Difference between revisions of "Timeline of model organisms"

Latest revision as of 21:33, 27 October 2024

This is a model organisms, which are non-human species extensively studied in biological research to gain insights into various processes that can be applicable to other organisms, including humans. These organisms are selected for their ease of maintenance, rapid life cycles, and well-characterized genetics. They play a crucial role in fields such as genetics, developmental biology, and neuroscience.

Sample questions

The following are some interesting questions that can be answered by reading this timeline:

Big picture

Time period	Development summary	More details
Ancient Times to Early 19th Century	Pre-Modern Era	During this period, early naturalists and philosophers such as Aristotle observe animals and plants, focusing primarily on descriptive biology. Humans and domestic animals are studied for agricultural, medical, and practical purposes, with limited experimentation. Comparative anatomy begins to emerge with scientists like Andreas Vesalius, who uses human cadavers and animals for anatomical studies, though the approach is largely observational and lacks the rigor of controlled experiments.^[1]
Mid-19th Century to Early 20th Century	Early Experimental Biology	The mid-19th century marks a turning point with the advent of systematic biological experimentation. Gregor Mendel’s experiments with pea plants in the 1850s uncover the basic laws of inheritance, forming the foundation of genetics. By the early 20th century, Thomas Hunt Morgan’s introduction of Drosophila melanogaster in genetics research help establish the chromosomal theory of inheritance, making the fruit fly a pioneering model organism. This period also sees early embryological studies using frogs and chickens, as researchers begin to explore developmental biology in a more structured way.^[2]
Mid-20th Century to Late 20th Century	The Rise of Molecular Biology	The mid-20th century brings revolutionary advancements in molecular biology, with model organisms playing central roles. Bacteria like Escherichia coli becomes vital for studying DNA replication, gene expression, and molecular genetics, while Caenorhabditis elegans provides insights into developmental biology and programmed cell death. Mice emerged as key models for human disease research, especially in cancer, immunology, and developmental studies. This era is characterized by the discovery of the DNA double helix, the rise of recombinant DNA technology in the 1970s, and the first successful gene cloning experiments, all of which rely on these model organisms.
Late 20th Century to Present	Genomics and Systems Biology Era	With the late 20th century comes the sequencing of genomes, including those of model organisms like Drosophila, C. elegans, and mice, ushering in the genomics era. New technologies like CRISPR-Cas9 revolutionize gene editing, allowing scientists to modify genes with precision in organisms like zebrafish and mice. This period sees the rise of systems biology, integrating data from various biological scales to understand complex processes such as disease mechanisms, development, and aging. Model organisms remain indispensable in fields like synthetic biology, drug discovery, and personalized medicine, as researchers now leverage the power of genomics to tackle complex biological questions on a large scale.

