer
mw-redirect
Mass spectrometer

values: Matrix-assisted laser desorption ionization

values:
/wiki/Microfluidic-based_tools
mw-redirect
Microfluidic-based tools

values:
/wiki/Isotope_affinity_tags
mw-redirect
Isotope affinity tags

values:
/wiki/Sanger_Centre
mw-redirect
Sanger Centre

values: nowraplinks collapsible autocollapse navbox-inner

values:
/wiki/Single-nucleotide_polymorphisms
mw-redirect
Single-nucleotide polymorphisms

values:
navbox-image
4
width:1px;padding:0px 0px 0px 2px

values:
row
navbox-group
width:1%;text-align: left;

values: Wellcome Trust Centre for Cell-Matrix Research

values: Wellcome Trust Centre for Gene Regulation and Expression

values: Wellcome Trust Centre for Human Genetics

values: Wellcome Trust Centre for Stem Cell Research

values: Wellcome Trust Centre for the History of Medicine

values:
navbox-image
7
width:1px;padding:0px 0px 0px 2px

values: Wellcome Trust Principal Research Fellow

values: nowraplinks collapsible collapsed navbox-inner

values:
/wiki/History_of_zoology_(since_1859)
mw-redirect
History of zoology (since 1859)

values:
/wiki/File:HomunculusLarge.png
image
The homonculus

values:
navbox-image
5
width:1px;padding:0px 0px 0px 2px

values:
/wiki/Rothamsted_Experimental_Station
mw-redirect
Rothamsted Experimental Station

values:
/wiki/RNA_world_hypothesis
mw-redirect
RNA world hypothesis

values:
/wiki/The_Renaissance
mw-redirect
The Renaissance

values:
/wiki/Early_Modern
mw-redirect
Early Modern

values: Exercitatio Anatomica de Motu Cordis et Sanguinis in Animalibus

values:
/wiki/The_Descent_of_Man
mw-redirect
The Descent of Man

values:
/wiki/Felix_d%27Herelle
mw-redirect
Felix d'Herelle

values:
/wiki/Walter_Gehring
mw-redirect
Walter Gehring

values:
/wiki/Eric_Wieschaus
mw-redirect
Eric Wieschaus

values:
/wiki/George_Wells_Beadle
mw-redirect
George Wells Beadle

values:
/wiki/James_D._Watson
mw-redirect
James D. Watson

values: Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid

values:
/wiki/Henry_Gleason
mw-redirect
Henry Gleason

values:
/wiki/Niko_Tinbergen
mw-redirect
Niko Tinbergen

values: History of the creation–evolution controversy

values: Relationship between religion and science

values: Timeline of biology and organic chemistry

values: nowraplinks hlist collapsible autocollapse navbox-inner

values:
noprint metadata navbox
navigation
Portals
font-weight:bold;padding:0.4em 2em

values: Category:Human Genome Project scientists

values: Category:1990 establishments in the United States

values: Category:Pages with citations lacking titles

values: Category:Pages with citations having bare URLs

values: Category:All articles with dead external links

values: Category:Articles with dead external links from August 2014

values: Category:Articles needing additional references from April 2017

values: Category:All articles needing additional references

values: Category:All articles with unsourced statements

values: Category:Articles with unsourced statements from January 2015

values: Category:Articles with Project Gutenberg links

values: Category:Articles with Internet Archive links

values: Discussion about edits from this IP address [ctrl-alt-n]

values: A list of edits made from this IP address [ctrl-alt-y]

values: You are encouraged to create an account and log in; however, it is not mandatory

values: You're encouraged to log in; however, it's not mandatory. [ctrl-alt-o]

values:
/wiki/Human_Genome_Project
View the content page [ctrl-alt-c]
c

values: Discussion about the content page [ctrl-alt-t]

values: p-namespaces navigation vectorTabs p-namespaces-label

values:
p-variants
navigation
vectorMenu emptyPortlet
p-variants-label

values: Past revisions of this page [ctrl-alt-h]