Full timeline

Year	Species	Event type	Details	Location/researcher affiliation
1822	Danio rerio	Discovery	Danio rerio, initially designated as Brachydanio rerio, is first described by English physician Sir Francis Hamilton in his book on the fishes of the Ganges River. Hamilton identifies ten species at that time, (the Danio genus now comprises 45 known species). D. rerio is a monophyletic species within the Cyprinidae family, characterized by a bilobate caudal fin.^[3]	United Kingdom
1833	Chlamydomonas reinhardtii	Discovery	German scientist Christian Gottfried Ehrenberg describes the genus Chlamydomonas, which includes the species Chlamydomonas reinhardtii, a single-celled green alga found in temperate soil habitats. By the mid-20th century, C. reinhardtii would become a vital model organism for investigating fundamental cellular processes, such as photosynthesis, light perception, and the structure, function, and biogenesis of cilia. Research on this alga would notably advance plant and cell biology, contributing to algal biotechnology, and enhancing our understanding of human diseases.^[4]	Germany
1853	Pisum sativum	Experiment	Gregor Mendel begins his groundbreaking experiments with pea plants, which lays the foundation for classical genetics. He focuses on easily observable traits like plant height, seed color, and flower color. Mendel employs controlled breeding, cross-pollinating plants with distinct traits to study inheritance patterns. Unlike earlier researchers, he meticulously counted and analyzed his results, identifying consistent patterns of trait transmission. His findings led to the formulation of key genetic principles, the "Law of Segregation" and the "Law of Independent Assortment," which explain how traits are passed from parents to offspring.^[5]^[6]^[7]^[8]	Czech Republic (Austrian Empire)
1857	Saccharomyces cerevisiae	Discovery	Louis Pasteur discovers Saccharomyces cerevisiae, identifying it as the key microbe in winemaking and bread baking. He classifies this yeast as a facultative anaerobe, meaning it can switch to fermentation in the absence of oxygen. This discovery would not only revolutionize food and beverage production but also establishes S. cerevisiae as a model organism in scientific research. Its simple eukaryotic structure and genetic tractability would make it a powerful tool for studying fundamental biological processes, including genetics, cell biology, and biochemistry, further enhancing our understanding of more complex organisms.^[9]^[10]	France
1900	Zea mays	Genetic validation	German botanist Carl Correns conducts experiments on Zea mays, commonly known as corn or maize. Correns confirms the findings of Gregor Mendel, an Austrian monk, regarding the principles of inheritance and genetic traits. Mendel's work, initially published in 1866, outlines the laws of inheritance based on his experiments with pea plants. Correns' validation of Mendel's findings with Zea mays provides further evidence for the existence of discrete units of inheritance, which would be later known as genes. This confirmation plays a crucial role in the establishment of modern genetics and lays the foundation for understanding heredity in plants and animals.^[11]	Germany
1902	Mus musculus	Genetic research	American biologist William Ernest Castle begins genetic studies on Mus musculus, commonly known as the house mouse. This marks the initiation of systematic genetic research on this species. Castle's work contributes to the understanding of inheritance patterns and genetic variation in mice, laying the groundwork for further investigations into the genetic basis of traits and the mechanisms of heredity. His studies on Mus musculus were instrumental in the development of mouse models for genetic research, which continue to be crucial in biomedical research and the study of human genetics.^[12]^[13]	United States
1909	Drosophila melanogaster	Gene mapping	American scientist Thomas Hunt Morgan begins his groundbreaking work with the fruit fly Drosophila melanogaster, which would become synonymous with his name. Prior to this, C. W. Woodworth and W. E. Castle had shown interest in Drosophila for genetic studies. Morgan's research with Drosophila would lead to the discovery of sex linkage of the gene for white eyes, demonstrating the phenomenon of linkage. He bred Drosophila in large quantities, facilitating the analysis of spontaneous mutations and the localization of genes. Morgan's work laid the foundation for understanding the linear arrangement of genes in chromosomes and significantly advanced the field of genetics.^[14]	United States (Columbia University)
1913	Zea mays	Research	Edgar Nelson Emerson and Roland McMillan East publish a significant paper on quantitative genetics in Zea mays. This paper marks an important milestone in the understanding of genetic principles governing quantitative traits, which are traits controlled by multiple genes and influenced by environmental factors. Emerson and East's work would contribute to the development of quantitative genetics as a field by elucidating the complex inheritance patterns of traits such as height, yield, and other quantitative characteristics in maize. Their research lays the foundation for further studies in the genetics of complex traits in various organisms.^[15]	United States
1915	Drosophila melanogaster	Publication	The Morgan Group, led by Thomas Hunt Morgan, publishes the first book on Mendelian genetics focusing on Drosophila melanogaster, commonly known as the fruit fly. This publication represents a significant milestone in the field of genetics, as it provides a comprehensive overview of the principles of Mendelian inheritance as observed in Drosophila. The book likely covers topics such as the inheritance of traits, the mapping of genes, and the understanding of genetic linkage. This work serves as a foundational resource for researchers studying genetics and paves the way for further investigations into the mechanisms of inheritance in various organisms.^[16]	United States
1927	Neurospora crassa	Discovery	Researchers Shear and Dodge make a notable discovery regarding Neurospora crassa, a type of bread mold. They identify and describe the sexual cycle of Neurospora crassa, shedding light on its reproductive mechanisms. Additionally, they characterize different mating types within the species, which are essential for sexual reproduction. This finding is significant in advancing the understanding of fungal genetics and reproductive biology. It lays the groundwork for further studies on the genetics and life cycle of Neurospora crassa, making it an essential model organism in genetic research.^[17]^[18]^[19]	United States
1930	Chlamydomonas	Genetics research	German biologist Franz Moewus pioneer biochemical genetics research on Chlamydomonas, focusing on Chlamydomonas eugametos. He claims to have identified carotenoid-related hormones that selectively activate male or female gametes and reports isolating and genetically mapping mutants along their biosynthetic pathways. Over a decade, he would analyze 200,000 zygotes for ten phenotypes, though many of his findings would be later deemed irreproducible, raising doubts about his conclusions. Despite these issues, Moewus significantly influences Chlamydomonas research, inspiring further studies and reinforcing concepts like gene-chromosome localization and the one gene-one enzyme hypothesis, which later led to the use of C. reinhardtii as a model organism.^[20]	Germany
1935	Saccharomyces cerevisiae	Foundational research	Danish biologist Øjvind Winge, later regarded as the "Father of Yeast Genetics," elucidates the life cycle of Saccharomyces cerevisiae, marking the beginning of yeast genetics research. Winge’s work in Denmark laid the foundation for subsequent studies in yeast genetics, which were later pursued by researchers such as Lindegren in the U.S. and Ephrussi in France. Meanwhile, Leupold pioneered genetic studies on Schizosaccharomyces pombe in Switzerland during the 1940s. Over the next four decades, yeast became an essential model organism in eukaryotic molecular biology, with early researchers contributing significantly to the field's growth and recognition.^[21]^[22]
1935		Genetic research	When Emil Winge begins conducting genetic research with yeast, advancements in genetics using other model organisms pave the way for his experiments. Key developments include the understanding of four-strand crossing over, which improves insights into genetic recombination; chromosome mapping, which facilitates the identification of gene locations; the discovery of lethal genes, which helps researchers understand gene functions and interactions; and the transformation of Pneumococcus, which demonstrates the principles of genetic transfer and transformation. These foundational discoveries provide a practical framework for genetic studies in yeast, ultimately contributing to the field of genetics.^[2]
1937	Paramecium	Reproduction	American biologist Tracy Sonneborn and H.S. Jennings make a significant breakthrough in Paramecium genetics by successfully establishing the conditions necessary for mating. They also define a system of mating types, with most Paramecium species exhibiting two distinct mating types. This discovery marks a key advancement in the understanding of reproduction and genetics in Paramecium species, contributing to its establishment as a model organism in cellular and genetic research. Their work lay the groundwork for future studies on sexual reproduction and genetic exchange in single-celled organisms.^[23]^[24]^[25]	United States
1939	T2 bacteriophage	One-step growth experiment	American biochemist Emory Ellis and German–American biophysicist Max Delbrück conduct a pivotal one-step growth experiment using T2 bacteriophages to investigate viral replication. They mix bacteria with sufficient viruses to ensure each host cell had a virus attached, then remove the free viruses and sample at regular intervals for a plaque assay. Initially, the number of plaques remain constant during the latent period, followed by a progressive increase, indicating that infected cells were lysing and releasing phages. This experiment illustrates the three main phases of virus replication: initiation of infection, genome replication and expression, and release of mature virions. The method would be also used to estimate burst size, although it may not account for variations in lysis timing, potentially biasing viral trait estimates.^[26]^[27]^[28]	United States
1941	Neurospora crassa	Genetics	American geneticists George Beadle and Edward Tatum's experiments with Neurospora crassa lead to the isolation of the first biochemical mutants, providing key insights into gene function. By exposing the fungus to X-rays, they induce mutations and identified strains requiring specific nutrients for growth. This indicates defects in certain metabolic pathways, and their work formulate the "one gene, one enzyme" hypothesis, which proposes that each gene is responsible for producing a single enzyme. This groundbreaking research marks a pivotal step in understanding gene roles in metabolism and would earn them the Nobel Prize in 1958.^[29]^[30]^[31]	United States
1943	Arabidopsis thaliana	Model introduction	German botanist Friedrich Laibach first promotes Arabidopsis thaliana as a model organism for genetic studies. Despite possessing ideal traits for genetic analysis, such as its small size, short generation time, and simple genome, Arabidopsis does not gain widespread popularity immediately. Laibach's pioneering efforts to use Arabidopsis for understanding genetics and development lays the groundwork for its eventual adoption as a model organism, which would later become central to plant molecular biology research.^[32]	Germany
1943	Saccharomyces cerevisiae	Genetic mapping	In St. Louis, Missouri, Carl and Gertrude Lindegren become the first to report the existence of two opposite mating types in Saccharomyces cerevisiae: a and α. Their work with heterothallic strains lays the groundwork for understanding the genetic basis of mating in yeast. Subsequently, they publish the first genetic maps of S. cerevisiae in 1949 and 1951, significantly advancing the field of yeast genetics and establishing S. cerevisiae as a key model organism for studying eukaryotic genetics and cellular processes.^[33]^[34]^[35]	United States
1944	Bacteriophage	Phage Group and Phage Treaty	German–American biophysicist Max Delbrück initiates the Phage Group, advocating for the "Phage Treaty," which encourages phage researchers to concentrate on a limited number of phage and bacterial strains under standardized experimental conditions. This initiative aims to enhance the comparability and replicability of research across different laboratories, thereby unifying the field of bacterial genetics. Delbrück's efforts would significantly advance the study of bacteriophages and lay a foundation for future research in molecular biology and genetics.^[36]	United States
1946	Escherichia coli	Genetic exchange	Joshua Lederberg and Edward Tatum discover bacterial conjugation through experiments with Escherichia coli (E. coli). They find that bacteria can exchange genetic information via a process called conjugation, which involves a bridge-like connection between two cells that facilitates the transfer of DNA from the donor to the recipient. The agent responsible for this transfer is the F ("fertility") factor, an extra-chromosomal genetic element that can replicate and move across cell membranes. For their pioneering work on bacterial genetics, Lederberg, Tatum, and George Beadle would be awarded the Nobel Prize in Physiology or Medicine in 1958. This discovery highlights that bacteria can change their genetic makeup in a manner analogous to sexual reproduction in more complex organisms, revealing conserved processes across different life forms.^[37]^[38]^[39]^[40]^[41]	United States
1949	Saccharomyces cerevisiae	Model introduction	Jean Ephrussi and colleagues discover petite mutants in Saccharomyces cerevisiae (baker's yeast). These mutants have smaller colonies compared to normal strains and arise either spontaneously or after acriflavine treatment. They lack functional mitochondrial DNA (mtDNA), resulting in impaired mitochondrial function, respiratory deficiency, and altered metabolism. The discovery is significant for mitochondrial genetics, as S. cerevisiae would become a model organism for studying the relationship between mitochondrial DNA and cellular processes, providing insights into genetic inheritance and cellular respiration in eukaryotic organisms.^[42]	France
1950	Zea mays	Publication	American scientist Barbara McClintock publishes her seminal paper titled The Origin and Behavior of Mutable Loci in Maize in the Proceedings of the National Academy of Sciences. In this work, she describes the discovery of transposable elements, or "jumping genes," which can move along chromosomes and influence genetic inheritance. McClintock's research demonstrates that genomic replication does not always follow a predictable pattern, revolutionizing the understanding of genetic inheritance. Her findings on the Suppressor-Mutator (Spm) element, which can toggle between active and inactive forms due to methylation, further highlights the complexity of gene regulation. Her groundbreaking contributions would ultimately earn her a Nobel Prize.^[43]^[44]^[45]^[46]	United States
1951	Escherichia virus Lambda	Discovery	American microbiologist Esther Lederberg, while a Ph.D. student at the University of Wisconsin, isolates bacteriophage lambda (λ), marking a significant discovery in microbial genetics. The details of this isolation and its implications for specialized transduction would be later elaborated upon in a 1953 paper co-authored with Joshua Lederberg. The discovery of λ facilitates a deeper understanding of genetic exchange mechanisms in bacteria, specifically through specialized transduction, where a bacteriophage transfers specific bacterial genes from one host to another, enhancing insights into gene mapping and bacterial genetics. This work establishes λ as a crucial model organism in genetic research.^[47]	United States
1952	Salmonella typhimurium	Discovery	Norton Zinder and Joshua Lederberg discover transduction, a process in which a virus (bacteriophage) transfers genetic material from a donor to a recipient bacterium. This breakthrough occurs at the University of Wisconsin–Madison, where they identify generalized transduction using the bacteriophage P22, which infects Salmonella typhimurium. P22's unique DNA packaging allows some virions to carry fragments of the host's DNA, which can then be transferred to another bacterial cell. This discovery is the first identification of phage-mediated gene transfer, significantly advancing microbial genetics and providing a valuable tool for studying bacterial gene exchange.^[48]^[49]^[50]^[51]^[52]	United States
1953	Tetrahymena thermophila	Model introduction	David L. Nanney first describes the MAT locus in Tetrahymena thermophila. This discovery is instrumental in understanding the organism's mating types and genetic mechanisms, further establishing Tetrahymena as a model organism for studying cellular biology and genetics.^[53]	United States
1953	Aspergillus nidulans	Model introduction	Italian-born Scottish geneticist Guido Pontecorvo and his team publish a key study in Advances in Genetics, describing both the genetic and parasexual systems of Aspergillus nidulans. This work is pivotal in establishing A. nidulans as a genetically tractable model organism for studying fungal genetics. Pontecorvo's discovery of the parasexual cycle, an alternative to sexual reproduction, allows for genetic analysis in organisms lacking a traditional sexual cycle. This groundbreaking research opens doors for genetic mapping in fungi and even has implications for studying genetic recombination in human somatic cells, shaping the future of microbial genetics and genetic analysis techniques.^[54]	United Kingdom
1954	Neurospora crassa	Genetic mapping	Barratt and colleagues publish the first major article on map construction in Neurospora crassa titled Map construction in Neurospora crassa, marking a key moment in fungal genetics. This work, which would become a foundational resource in the field, combines theoretical insights with experimental findings and provides a comprehensive compendium of N. crassa mutants. The study contributes significantly to the standardization of genetic methods, strains, and nomenclature. It also advances the understanding of meiotic phenomena like segregation and intrachromosomal recombination, paving the way for deeper insights into gene conversion and molecular recombination.^[55]	United States
1965	Arabidopsis thaliana	Model introduction	Gerhard Röbbelen organizes the first International Arabidopsis Symposium. This event marks a significant step in the recognition and promotion of Arabidopsis thaliana as a model organism in plant genetics and developmental biology. The symposium brings together researchers interested in using Arabidopsis for genetic studies, helping to solidify its status as a key model organism due to its small genome, short generation time, and ease of genetic manipulation. This symposium plays a crucial role in fostering a community of scientists focused on Arabidopsis, which would later become one of the most widely studied organisms in plant biology.^[56]	Germany
1965	Caenorhabditis elegans	Model introduction	South African biologist Sydney Brenner selects Caenorhabditis elegans as a model organism to study genetics, development, and neural function. Over the years, this nematode would become one of the most comprehensively understood metazoans, particularly in terms of anatomy, genetics, and behavior. Two major accomplishments in C. elegans research include the mapping of its complete cell lineage and the creation of a "wiring diagram" of its cell contacts through electron microscopy. Advances also include molecular cloning of developmental genes using transposon tagging and the development of a physical genome map. This knowledge serves as a foundation for future research in biology.^[57]^[58]	United Kingdom
1966		Literature	American internist and medical geneticist Victor A. McKusick publishes the first edition of Mendelian Inheritance in Man (MIM), a comprehensive knowledge base of human genes and genetic disorders. Initially developed in the early 1960s, MIM categorizes autosomal dominant, autosomal recessive, and X-linked phenotypes. The first print edition launched in 1966, with 12 editions published by 1998. In 1987, MIM would transition online as OMIM (Online Mendelian Inheritance in Man) at Johns Hopkins University, later moving to the National Center for Biotechnology Information in 1995. Today, OMIM is updated daily and serves as a crucial resource for students, researchers, and clinicians in genetics and genomics.^[59]	United States
1974	Caenorhabditis elegans	Genetic screening	Sydney Brenner conducts a groundbreaking genetic screen and creates the first genetic map for Caenorhabditis elegans (C. elegans), isolating notable mutants such as Dumpy, Squat, Long, Blistered, and Roller. His genetic map identifies over 100 loci, and he estimates the genome to contain about 2,000 essential genes. As a nematode, C. elegans serves as a vital model organism in genetics due to its short lifespan, rapid generation time, and hermaphroditic nature, allowing for efficient isolation and characterization of mutants. Brenner's research also reveals the significance of a "nonsense" codon in mRNA, crucial for protein synthesis.^[60]^[61]^[62]^[63]^[64]^[65]	United Kingdom
1980	Drosophila melanogaster	Embryonic mutagenesis	In their study published in Nature, Christiane Nüsslein-Volhard and Eric Wieschaus investigate embryonic lethal mutants of Drosophila melanogaster, identifying 15 loci that significantly alter the segmental pattern of the larva. This comprehensive screening aims to uncover genes involved in segmentation, revealing that the segmentation process encompasses multiple levels of spatial organization: the egg as a developmental unit, a repeat unit spanning two segments, and the individual segment itself. Their findings not only provide insights into the genetic architecture governing body segmentation in Drosophila but also lays a foundation for understanding developmental biology. This work contributes to the broader use of Drosophila as a model organism in genetics, influencing research on gene function and developmental processes across various species.^[66]	Germany
1981	Danio rerio	Clonal production	American molecular biologist George Streisinger et al. publish a pivotal study in Nature detailing the production of clones of homozygous diploid zebrafish (danio rerio). This research demonstrates that large-scale production of homozygous diploid zebrafish can be achieved through simple physical treatments. By cloning individual homozygotes, the researchers established a reliable method for generating genetically identical fish, which is crucial for conducting genetic analyses in vertebrates. The availability of these cloned homozygous zebrafish not only advances the understanding of genetic inheritance and development in this model organism but also paves the way for further research in genetics, developmental biology, and toxicology, solidifying zebrafish's role as a valuable tool in scientific inquiry.^[67]^[68]	United States
1984	Arabidopsis thaliana	Genome estimation	Leutwiler et al. utilize reassociation kinetics and quantitative gel blot hybridization to estimate the genome size of Arabidopsis thaliana at approximately 70 megabases (Mb). However, other techniques, including Feulgen photometry and flow cytometry, suggest a range of 0.085–0.215 picograms (pg) for the genome size. The actual genome size of A. thaliana is closer to 135 Mb, consisting of five pairs of chromosomes, making it the smallest genome among flowering plants. The complete genome sequence would be published in 2000, revealing around 25,498 predicted genes. Variations in genome size among organisms can be attributed to differences in the amplification, deletion, and divergence of repetitive sequences, highlighting the complexity of genomic structure and evolution.^[69]^[70]^[71]^[72]^[73]	United States
1996 (April)	Saccharomyces cerevisiae	Genome sequencing	The complete genome of Saccharomyces cerevisiae (baker’s yeast) strain S288C is sequenced, making it the first eukaryotic genome to be fully mapped. This significant milestone involves over 600 scientists from North America, Japan, and Europe, who collaboratively sequenced its 12 million base pairs across 16 chromosomes. The S. cerevisiae genome contains about 6,000 genes, with roughly one-third having human counterparts. This achievement underscores S. cerevisiae's role as a model organism in eukaryotic genomics, proving invaluable for studying fundamental processes, such as cell cycle regulation, which is crucial for cancer research. Additionally, this unicellular fungus is widely used in the food and beverage industry.^[74]^[75]^[76]^[77]^[78]	North America, Japan, Europe
1996	Danio rerio	Mutant screening	A large-scale screen for developmental mutants in Danio rerio (zebrafish) is led by researchers Christiane Nüsslein-Volhard in Tübingen, Germany, and Wolfgang Driever in Boston, USA. This extensive effort results in thousands of mutant lines exhibiting defects in major organ systems and embryo patterning, with the findings published in a special issue of the journal Development, which would remain its largest issue to date. The success of this screen would inspire many research centers to continue exploring mutant genetics. The zebrafish would emerge as a prominent model organism in vertebrate developmental biology due to its small size, low maintenance cost, and the transparency of its early embryos, which facilitates the screening of morphological defects.^[79]^[80]^[81]^[82]	Germany, United States
1997	Escherichia coli	Genome sequencing	The complete genome sequence of Escherichia coli is published, providing significant insights into one of the most extensively studied bacteria in molecular biology. E. coli resides in the lower intestinal tract of animals and is typically harmless, making it an ideal model organism for biochemists and geneticists. By this time, researchers had long used E. coli to investigate fundamental biochemical processes and gene regulation. The sequenced genome, comprising approximately 4.6 million base pairs and around 4,000 genes, enables scientists to deepen their understanding of its biology. Moreover, comparing non-pathogenic strains with pathogenic ones can aid in developing treatments and preventive strategies for illnesses such as food poisoning.^[83]	United States
1998	Caenorhabditis elegans	Genome sequencing	The genome of Caenorhabditis elegans, a small soil-dwelling nematode, is fully sequenced and published in Science, making it the first multicellular organism to have its genome completed. This effort, led by the C. elegans Sequencing Consortium—a collaboration between the Genome Sequencing Center in St. Louis and the Sanger Centre in Hinxton—demonstrates the feasibility of high-throughput sequencing techniques, crucial for the Human Genome Project. With an accuracy of one error per 10,000 bases, the C. elegans genome becomes a valuable resource for gene discovery. Its use as a model organism provides insights into genetic function and developmental biology, further influencing biomedical research.^[84]^[85]^[86]	United States, United Kingdom
2000 (December)	Arabidopsis thaliana	Genome sequencing	The genome of Arabidopsis thaliana, a small flowering plant, is fully sequenced, marking a historic milestone as the first complete genome of a flowering plant, and launching the era of plant genomics. This achievement provides an essential genetic reference that would be freely accessible to scientists, revolutionizing plant science by enabling in-depth studies on plant genetics, growth, and development. The global collaboration involves major institutions such as Stanford Genome Technology Center and Cold Spring Harbor Laboratory. The A. thaliana genome spans around 125 megabases, contains roughly 25,500 genes, and features about 35% unique genes, with evidence of ancient polyploidy in large segmental duplications.^[87]^[88]^[89]	United States
2000	Drosophila melanogaster	Genome sequencing	The genome of the fruit fly Drosophila melanogaster is sequenced in a groundbreaking effort published in the March 24, 2000 issue of Science. This project, a collaboration between Celera Genomics and the Drosophila Genome Projects, marks the first successful application of the whole genome shotgun (WGS) method in a multicellular organism. Researchers sequence approximately 97 to 98 percent of the genome, capturing nearly all of the estimated 13,600 genes. This achievement is significant in genetic research, establishing a precedent for future genome projects. Further improvements and annotations would since be made by the Berkeley Drosophila Genome Project and FlyBase.^[90]^[91]^[92]	United States
2001	Human	Genome sequencing	The International Human Genome Sequencing Consortium publishes a draft sequence of the human genome in Nature, marking a landmark achievement in genomics. This collaborative project involves 16 genome centers worldwide, which coordinated efforts through regular meetings and phone conferences. Following a policy of rapid data release, the consortium made the assembled genome sequence publicly available within 24 hours. The draft reveals key insights: the human genome spans 2.85 billion bases, with DNA sequences that are 99.9% identical between individuals. It includes approximately 22,300 protein-coding genes, numerous segmental duplications, and over 3 million single nucleotide polymorphisms (SNPs). The Human Genome Project lays a crucial foundation for biomedical research, enabling advancements in disease research and personalized medicine.^[93]^[94]^[95]	United States
2002		Genome sequencing	The Mouse Genome Sequencing Consortium—including the Broad Institute, Washington University, and the Sanger Institute—publish a draft of the mouse genome, using the C57BL/6J strain, the most widely studied inbred strain. This draft revealed around 24,500 protein-coding genes across 19 autosomal pairs, plus X and Y sex chromosomes. The mouse genome, about 14% shorter than the human genome, aligns with large human chromosomal segments, and 75% of mouse genes have direct human counterparts. The similarities between mouse and human genomes make the mouse an essential model for studying human disease, with its short lifecycle and rapid breeding facilitating genetic studies on a large scale.^[96]^[97]^[98]	United States
2003	Neurospora crassa	Genome sequencing	The complete genome sequence of Neurospora crassa is reported, revealing a genome size of 43 million base pairs that encodes approximately 10,000 genes organized into seven linkage groups. The sequencing is conducted using whole genome shotgun sequencing and paired-end sequencing methods. Notably, N. crassa features a G + C content of about 50% and exhibits minimal repetitive DNA, making it a valuable model organism for studying genetics, gene regulation, and metabolic processes. This comprehensive genomic information enhances the understanding of fungal biology and facilitates comparative studies across various organisms.^[99]	United States
2003	Nothobranchius furzeri	Model introduction	Nothobranchius furzeri (Turquoise killifish) is introduced as model organism for aging and longevity research due to its short lifespan and vulnerability to age-related diseases. With a maximum survival time of under 12 weeks in laboratory conditions, N. furzeri exhibits a typical age-dependent increase in mortality. This species' short lifespan, small size, and ability to be propagated in captivity make it ideal for studying the biology of aging. Researchers, including Stefano Valdesalici and Alessandro Cellerino, highlighted its potential in evolutionary studies related to senescence and the effects of natural selection on lifespan.^[100]
2004		Literature	Pamela M. Carroll and Kevin Fitzgerald publish Model Organisms in Drug Discovery, which explores the use of model organisms like Drosophila (fruit flies), C. elegans worms, yeast, mice, and zebrafish in drug discovery and biomedical research. These organisms share many biological pathways with humans, making them invaluable for studying diseases and testing therapeutic compounds. The book highlights how advances in bioinformatics, proteomics, and automation technologies have enhanced the use of these models on an industrial scale. Leading experts discuss real-life applications of model organisms in therapeutic areas, their role in drug discovery, and their potential future impact.^[101]	United States
2007	Chlamydomonas reinhardtii	Genome sequencing	The complete nuclear genome sequence of Chlamydomonas reinhardtii is published. This organism diverged from land plants over 1 billion years ago and serves as a model for studying chloroplast-based photosynthesis and the structure and function of eukaryotic flagella, which are absent in land plants. The ∼120-megabase genome is sequenced, and comparative phylogenomic analyses identify genes encoding uncharacterized proteins likely related to chloroplast and flagellar function. This research enhances our understanding of the ancestral eukaryotic cell, uncovers novel genes associated with photosynthesis and flagellar functions, and links ciliopathy with flagellar composition and function.^[102]
2009		Literature	David A. Crotty and Alexander Gann publish Emerging Model Organisms: A Laboratory Manual, which explores the growing diversity of model organisms used in molecular, cellular, and developmental biology. With advances in gene expression technology, lower genome sequencing costs, and a rising interest in evolutionary biology, by this time researchers expand beyond traditional models. The book introduces 23 emerging model organisms, such as bats, butterflies, snails, and tomatoes, each offering unique research opportunities.^[103]
2019		Literature	Daiana S. Avila publishes Caenorhabditis Elegans: An Overview and Emerging Roles in Studying Disease, which examines the use of the nematode C. elegans as a model organism for studying various diseases. By this time, this nonpathogenic organism had been instrumental in uncovering genetic mechanisms linked to cancer, aging, and nervous system development. The book covers disease models in C. elegans for aging, metabolic syndrome, neurodegenerative diseases, and chemically induced neurodegeneration. It reviews the creation of transgenic animals mimicking human diseases, advances in molecular understanding, and potential treatments, highlighting both the strengths and limitations of these models.^[104]