values: p-views navigation vectorTabs p-views-label

values:
p-cactions
navigation
vectorMenu emptyPortlet
p-cactions-label

values:
mw-wiki-logo
/wiki/Main_Page
Visit the main page

values:
/wiki/Main_Page
Visit the main page [ctrl-alt-z]
z

values: Featured content – the best of Wikipedia

values: Find background information on current events

values:
/wiki/Special:Random
Load a random article [ctrl-alt-x]
x

values: portal navigation p-navigation p-navigation-label

values: Guidance on how to use and edit Wikipedia

values: About the project, what you can do, where to find things

values: A list of recent changes in the wiki [ctrl-alt-r]

values: portal navigation p-interaction p-interaction-label

values: List of all English Wikipedia pages containing links to this page [ctrl-alt-j]

values: portal navigation p-wikibase-otherprojects p-wikibase-otherprojects-label

values: مشروع الجينوم البشري – Arabic

values: হিউম্যান জিনোম প্রজেক্ট – Bangla

values:
interlanguage-link interwiki-bn
display: none;

values: Jîn-lūi Ûi-thoân-thé Kè-ōe – Chinese (Min Nan)

values:
interlanguage-link interwiki-zh-min-nan
display: none;

values:
interlanguage-link interwiki-bs
display: none;

values:
interlanguage-link interwiki-ca
display: none;

values:
interlanguage-link interwiki-cs
display: none;

values:
https://da.wikipedia.org/wiki/HUGO
HUGO – Danish
da
da
interlanguage-link-target

values:
interlanguage-link interwiki-da
display: none;

values:
interlanguage-link interwiki-et
display: none;

values: پروژه ژنوم انسان – Persian

values:
interlanguage-link interwiki-fa
display: none;

values:
interlanguage-link interwiki-gl
display: none;

values: 인간 게놈 프로젝트 – Korean

values:
interlanguage-link interwiki-ko
display: none;

values:
interlanguage-link interwiki-hr
display: none;

values:
interlanguage-link interwiki-it
display: none;

values: פרויקט גנום האדם – Hebrew

values:
interlanguage-link interwiki-lt
display: none;

values:
interlanguage-link interwiki-hu
display: none;

values: Проект за човечки геном – Macedonian

values:
interlanguage-link interwiki-mk
display: none;

values: ഹ്യുമൻ ജിനോം പ്രൊജക്റ്റ് – Malayalam

values:
interlanguage-link interwiki-ml
display: none;

values: ヒトゲノム計画 – Japanese

values:
interlanguage-link interwiki-ja
display: none;

values:
interlanguage-link interwiki-no
display: none;

values: Projekt poznania ludzkiego genomu – Polish

values:
interlanguage-link interwiki-ro
display: none;

values: Проект «Геном человека» – Russian

values: මානව ගෙනෝම ව්‍යාපෘතිය – Sinhala

values:
interlanguage-link interwiki-si
display: none;

values:
interlanguage-link interwiki-sk
display: none;

values:
interlanguage-link interwiki-sl
display: none;

values: پرۆژەی جینۆمی مرۆڤ – Central Kurdish

values:
interlanguage-link interwiki-ckb
display: none;

values:
interlanguage-link interwiki-sr
display: none;

values: Projekat ljudskog genoma – Serbo-Croatian

values:
interlanguage-link interwiki-sh
display: none;

values:
interlanguage-link interwiki-fi
display: none;

values: மனித மரபணுத்தொகைத் திட்டம் – Tamil

values:
interlanguage-link interwiki-ta
display: none;

values: โครงการจีโนมมนุษย์ – Thai

values:
interlanguage-link interwiki-th
display: none;

values:
interlanguage-link interwiki-tr
display: none;

values: Проект геному людини – Ukrainian

values:
interlanguage-link interwiki-uk
display: none;

values:
interlanguage-link interwiki-vi
display: none;

values: 人类基因组计划 – Chinese

values:
suggestions
display: none; font-size: 13px;

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-Human_Genome_Project rootpage-Human_Genome_Project skin-vector action-view