Numerical and visual data

Google trends

The image below shows Google Trends data for model organism (tipic) from January 2004 to October 2024, as well as interest by region.^[105]

Google Ngram Viewer

The chart below shows Google Ngram Viewer data for "model organism" from 1800 to 2022.^[106]

Wikipedia views

The chart below shows pageviews of the English Wikipedia article Model organism, from July 2015 to October 2024, when the screenshot was taken.^[107]

Meta information on the timeline

How the timeline was built

The initial version of the timeline was written by Sebastian.

Funding information for this timeline is available.

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What the timeline is still missing

Books
Recognition. Nobel
Species discovery dates
Davis, R. H. (2004). The age of model organisms. Nature Reviews Genetics, 5(1), 69–76. doi:10.1038/nrg1250
A column for kingdom
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Model organism
History of model organisms
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Timeline update strategy

External links

References

↑ "Andreas Vesalius (1514–1564)". The Embryo Project Encyclopedia. Arizona State University. Retrieved 19 October 2024.
↑ ^2.0 ^2.1 Bruno Müller, Ueli Grossniklaus (2010). "Model organisms--A historical perspective". Journal of Proteomics. 73 (12): 2182–2183. PMID 20727995. doi:10.1016/j.jprot.2010.08.002.
↑ Laura D. Mackey, D. Allan Smith, Richard D. Wood. "Probing the Mechanism of Nucleotide Excision Repair with Xenopus Egg Extracts". Nature Protocols. pp. 846–854. doi:10.1038/nprot.2016.049. Retrieved 20 October 2024.
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[1] "Andreas Vesalius (1514–1564)". The Embryo Project Encyclopedia. Arizona State University. Retrieved 19 October 2024.

[historical-2] 2.0 ^2.1 Bruno Müller, Ueli Grossniklaus (2010). "Model organisms--A historical perspective". Journal of Proteomics. 73 (12): 2182–2183. PMID 20727995. doi:10.1016/j.jprot.2010.08.002.

[3] Laura D. Mackey, D. Allan Smith, Richard D. Wood. "Probing the Mechanism of Nucleotide Excision Repair with Xenopus Egg Extracts". Nature Protocols. pp. 846–854. doi:10.1038/nprot.2016.049. Retrieved 20 October 2024.

[4] Severin Sasso, Herwig Stibor, Maria Mittag, Arthur R Grossman (2018). "From molecular manipulation of domesticated Chlamydomonas reinhardtii to survival in nature". eLife. 7: e39233. doi:10.7554/eLife.39233. |access-date= requires |url= (help)

[5] "Gregor Mendel: The father of genetics who opened a biological world full of wonders" (PDF). Cell.com. Retrieved 24 October 2024.

[6] "Gregor Mendel's Pea Plant Experiment" (PDF). Michigan.gov. Retrieved 24 October 2024.

[7] "Mendel and his peas". LibreTexts. Retrieved 24 October 2024.

[8] "Gregor Mendel: Genetics & Experiments". Study.com. Retrieved 24 October 2024.

[9] "An Introduction to Saccharomyces cerevisiae". JoVE. Retrieved 20 October 2024.

[10] "Compensatory evolution reveals ecological constraints on morphological evolution". eLife. doi:10.7554/eLife.05835. Retrieved 20 October 2024.

[11] Rheinberger, H. J. (December 2000). "Mendelian inheritance in Germany between 1900 and 1910. The case of Carl Correns (1864-1933)". Comptes rendus de l'Academie des sciences. Serie III, Sciences de la vie. 323 (12): 1089–1096. ISSN 0764-4469. doi:10.1016/s0764-4469(00)01267-1.

[12] Phifer-Rixey, Megan; Nachman, Michael W (15 April 2015). "Insights into mammalian biology from the wild house mouse Mus musculus". eLife. 4. doi:10.7554/eLife.05959.

[13] "Chapter 1 - The Laboratory Mouse". www.informatics.jax.org. Retrieved 2 June 2024.

[14] "The Nobel Prize in Physiology or Medicine 1933". NobelPrize.org. Retrieved 2 June 2024.

[15] "Google Scholar". scholar.google.com. Retrieved 2 June 2024.

[16] Hugo J Bellen, Shinya Yamamoto (24 September 2015). "Morgan's Legacy: Fruit Flies and the Functional Annotation of Conserved Genes". Cell. 163 (1): 12–14. PMC PMC4783153 Check |pmc= value (help). PMID 26406362. doi:10.1016/j.cell.2015.09.009. Retrieved 24 October 2024.

[17] Christine M. Roche, Jennifer J. Loros, Kevin McCluskey, N. Louise Glass (2014). "Neurospora crassa: Looking Back and Looking Forward at a Model Microbe". American Journal of Botany. 101 (12): 2022–2035. doi:10.3732/ajb.1400377.

[18] Shirley B. R. R. A. (2000). "A History of Research on the Fungal Genus Neurospora". Fungal Genetics Reports. Retrieved 25 October 2024.

[19] Rowland H Davis (March 2007). "Tending Neurospora: David Perkins, 1919–2007, and Dorothy Newmeyer Perkins, 1922–2007". Mycologia. 99 (2): 171–175. PMC PMC1855115 Check |pmc= value (help). PMID 17449866. Retrieved 25 October 2024.

[20] Patrice A Salomé; Sabeeha S Merchant (19 June 2019). "A Series of Fortunate Events: Introducing Chlamydomonas as a Reference Organism". Plant Cell. 31 (8): 1682–1707. doi:10.1105/tpc.18.00952. Retrieved 16 October 2024.

[21] "Full Catalog of Titles from Cold Spring Harbor Laboratory Press". cshlpress.com. Retrieved 16 October 2024.

[22] "Elucidation of the life cycle of Saccharomyces cerevisiae yeast". winehistory.com.au. Retrieved 16 October 2024.

[23] Helmut Plattner (2022). "Ciliate research: From myth to trendsetting science". Journal of Eukaryotic Microbiology. Wiley Periodicals LLC. doi:10.1111/jeu.12926. Retrieved 16 October 2024.

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[25] Helmut Plattner (2022). "Ciliate research: From myth to trendsetting science". Journal of Eukaryotic Microbiology. doi:10.1111/jeu.12926. Retrieved 16 October 2024.

[26] Marian Dominguez-Mirazo, Jeremy D. Harris, David Demory, Joshua S. Weitz. "Accounting for cellular-level variation in lysis: implications for virus–host dynamics". MBio. doi:https://doi.org/10.1128/mbio.01376-24 Check |doi= value (help). Retrieved 17 October 2024.

[27] Sherin Kannoly, Gabriella Oken, Jonathan Shadan, David Musheyev, Kevin Singh, Abhyudai Singh, John J Dennehy (2022-12-21). "Single-Cell Approach Reveals Intercellular Heterogeneity in Phage-Producing Capacities". Microbiol Spectr. 11 (1): e02663-21 pmid=36541779. doi:10.1128/spectrum.02663-21. Retrieved 17 October 2024.

[28] Alan J Cann (2015-07-24). "Replication". Principles of Molecular Virology: 105–133. doi:10.1016/B978-0-12-801946-7.00004-3. Retrieved 17 October 2024.

[29] "Beadle and Tatum's 1941 Experiments with Neurospora Revealed that Genes Produce Enzymes". embryo.asu.edu. Retrieved 17 October 2024.

[30] Charles Yanofsky (2005). "The Favorable Features of Tryptophan Synthase for Proving Beadle and Tatum's One Gene–One Enzyme Hypothesis". Genetics. 169 (2): 511–516. PMID 15731515. doi:10.1093/genetics/169.2.511. Retrieved 17 October 2024.

[31] Vincenzo E. A. Russo, Niketan N. Pandit. "Development". Development in Neurospora crassa. Springer. pp. 88–102. Retrieved 17 October 2024.

[32] George Haughn, Ljerka Kunst. "Arabidopsis Thaliana, a Model Organism for Molecular Genetic Studies in Plants: How and Why Was Arabidopsis Chosen Over Other Plants?" (PDF). blogs.ubc.ca. University of British Columbia. Retrieved 17 October 2024.

[33] Lindegren C.C., Lindegren G. (1943). "A New method for hybridizing yeast". Proceedings of the National Academy of Sciences USA. 29: 306. doi:10.1073/pnas.29.10.306. |access-date= requires |url= (help)

[34] Szymanski E., Vermeulen N., Wong M. (2019). "Yeast: One cell, one reference sequence, many genomes?". New Genetics and Society. 38: 430–450. doi:10.1080/14636778.2019.1677150. |access-date= requires |url= (help)

[35] Thomas A Dixon, Isak S Pretorius (2020-09-28). "Drawing on the Past to Shape the Future of Synthetic Yeast Research". International Journal of Molecular Sciences. 21 (19): 7156. doi:10.3390/ijms21197156. Retrieved 17 October 2024.

[36] "History: The Phage Group". Cold Spring Harbor Laboratory. Archived from the original on 2007-05-17. Retrieved 2007-05-04.

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@@ Line 1: / Line 1: @@
-This is a '''timeline of FIXME'''.
+This is a '''{{w|model organism}}s''', which are non-human species extensively studied in biological research to gain insights into various processes that can be applicable to other organisms, including humans. These organisms are selected for their ease of maintenance, rapid life cycles, and well-characterized genetics. They play a crucial role in fields such as {{w|genetics}}, {{w|developmental biology}}, and {{w|neuroscience}}.
 == Sample questions ==
@@ Line 9: / Line 9: @@
 {| class="wikitable"
 ! Time period !! Development summary !! More details
+|-
+| Ancient Times to Early 19th Century || Pre-Modern Era || During this period, early naturalists and philosophers such as Aristotle observe animals and plants, focusing primarily on descriptive biology. Humans and domestic animals are studied for agricultural, medical, and practical purposes, with limited experimentation. Comparative anatomy begins to emerge with scientists like {{w|Andreas Vesalius}}, who uses human cadavers and animals for anatomical studies, though the approach is largely observational and lacks the rigor of controlled experiments.<ref>{{cite web |title=Andreas Vesalius (1514–1564) |url=https://embryo.asu.edu/pages/andreas-vesalius-1514-1564 |website=The Embryo Project Encyclopedia |publisher=Arizona State University |accessdate=19 October 2024}}</ref>
+|-
+| Mid-19th Century to Early 20th Century || Early Experimental Biology || The mid-19th century marks a turning point with the advent of systematic biological experimentation. Gregor Mendel’s experiments with pea plants in the 1850s uncover the basic laws of inheritance, forming the foundation of genetics. By the early 20th century, Thomas Hunt Morgan’s introduction of ''{{w|Drosophila melanogaster}}'' in genetics research help establish the chromosomal theory of inheritance, making the fruit fly a pioneering model organism. This period also sees early embryological studies using frogs and chickens, as researchers begin to explore developmental biology in a more structured way.<ref name="historical"/>
+|-
+| Mid-20th Century to Late 20th Century || The Rise of Molecular Biology || The mid-20th century brings revolutionary advancements in molecular biology, with model organisms playing central roles. Bacteria like ''{{w|Escherichia coli}}'' becomes vital for studying DNA replication, gene expression, and molecular genetics, while ''{{w|Caenorhabditis elegans}}'' provides insights into developmental biology and programmed cell death. Mice emerged as key models for human disease research, especially in cancer, immunology, and developmental studies. This era is characterized by the discovery of the DNA double helix, the rise of recombinant DNA technology in the 1970s, and the first successful gene cloning experiments, all of which rely on these model organisms.
+|-
+| Late 20th Century to Present ||  Genomics and Systems Biology Era || With the late 20th century comes the sequencing of genomes, including those of model organisms like Drosophila, C. elegans, and mice, ushering in the genomics era. New technologies like CRISPR-Cas9 revolutionize gene editing, allowing scientists to modify genes with precision in organisms like zebrafish and mice. This period sees the rise of systems biology, integrating data from various biological scales to understand complex processes such as disease mechanisms, development, and aging. Model organisms remain indispensable in fields like synthetic biology, drug discovery, and personalized medicine, as researchers now leverage the power of genomics to tackle complex biological questions on a large scale.
 |-
 |}
@@ Line 15: / Line 23: @@
 {| class="sortable wikitable"
-! Year !! Kingdom !! Event type !! Details !! Location/researcher affiliation
+! Year !! Species !! Event type !! Details !! Location/researcher affiliation
+|-
+| 1822 || ''Danio rerio'' || Discovery || ''Danio rerio'', initially designated as ''Brachydanio rerio'', is first described by English physician Sir Francis Hamilton in his book on the fishes of the {{w|Ganges River}}. Hamilton identifies ten species at that time, (the Danio genus now comprises 45 known species). ''D. rerio'' is a monophyletic species within the ''Cyprinidae'' family, characterized by a bilobate caudal fin.<ref>{{cite web |title=Probing the Mechanism of Nucleotide Excision Repair with Xenopus Egg Extracts |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC4937470/ |journal=Nature Protocols |volume=11 |issue=5 |pages=846–854 |doi=10.1038/nprot.2016.049 |author=Laura D. Mackey, D. Allan Smith, Richard D. Wood |accessdate=20 October 2024}}</ref> || {{w|United Kingdom}}
 |-
-| 1900 || || || German botanist {{w|Carl Correns}} conducts experiments on ''{{w|Zea mays}}'', commonly known as corn or maize. Correns confirms the findings of {{w|Gregor Mendel}}, an Austrian monk, regarding the principles of inheritance and genetic traits. Mendel's work, initially published in 1866, outlines the laws of inheritance based on his experiments with pea plants. Correns' validation of Mendel's findings with ''Zea mays'' provided further evidence for the existence of discrete units of inheritance, which we now know as genes. This confirmation plays a crucial role in the establishment of modern genetics and lays the foundation for understanding heredity in plants and animals.<ref>{{cite journal |last1=Rheinberger |first1=H. J. |title=Mendelian inheritance in Germany between 1900 and 1910. The case of Carl Correns (1864-1933) |journal=Comptes rendus de l'Academie des sciences. Serie III, Sciences de la vie |date=December 2000 |volume=323 |issue=12 |pages=1089–1096 |doi=10.1016/s0764-4469(00)01267-1 |url=https://pubmed.ncbi.nlm.nih.gov/11147095/ |issn=0764-4469}}</ref> || {{w|Germany}}
+| 1833 || ''Chlamydomonas reinhardtii'' || Discovery || German scientist {{w|Christian Gottfried Ehrenberg}} describes the genus ''Chlamydomonas'', which includes the species ''Chlamydomonas reinhardtii'', a single-celled green alga found in temperate soil habitats. By the mid-20th century, C. reinhardtii would become a vital model organism for investigating fundamental cellular processes, such as photosynthesis, light perception, and the structure, function, and biogenesis of cilia. Research on this alga would notably advance plant and cell biology, contributing to algal biotechnology, and enhancing our understanding of human diseases.<ref>{{cite journal |title=From molecular manipulation of domesticated Chlamydomonas reinhardtii to survival in nature |author=Severin Sasso, Herwig Stibor, Maria Mittag, Arthur R Grossman |journal=eLife |volume=7 |pages=e39233 |year=2018 |doi=10.7554/eLife.39233|accessdate=20 October 2024}}</ref> || {{w|Germany}}
 |-
-| 1902 || || || American biologist {{w|William Ernest Castle}} begins genetic studies on ''{{w|Mus musculus}}'', commonly known as the house mouse. This marks the initiation of systematic genetic research on this species. Castle's work contributes to the understanding of inheritance patterns and genetic variation in mice, laying the groundwork for further investigations into the genetic basis of traits and the mechanisms of heredity. His studies on ''Mus musculus'' were instrumental in the development of mouse models for genetic research, which continue to be crucial in biomedical research and the study of human genetics.<ref>{{cite journal |last1=Phifer-Rixey |first1=Megan |last2=Nachman |first2=Michael W |title=Insights into mammalian biology from the wild house mouse Mus musculus |journal=eLife |date=15 April 2015 |volume=4 |doi=10.7554/eLife.05959 |url=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4397906/}}</ref><ref>{{cite web |title=Chapter 1 - The Laboratory Mouse |url=https://www.informatics.jax.org/greenbook/chapters/chapter1.shtml |website=www.informatics.jax.org |access-date=2 June 2024}}</ref> || {{w|United States}}
+| 1853 || ''Pisum sativum'' || Experiment || {{w|Gregor Mendel}} begins his groundbreaking experiments with pea plants, which lays the foundation for classical genetics. He focuses on easily observable traits like plant height, seed color, and flower color. Mendel employs controlled breeding, cross-pollinating plants with distinct traits to study inheritance patterns. Unlike earlier researchers, he meticulously counted and analyzed his results, identifying consistent patterns of trait transmission. His findings led to the formulation of key genetic principles, the "Law of Segregation" and the "Law of Independent Assortment," which explain how traits are passed from parents to offspring.<ref>{{cite web |title=Gregor Mendel: The father of genetics who opened a biological world full of wonders |url=https://www.cell.com/molecular-plant/pdf/S1674-2052(22)00365-3.pdf |website=Cell.com |accessdate=24 October 2024}}</ref><ref>{{cite web |title=Gregor Mendel’s Pea Plant Experiment |url=https://www.michigan.gov/explorelabscience/-/media/Project/Websites/explorelabscience/pdf/Lab-Teens-pdf/Gregor_Mendel_Pea_Experiment.pdf |website=Michigan.gov |accessdate=24 October 2024}}</ref><ref>{{cite web |title=Mendel and his peas |url=https://bio.libretexts.org/Bookshelves/Genetics/Classical_Genetics_(Khan_Academy)/01%3A_Introduction_to_heredity/1.04%3A_Mendel_and_his_peas |website=LibreTexts |accessdate=24 October 2024}}</ref><ref>{{cite web |title=Gregor Mendel: Genetics & Experiments |url=https://study.com/academy/lesson/gregor-mendel-genetics-experiments-laws-discovery.html |website=Study.com |accessdate=24 October 2024}}</ref> || {{w|Czech Republic}} ({{w|Austrian Empire}})
 |-
-| 1909 || || || Thomas Hunt Morgan begins his groundbreaking work with the fruit fly ''{{w|Drosophila melanogaster}}'', which would become synonymous with his name. Prior to this, C. W. Woodworth and W. E. Castle had shown interest in Drosophila for genetic studies. Morgan's research with Drosophila would lead to the discovery of sex linkage of the gene for white eyes, demonstrating the phenomenon of linkage. He bred Drosophila in large quantities, facilitating the analysis of spontaneous mutations and the localization of genes. Morgan's work laid the foundation for understanding the linear arrangement of genes in chromosomes and significantly advanced the field of genetics.<ref>{{cite web |title=The Nobel Prize in Physiology or Medicine 1933 |url=https://www.nobelprize.org/prizes/medicine/1933/morgan/biographical/ |website=NobelPrize.org |access-date=2 June 2024}}</ref> ||
+| 1857 || ''Saccharomyces cerevisiae'' || Discovery || {{w|Louis Pasteur}} discovers ''Saccharomyces cerevisiae'', identifying it as the key microbe in winemaking and bread baking. He classifies this yeast as a facultative anaerobe, meaning it can switch to fermentation in the absence of oxygen. This discovery would not only revolutionize food and beverage production but also establishes ''S. cerevisiae'' as a model organism in scientific research. Its simple eukaryotic structure and genetic tractability would make it a powerful tool for studying fundamental biological processes, including genetics, cell biology, and biochemistry, further enhancing our understanding of more complex organisms.<ref>{{cite web |title=An Introduction to Saccharomyces cerevisiae |url=https://app.jove.com/v/5081/an-introduction-to-saccharomyces-cerevisiae |website=JoVE |accessdate=20 October 2024}}</ref><ref>{{cite web |title=Compensatory evolution reveals ecological constraints on morphological evolution |url=https://elifesciences.org/articles/05835 |website=eLife |accessdate=20 October 2024 |doi=10.7554/eLife.05835}}</ref> || {{w|France}}
 |-
-| 1913 || || || ''Z. mays'': Emerson and East publish important quantitative genetics paper. ||
+| 1900 || ''Zea mays'' || Genetic validation || German botanist {{w|Carl Correns}} conducts experiments on ''{{w|Zea mays}}'', commonly known as corn or maize. Correns confirms the findings of {{w|Gregor Mendel}}, an Austrian monk, regarding the principles of inheritance and genetic traits. Mendel's work, initially published in 1866, outlines the laws of inheritance based on his experiments with pea plants. Correns' validation of Mendel's findings with ''Zea mays'' provides further evidence for the existence of discrete units of inheritance, which would be later known as genes. This confirmation plays a crucial role in the establishment of modern genetics and lays the foundation for understanding heredity in plants and animals.<ref>{{cite journal |last1=Rheinberger |first1=H. J. |title=Mendelian inheritance in Germany between 1900 and 1910. The case of Carl Correns (1864-1933) |journal=Comptes rendus de l'Academie des sciences. Serie III, Sciences de la vie |date=December 2000 |volume=323 |issue=12 |pages=1089–1096 |doi=10.1016/s0764-4469(00)01267-1 |url=https://pubmed.ncbi.nlm.nih.gov/11147095/ |issn=0764-4469}}</ref> || {{w|Germany}}
 |-
-| 1915 || || || D. melanogaster: First book on Mendelian genetics is published from the Morgan Group
+| 1902 || ''Mus musculus'' || Genetic research || American biologist {{w|William Ernest Castle}} begins genetic studies on ''{{w|Mus musculus}}'', commonly known as the house mouse. This marks the initiation of systematic genetic research on this species. Castle's work contributes to the understanding of inheritance patterns and genetic variation in mice, laying the groundwork for further investigations into the genetic basis of traits and the mechanisms of heredity. His studies on ''Mus musculus'' were instrumental in the development of mouse models for genetic research, which continue to be crucial in biomedical research and the study of human genetics.<ref>{{cite journal |last1=Phifer-Rixey |first1=Megan |last2=Nachman |first2=Michael W |title=Insights into mammalian biology from the wild house mouse Mus musculus |journal=eLife |date=15 April 2015 |volume=4 |doi=10.7554/eLife.05959 |url=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4397906/}}</ref><ref>{{cite web |title=Chapter 1 - The Laboratory Mouse |url=https://www.informatics.jax.org/greenbook/chapters/chapter1.shtml |website=www.informatics.jax.org |access-date=2 June 2024}}</ref> || {{w|United States}}
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-| 1927 || || || Neurospora crassa: Shear and Dodge discover sexual cycle and describe mating types
+| 1909 || ''Drosophila melanogaster'' || Gene mapping || American scientist {{w|Thomas Hunt Morgan}} begins his groundbreaking work with the fruit fly ''{{w|Drosophila melanogaster}}'', which would become synonymous with his name. Prior to this, C. W. Woodworth and W. E. Castle had shown interest in Drosophila for genetic studies. Morgan's research with Drosophila would lead to the discovery of sex linkage of the gene for white eyes, demonstrating the phenomenon of linkage. He bred Drosophila in large quantities, facilitating the analysis of spontaneous mutations and the localization of genes. Morgan's work laid the foundation for understanding the linear arrangement of genes in chromosomes and significantly advanced the field of genetics.<ref>{{cite web |title=The Nobel Prize in Physiology or Medicine 1933 |url=https://www.nobelprize.org/prizes/medicine/1933/morgan/biographical/ |website=NobelPrize.org |access-date=2 June 2024}}</ref> || {{w|United States}} ({{w|Columbia University}})
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-| 1930 || || || ''Chlamydomonas reinhardtii'': Moewus develops genetic system. ||
+| 1913 || ''Zea mays'' || Research || Edgar Nelson Emerson and Roland McMillan East publish a significant paper on quantitative genetics in ''{{w|Zea mays}}''. This paper marks an important milestone in the understanding of genetic principles governing quantitative traits, which are traits controlled by multiple genes and influenced by environmental factors. Emerson and East's work would contribute to the development of quantitative genetics as a field by elucidating the complex inheritance patterns of traits such as height, yield, and other quantitative characteristics in maize. Their research lays the foundation for further studies in the genetics of complex traits in various organisms.<ref>{{cite web |title=Google Scholar |url=https://scholar.google.com/scholar_lookup?&title=The%20inheritance%20of%20quantitative%20characters%20in%20maize&pages=1-120&publication_year=1913&author=Emerson%2CRA&author=East%2CEM |website=scholar.google.com |access-date=2 June 2024}}</ref> || {{w|United States}}
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-| 1935 || || || ''Saccharomyces cerevisiae'': Winge describes haplo- and diplophase of life cycle. ||
+| 1915 || ''{{w|Drosophila melanogaster}}'' || Publication || The Morgan Group, led by Thomas Hunt Morgan, publishes the first book on Mendelian genetics focusing on ''{{w|Drosophila melanogaster}}'', commonly known as the fruit fly. This publication represents a significant milestone in the field of genetics, as it provides a comprehensive overview of the principles of Mendelian inheritance as observed in ''Drosophila''. The book likely covers topics such as the inheritance of traits, the mapping of genes, and the understanding of genetic linkage. This work serves as a foundational resource for researchers studying genetics and paves the way for further investigations into the mechanisms of inheritance in various organisms.<ref>{{cite journal |title=Morgan’s Legacy: Fruit Flies and the Functional Annotation of Conserved Genes |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC4783153/ |author=Hugo J Bellen, Shinya Yamamoto |journal=Cell |volume=163 |issue=1 |pages=12–14 |date=24 September 2015 |doi=10.1016/j.cell.2015.09.009 |pmid=26406362 |pmc=PMC4783153 |accessdate=24 October 2024}}</ref> || {{w|United States}}
 |-
-| 1937 || || || Paramecium spp.: Sonneborn and Jennings domesticate crosses and define mating types. ||
+| 1927 || ''{{w|Neurospora crassa}}'' || Discovery || Researchers Shear and Dodge make a notable discovery regarding ''Neurospora crassa'', a type of bread mold. They identify and describe the sexual cycle of ''Neurospora crassa'', shedding light on its reproductive mechanisms. Additionally, they characterize different mating types within the species, which are essential for sexual reproduction. This finding is significant in advancing the understanding of fungal genetics and reproductive biology. It lays the groundwork for further studies on the genetics and life cycle of Neurospora crassa, making it an essential model organism in genetic research.<ref>{{cite journal |title=Neurospora crassa: Looking Back and Looking Forward at a Model Microbe |url=https://bsapubs.onlinelibrary.wiley.com/doi/pdf/10.3732/ajb.1400377 |author=Christine M. Roche, Jennifer J. Loros, Kevin McCluskey, N. Louise Glass |journal=American Journal of Botany |volume=101 |issue=12 |pages=2022–2035 |year=2014 |doi=10.3732/ajb.1400377}}</ref><ref>{{cite web |title=A History of Research on the Fungal Genus Neurospora |url=https://newprairiepress.org/cgi/viewcontent.cgi?article=1121&context=fgr |author=Shirley B. R. R. A. |journal=Fungal Genetics Reports |volume=41 |issue=1 |date=2000 |accessdate=25 October 2024}}</ref><ref>{{cite journal |title=Tending Neurospora: David Perkins, 1919–2007, and Dorothy Newmeyer Perkins, 1922–2007 |url=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1855115/ |author=Rowland H Davis |journal=Mycologia |volume=99 |issue=2 |pages=171–175 |date=March 2007 |pmid=17449866 |pmc=PMC1855115 |accessdate=25 October 2024}}</ref> || {{w|United States}}
 |-
-| 1939 || || || T phages: Ellis and Delbrück describe replication cycle, ‘one-step growth. ||
+| 1930 || ''{{w|Chlamydomonas}}'' || Genetics research || German biologist {{w|Franz Moewus}} pioneer biochemical genetics research on ''{{w|Chlamydomonas}}'', focusing on ''{{w|Chlamydomonas eugametos}}''. He claims to have identified carotenoid-related hormones that selectively activate male or female gametes and reports isolating and genetically mapping mutants along their biosynthetic pathways. Over a decade, he would analyze 200,000 zygotes for ten phenotypes, though many of his findings would be later deemed irreproducible, raising doubts about his conclusions. Despite these issues, Moewus significantly influences ''Chlamydomonas'' research, inspiring further studies and reinforcing concepts like gene-chromosome localization and the one gene-one enzyme hypothesis, which later led to the use of *C. reinhardtii* as a model organism.<ref>{{cite journal |title=A Series of Fortunate Events: Introducing Chlamydomonas as a Reference Organism |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC6713297/ |journal=Plant Cell |date=19 June 2019 |volume=31 |issue=8 |pages=1682–1707 |doi=10.1105/tpc.18.00952 |author1=Patrice A Salomé |author2=Sabeeha S Merchant |accessdate=16 October 2024}}</ref> || {{w|Germany}}
 |-
-| 1941 || || || N. crassa: Beadle and Tatum isolate first biochemical mutants. ||
+| 1935 || ''{{w|Saccharomyces cerevisiae}}'' || Foundational research ||  Danish biologist {{w|Øjvind Winge}}, later regarded as the "Father of Yeast Genetics," elucidates the life cycle of ''{{w|Saccharomyces cerevisiae}}'', marking the beginning of yeast genetics research. Winge’s work in Denmark laid the foundation for subsequent studies in yeast genetics, which were later pursued by researchers such as Lindegren in the U.S. and Ephrussi in France. Meanwhile, Leupold pioneered genetic studies on *Schizosaccharomyces pombe* in Switzerland during the 1940s. Over the next four decades, yeast became an essential model organism in eukaryotic molecular biology, with early researchers contributing significantly to the field's growth and recognition.<ref>{{cite web |title=Full Catalog of Titles from Cold Spring Harbor Laboratory Press |url=https://cshlpress.com/default.tpl?cart=1729204217784511883&fromlink=T&linkaction=full&linksortby=oop_title&--eqSKUdatarq=676 |website=cshlpress.com |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Elucidation of the life cycle of Saccharomyces cerevisiae yeast |url=https://winehistory.com.au/wiki/Wine_History/Elucidation_of_the_life_cycle_of_Saccharomyces_cerevisiae_yeast |website=winehistory.com.au |accessdate=16 October 2024}}</ref> ||
 |-
-| 1943 || || || ''Arabidopsis thaliana'': Laibach initiates program in genetics and development. ||
+| 1935 || || Genetic research || When Emil Winge begins conducting genetic research with yeast, advancements in genetics using other model organisms pave the way for his experiments. Key developments include the understanding of four-strand crossing over, which improves insights into genetic recombination; chromosome mapping, which facilitates the identification of gene locations; the discovery of lethal genes, which helps researchers understand gene functions and interactions; and the transformation of ''Pneumococcus'', which demonstrates the principles of genetic transfer and transformation. These foundational discoveries provide a practical framework for genetic studies in yeast, ultimately contributing to the field of genetics.<ref name="historical">{{cite journal |title=Model organisms--A historical perspective |url=https://pubmed.ncbi.nlm.nih.gov/20727995/ |journal=Journal of Proteomics |year=2010 |volume=73 |issue=12 |pages=2182-2183 |author=Bruno Müller, Ueli Grossniklaus |PMID=20727995 |DOI=10.1016/j.jprot.2010.08.002}}</ref>
 |-
-| 1943 || || || ''S.cerevisiae'': Lindegren begins genetics with heterothallic strains. ||
+| 1937 || ''{{w|Paramecium}}'' || Reproduction || American biologist {{w|Tracy Sonneborn}} and H.S. Jennings make a significant breakthrough in ''{{w|Paramecium}}'' genetics by successfully establishing the conditions necessary for mating. They also define a system of mating types, with most ''Paramecium'' species exhibiting two distinct mating types. This discovery marks a key advancement in the understanding of reproduction and genetics in ''Paramecium'' species, contributing to its establishment as a model organism in cellular and genetic research. Their work lay the groundwork for future studies on sexual reproduction and genetic exchange in single-celled organisms.<ref>{{cite journal |title=Ciliate research: From myth to trendsetting science |author=Helmut Plattner |journal=Journal of Eukaryotic Microbiology |year=2022 |doi=10.1111/jeu.12926 |url=https://onlinelibrary.wiley.com/doi/full/10.1111/jeu.12926 |publisher=Wiley Periodicals LLC |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Ciliate research: From myth to trendsetting science |author=Helmut Plattner |journal=Journal of Eukaryotic Microbiology |year=2022 |doi=10.1111/jeu.12926 |url=https://onlinelibrary.wiley.com/doi/full/10.1111/jeu.12926 |publisher=Wiley Periodicals LLC |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Ciliate research: From myth to trendsetting science |author=Helmut Plattner |journal=Journal of Eukaryotic Microbiology |year=2022 |doi=10.1111/jeu.12926 |url=https://onlinelibrary.wiley.com/doi/full/10.1111/jeu.12926 |accessdate=16 October 2024}}</ref> || {{w|United States}}
 |-
-| 1944 || || || T phages: Delbrück initiates Phage Group. ||
+| 1939 || T2 bacteriophage || One-step growth experiment || American biochemist {{w|Emory Ellis}} and German–American biophysicist {{w|Max Delbrück}} conduct a pivotal one-step growth experiment using T2 bacteriophages to investigate viral replication. They mix bacteria with sufficient viruses to ensure each host cell had a virus attached, then remove the free viruses and sample at regular intervals for a plaque assay. Initially, the number of plaques remain constant during the latent period, followed by a progressive increase, indicating that infected cells were lysing and releasing phages. This experiment illustrates the three main phases of virus replication: initiation of infection, genome replication and expression, and release of mature virions. The method would be also used to estimate burst size, although it may not account for variations in lysis timing, potentially biasing viral trait estimates.<ref>{{cite journal |title=Accounting for cellular-level variation in lysis: implications for virus–host dynamics |author=Marian Dominguez-Mirazo, Jeremy D. Harris, David Demory, Joshua S. Weitz |url=https://journals.asm.org/doi/10.1128/mbio.01376-24 |doi=https://doi.org/10.1128/mbio.01376-24 |journal=MBio |accessdate=17 October 2024}}</ref><ref>{{cite journal |title=Single-Cell Approach Reveals Intercellular Heterogeneity in Phage-Producing Capacities |author=Sherin Kannoly, Gabriella Oken, Jonathan Shadan, David Musheyev, Kevin Singh, Abhyudai Singh, John J Dennehy |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC9927085/ |doi=10.1128/spectrum.02663-21 |journal=Microbiol Spectr |date=2022-12-21 |volume=11 |issue=1 |page=e02663-21 pmid=36541779 |accessdate=17 October 2024}}</ref><ref>{{cite journal |title=Replication |author=Alan J Cann |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC7149640/ |doi=10.1016/B978-0-12-801946-7.00004-3 |journal=Principles of Molecular Virology |date=2015-07-24 |pages=105–133|accessdate=17 October 2024}}</ref> || {{w|United States}}
 |-
-| 1946 || || || ''Escherichia coli'': Lederberg and Tatum discover gene exchange. ||
+| 1941 || ''{{w|Neurospora crassa}}'' || Genetics || American geneticists {{w|George Beadle}} and {{w|Edward Tatum}}'s experiments with ''{{w|Neurospora crassa}}'' lead to the isolation of the first biochemical mutants, providing key insights into gene function. By exposing the fungus to X-rays, they induce mutations and identified strains requiring specific nutrients for growth. This indicates defects in certain metabolic pathways, and their work formulate the "one gene, one enzyme" hypothesis, which proposes that each gene is responsible for producing a single enzyme. This groundbreaking research marks a pivotal step in understanding gene roles in metabolism and would earn them the Nobel Prize in 1958.<ref>{{cite web |title=Beadle and Tatum's 1941 Experiments with Neurospora Revealed that Genes Produce Enzymes |url=https://embryo.asu.edu/pages/beadle-and-tatums-1941-experiments-neurospora-revealed-genes-produce-enzymes |website=embryo.asu.edu |accessdate=17 October 2024}}</ref><ref>{{cite journal |title=The Favorable Features of Tryptophan Synthase for Proving Beadle and Tatum's One Gene–One Enzyme Hypothesis |author=Charles Yanofsky |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC1449131/ |doi=10.1093/genetics/169.2.511 |journal=Genetics |date=2005|volume=169 |issue=2 |pages=511–516 |pmid=15731515 |accessdate=17 October 2024}}</ref><ref>{{cite book |title=Development in Neurospora crassa |author=Vincenzo E. A. Russo, Niketan N. Pandit |chapter=Development |url=https://link.springer.com/chapter/10.1007/978-3-642-77043-2_7 |pages=88–102 |publisher=Springer |accessdate=17 October 2024}}</ref> || {{w|United States}}
 |-
-| 1946 || || || ''S. cerevisiae'': Ephrussi discovers cytoplasmic petite colonie variant. ||
+| 1943 || ''{{w|Arabidopsis thaliana}}'' || Model introduction || German botanist {{w|Friedrich Laibach}} first promotes ''{{w|Arabidopsis thaliana}}'' as a model organism for genetic studies. Despite possessing ideal traits for genetic analysis, such as its small size, short generation time, and simple genome, ''Arabidopsis'' does not gain widespread popularity immediately. Laibach's pioneering efforts to use ''Arabidopsis'' for understanding genetics and development lays the groundwork for its eventual adoption as a model organism, which would later become central to plant molecular biology research.<ref>{{cite web |title=Arabidopsis Thaliana, a Model Organism for Molecular Genetic Studies in Plants: How and Why Was Arabidopsis Chosen Over Other Plants? |author=George Haughn, Ljerka Kunst |url=https://blogs.ubc.ca/haughn/files/2011/05/Essay-Arabidopsis-as-a-model-organism-2010-proof.pdf |website=blogs.ubc.ca |publisher=University of British Columbia |accessdate=17 October 2024}}</ref> || {{w|Germany}}
 |-
-| 1949 || || || ''S. cerevisiae'': Roman begins major US genetic studies. ||
+| 1943 || ''{{w|Saccharomyces cerevisiae}}'' || Genetic mapping || In St. Louis, Missouri, Carl and Gertrude Lindegren become the first to report the existence of two opposite mating types in ''{{w|Saccharomyces cerevisiae}}'': a and α. Their work with heterothallic strains lays the groundwork for understanding the genetic basis of mating in yeast. Subsequently, they publish the first genetic maps of ''S. cerevisiae'' in 1949 and 1951, significantly advancing the field of yeast genetics and establishing ''S. cerevisiae'' as a key model organism for studying eukaryotic genetics and cellular processes.<ref>{{cite journal |title=A New method for hybridizing yeast |author=Lindegren C.C., Lindegren G. |journal=Proceedings of the National Academy of Sciences USA |date=1943 |volume=29 |pages=306 |doi=10.1073/pnas.29.10.306 |pmid= |accessdate=17 October 2024}}</ref><ref>{{cite journal |title=Yeast: One cell, one reference sequence, many genomes? |author=Szymanski E., Vermeulen N., Wong M. |journal=New Genetics and Society |date=2019 |volume=38 |pages=430–450 |doi=10.1080/14636778.2019.1677150 |accessdate=17 October 2024}}</ref><ref>{{cite journal |title=Drawing on the Past to Shape the Future of Synthetic Yeast Research |author=Thomas A Dixon, Isak S Pretorius |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC7583028/ |doi=10.3390/ijms21197156 |journal=International Journal of Molecular Sciences |date=2020-09-28 |volume=21 |issue=19 |page=7156 |accessdate=17 October 2024}}</ref> || {{w|United States}}
 |-
-| 1950 || || || ''C. reinhardtii'': Lewin and Sager begin nuclear and organelle genetic studies. ||
+| 1944 || Bacteriophage || Phage Group and Phage Treaty || German–American biophysicist {{w|Max Delbrück}} initiates the {{w|Phage Group}}, advocating for the "Phage Treaty," which encourages phage researchers to concentrate on a limited number of phage and bacterial strains under standardized experimental conditions. This initiative aims to enhance the comparability and replicability of research across different laboratories, thereby unifying the field of bacterial genetics. Delbrück's efforts would significantly advance the study of bacteriophages and lay a foundation for future research in molecular biology and genetics.<ref>{{cite web |title=History: The Phage Group |url=http://www.cshl.edu/History/phagegroup.html |website=Cold Spring Harbor Laboratory |accessdate=2007-05-04 |archiveurl=https://web.archive.org/web/20070517203936/http://www.cshl.edu/History/phagegroup.html |archivedate=2007-05-17}}</ref> || {{w|United States}}
 |-
-| 1950 || || || ''Z. mays'': McClintock describes transposable elements. ||
+| 1946 || ''{{w|Escherichia coli}}'' || Genetic exchange || {{w|Joshua Lederberg}} and {{w|Edward Tatum}} discover bacterial conjugation through experiments with ''{{w|Escherichia coli}}'' (E. coli). They find that bacteria can exchange genetic information via a process called conjugation, which involves a bridge-like connection between two cells that facilitates the transfer of DNA from the donor to the recipient. The agent responsible for this transfer is the F ("fertility") factor, an extra-chromosomal genetic element that can replicate and move across cell membranes. For their pioneering work on bacterial genetics, Lederberg, Tatum, and George Beadle would be awarded the Nobel Prize in Physiology or Medicine in 1958. This discovery highlights that bacteria can change their genetic makeup in a manner analogous to sexual reproduction in more complex organisms, revealing conserved processes across different life forms.<ref>{{cite web |title=Joshua Lederberg - Facts |url=https://www.nobelprize.org/prizes/medicine/1958/lederberg/facts/ |website=Nobel Prize |accessdate=17 October 2024}}</ref><ref>{{cite web |title=Bacterium Conjugation |url=https://www.sciencedirect.com/topics/medicine-and-dentistry/bacterium-conjugation |website=ScienceDirect |accessdate=17 October 2024 |publisher=Medical Clinics of North America |date=2006}}</ref><ref>{{cite journal |title=How Escherichia coli Became the Flagship Bacterium of Molecular Biology |author=Natividad Ruiz, Thomas J. Silhavy |url=https://journals.asm.org/doi/10.1128/jb.00230-22 |doi=10.1128/jb.00230-22 |journal=Journal of Bacteriology |accessdate=17 October 2024}}</ref><ref>{{cite journal |title=Plasmid Transfer by Conjugation in Gram-Negative Bacteria: From the Cellular to the Community Level |author=Chloé Virolle, Kelly Goldlust, Sarah Djermoun, Sarah Bigot, Christian Lesterlin |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC7690428/ |doi=10.3390/genes11111239 |journal=Genes |date=2020-10-22 |volume=11 |issue=11 |page=1239 |pmid=33105635 |accessdate=17 October 2024}}</ref><ref>{{cite web |title=Bacterial Conjugation |url=https://www.sciencedirect.com/topics/biochemistry-genetics-and-molecular-biology/bacterial-conjugation |website=ScienceDirect |accessdate=17 October 2024}}</ref> || {{w|United States}}
 |-
-| 1951 || || || Phage lambda: Lederberg laboratory discovers phage and specialized transduction ||
+| 1949 || ''{{w|Saccharomyces cerevisiae}}'' || Model introduction || Jean Ephrussi and colleagues discover petite mutants in ''{{w|Saccharomyces cerevisiae}}'' (baker's yeast). These mutants have smaller colonies compared to normal strains and arise either spontaneously or after {{w|acriflavine}} treatment. They lack functional mitochondrial DNA (mtDNA), resulting in impaired mitochondrial function, respiratory deficiency, and altered metabolism. The discovery is significant for mitochondrial genetics, as ''S. cerevisiae'' would become a model organism for studying the relationship between mitochondrial DNA and cellular processes, providing insights into genetic inheritance and cellular respiration in eukaryotic organisms.<ref>{{cite journal |title=Transition of the ability to generate petites in the Saccharomyces/Kluyveromyces complex |author=Veronika Fekete, Mária Čierna, Silvia Poláková, Jure Piškur, Pavol Sulo |url=https://academic.oup.com/femsyr/article/7/8/1237/548562 |doi=10.1111/j.1567-1364.2007.00287.x |journal=FEMS Yeast Research |volume=7 |issue=8 |pages=1237–1247 |date=01 December 2007 |accessdate=17 October 2024}}</ref> || {{w|France}}
 |-
-| 1952 || || || Phage P22: Zinder and Lederberg discover transduction ||
+| 1950 || ''{{w|Zea mays}}'' || Publication || American scientist {{w|Barbara McClintock}} publishes her seminal paper titled ''The Origin and Behavior of Mutable Loci in Maize'' in the ''Proceedings of the National Academy of Sciences''. In this work, she describes the discovery of transposable elements, or "jumping genes," which can move along chromosomes and influence genetic inheritance. McClintock's research demonstrates that genomic replication does not always follow a predictable pattern, revolutionizing the understanding of genetic inheritance. Her findings on the Suppressor-Mutator (Spm) element, which can toggle between active and inactive forms due to methylation, further highlights the complexity of gene regulation. Her groundbreaking contributions would ultimately earn her a Nobel Prize.<ref>{{cite web |title=Barbara McClintock and the discovery of jumping genes |url=https://www.pnas.org/doi/pdf/10.1073/pnas.1219372109 |website=Proceedings of the National Academy of Sciences |accessdate=17 October 2024}}</ref><ref>{{cite web |title=Barbara McClintock and the Discovery of Jumping Genes |url=https://www.nature.com/scitable/topicpage/barbara-mcclintock-and-the-discovery-of-jumping-34083/ |website=Nature Scitable |accessdate=19 October 2024}}</ref><ref>{{cite journal |title=McClintock's challenge in the 21st century |author=Nina V. Fedoroff |journal=Proceedings of the National Academy of Sciences |volume=109 |issue=50 |pages=20200–20203 |date=2012-12-11 |doi=10.1073/pnas.1215482109 |pmid=23150590 |accessdate=17 October 2024}}</ref><ref>{{cite web |title=The Origin and Behavior of Mutable Loci in Maize |author=Barbara McClintock |url=https://embryo.asu.edu/pages/origin-and-behavior-mutable-loci-maize-1950-barbara-mcclintock |website=Embryo Project Encyclopedia |date=2014-10-24 |accessdate=17 October 2024}}</ref> || {{w|United States}}
 |-
-| 1953 || || || ''Aspergillus nidulans'': Pontecorvo describes genetic and parasexual systems ||
+| 1951 || ''{{w|Escherichia virus Lambda}}'' || Discovery || American microbiologist {{w|Esther Lederberg}}, while a Ph.D. student at the {{w|University of Wisconsin}}, isolates [[w:Lambda phage|bacteriophage lambda]] (λ), marking a significant discovery in microbial genetics. The details of this isolation and its implications for specialized transduction would be later elaborated upon in a 1953 paper co-authored with {{w|Joshua Lederberg}}. The discovery of λ facilitates a deeper understanding of genetic exchange mechanisms in bacteria, specifically through specialized transduction, where a bacteriophage transfers specific bacterial genes from one host to another, enhancing insights into gene mapping and bacterial genetics. This work establishes λ as a crucial model organism in genetic research.<ref>{{cite journal |last=Gottesman |first=Max E. |last2=Weisberg |first2=Robert A. |title=Little Lambda, Who Made Thee? |journal=Microbiology and Molecular Biology Reviews |date=December 2004 |volume=68 |issue=4 |pages=796–813 |doi=10.1128/MMBR.68.4.796-813.2004 |pmid=15590784 |pmc=539004}}</ref> || {{w|United States}}
 |-
-| 1954 || || || N. crassa: First major article on map construction in N. crassa ||
+| 1952 || ''{{w|Salmonella typhimurium}}'' || Discovery || {{w|Norton Zinder}} and {{w|Joshua Lederberg}} discover {{w|transduction}}, a process in which a virus ({{w|bacteriophage}}) transfers genetic material from a donor to a recipient bacterium. This breakthrough occurs at the {{w|University of Wisconsin–Madison}}, where they identify generalized transduction using the bacteriophage P22, which infects ''{{w|Salmonella typhimurium}}''. P22's unique DNA packaging allows some virions to carry fragments of the host's DNA, which can then be transferred to another bacterial cell. This discovery is the first identification of phage-mediated gene transfer, significantly advancing microbial genetics and providing a valuable tool for studying bacterial gene exchange.<ref>{{cite journal |title=The bacteriophage decides own tracks: When they are with or against the bacteria |url=https://pubmed.ncbi.nlm.nih.gov/34841341/ |journal=Current Research in Microbial Science |year=2022 |volume=2 |pages=100050 |author=Salsabil Makky, Alyaa Dawoud, Anan Safwat, Abdallah S Abdelsattar, Nouran Rezk, Ayman El-Shibiny|DOI=10.1016/j.crmicr.2021.100050}}</ref><ref>{{cite journal |title=Genetic transduction by phages and chromosomal islands: The new and noncanonical |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC6687093/ |journal=Current Opinion in Microbiology |year=2019 |volume=51 |pages=34-40 |author=Yin Ning Chiang, José R Penadés, John Chen |editor=Kimberly A Kline |PMID=31393945}}</ref><ref>{{cite book |title=DNA Packaging by Bacteriophage P22 |author=Sherwood Casjens, Peter Weigele |date=2005 |publisher=NCBI Bookshelf |website=ncbi.nlm.nih.gov |url=https://www.ncbi.nlm.nih.gov/books/NBK6430/ |accessdate=19 October 2024}}</ref><ref>{{cite web |title=Enterobacteria Phage P22 |website=ScienceDirect |url=https://www.sciencedirect.com/topics/biochemistry-genetics-and-molecular-biology/enterobacteria-phage-p22 |publisher=ScienceDirect |accessdate=19 October 2024}}</ref><ref>{{cite journal |last=Parkinson |first=John S. |title=Classic Spotlight: The Discovery of Bacterial Transduction |journal=Journal of Bacteriology |date=7 October 2016 |volume=198 |issue=21 |pages=2899–2900 |doi=10.1128/JB.00635-16 |pmid=27736750 |pmc=5055593}}</ref> || {{w|United States}}
 |-
-| 1956 || || || C. reinhardtii: Levine develops important genetic programme ||
+| 1953 || ''{{w|Tetrahymena thermophila}}'' || Model introduction || David L. Nanney first describes the MAT locus in ''{{w|Tetrahymena thermophila}}''. This discovery is instrumental in understanding the organism's mating types and genetic mechanisms, further establishing ''Tetrahymena'' as a model organism for studying cellular biology and genetics.<ref>{{cite journal |title=Selecting One of Several Mating Types through Gene Segment Joining and Deletion in Tetrahymena thermophila |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC3608545/ |journal=PLoS Biol |volume=11 |issue=3 |pages=e1001518 |doi=10.1371/journal.pbio.1001518 |author=Marcella D Cervantes, Eileen P Hamilton, Jie Xiong, Michael J Lawson, Dongxia Yuan, Michalis Hadjithomas, Wei Miao, Eduardo Orias |date=26 March 2013 |accessdate=20 October 2024}}</ref> || {{w|United States}}
+|-
+| 1953 || ''{{w|Aspergillus nidulans}}'' || Model introduction || Italian-born Scottish geneticist {{w|Guido Pontecorvo}} and his team publish a key study in ''Advances in Genetics'', describing both the genetic and parasexual systems of ''{{w|Aspergillus nidulans}}''. This work is pivotal in establishing ''A. nidulans'' as a genetically tractable model organism for studying fungal genetics. Pontecorvo's discovery of the parasexual cycle, an alternative to sexual reproduction, allows for genetic analysis in organisms lacking a traditional sexual cycle. This groundbreaking research opens doors for genetic mapping in fungi and even has implications for studying genetic recombination in human somatic cells, shaping the future of microbial genetics and genetic analysis techniques.<ref>{{cite journal |last=Cohen |first=Bernard L. |title=Guido Pontecorvo (“Ponte”): A Centenary Memoir |journal=Genetics |date=November 2007 |volume=177 |issue=3 |pages=1439–1444 |doi=10.1093/genetics/177.3.1439 |pmid=18039877 |pmc=2147990}}</ref> || {{w|United Kingdom}}
 |-
-| 1958 || || || C. reinhardtii: Gillham begins genetics of chloroplast ||
+| 1954 || ''{{w|Neurospora crassa}}'' || Genetic mapping || Barratt and colleagues publish the first major article on map construction in ''{{w|Neurospora crassa}}'' titled ''Map construction in Neurospora crassa'', marking a key moment in fungal genetics. This work, which would become a foundational resource in the field, combines theoretical insights with experimental findings and provides a comprehensive compendium of ''N. crassa'' mutants. The study contributes significantly to the standardization of genetic methods, strains, and nomenclature. It also advances the understanding of meiotic phenomena like segregation and intrachromosomal recombination, paving the way for deeper insights into gene conversion and molecular recombination.<ref>{{cite journal |title=The fungal genetic system: a historical overview |author=Rowland H. Davis |journal=Journal of Genetics |volume=75 |number=3 |pages=245–253 |date=December 1996 |url=https://www.ias.ac.in/article/fulltext/jgen/075/03/0245-0253 |publisher=Indian Academy of Sciences |accessdate=17 October 2024}}</ref> || {{w|United States}}
 |-
-| 1958 || || || Tetrahymena thermophila: Allen and Nanney describe genetic system ||
+| 1965 || ''{{w|Arabidopsis thaliana}}'' || Model introduction || Gerhard Röbbelen organizes the first International ''{{w|Arabidopsis}}'' Symposium. This event marks a significant step in the recognition and promotion of ''{{w|Arabidopsis thaliana}}'' as a model organism in plant genetics and developmental biology. The symposium brings together researchers interested in using ''Arabidopsis'' for genetic studies, helping to solidify its status as a key model organism due to its small genome, short generation time, and ease of genetic manipulation. This symposium plays a crucial role in fostering a community of scientists focused on ''Arabidopsis'', which would later become one of the most widely studied organisms in plant biology.<ref>{{cite journal |title=A fortunate choice: the history of Arabidopsis as a model plant |author=Chris Somerville, Maarten Koornneef |journal=Nature Reviews Genetics |volume=3 |issue=11 |pages=883–889 |date=2002 |doi=10.1038/nrg927 |issn=1471-0056 |pmid=12415318 |accessdate=17 October 2024}}</ref> || {{w|Germany}}
 |-
-| 1960 || || || ''E. coli'': Jacob and Wollman fully describe genetic system. ||
+| 1965 || ''{{w|Caenorhabditis elegans}}'' || Model introduction || South African biologist {{w|Sydney Brenner}} selects ''{{w|Caenorhabditis elegans}}'' as a model organism to study genetics, development, and neural function. Over the years, this {{w|nematode}} would become one of the most comprehensively understood metazoans, particularly in terms of {{w|anatomy}}, {{w|genetics}}, and behavior. Two major accomplishments in ''C. elegans'' research include the mapping of its complete cell lineage and the creation of a "wiring diagram" of its cell contacts through electron microscopy. Advances also include molecular cloning of developmental genes using transposon tagging and the development of a physical genome map. This knowledge serves as a foundation for future research in biology.<ref>{{cite web |title=Genome Research |url=https://www.cshlpress.com/default.tpl?cart=1729217571804803394&fromlink=T&linkaction=full&linksortby=oop_title&--eqSKUdatarq=39 |website=cshlpress.com |accessdate=16 October 2024}}</ref><ref>{{cite web |title=C. elegans II. 2nd edition |url=https://www.ncbi.nlm.nih.gov/books/NBK20086/ |website=ncb.nlm.nih.gov |accessdate=17 October 2024}}</ref> || {{w|United Kingdom}}
 |-
-| 1965 || || || ''A. thaliana'': First International Arabidopsis Symposium. ||
+| 1966 || || Literature || American internist and medical geneticist {{w|Victor A. McKusick}} publishes the first edition of ''Mendelian Inheritance in Man'' (MIM), a comprehensive knowledge base of human genes and genetic disorders. Initially developed in the early 1960s, MIM categorizes autosomal dominant, autosomal recessive, and X-linked phenotypes. The first print edition launched in 1966, with 12 editions published by 1998. In 1987, MIM would transition online as OMIM (Online Mendelian Inheritance in Man) at {{w|Johns Hopkins University}}, later moving to the National Center for Biotechnology Information in 1995. Today, OMIM is updated daily and serves as a crucial resource for students, researchers, and clinicians in genetics and genomics.<ref>{{cite web |title=Mendelian Inheritance in Man and Its Online Version, OMIM |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC1852721/ |author=Victor A McKusick |website=PMC |accessdate=16 October 2024}}</ref> || {{w|United States}}
 |-
-| 1965 || || || ''Caenorhabditis elegans'': Brenner proposes programme in genetics of neural development. ||
+| 1974 || ''{{w|Caenorhabditis elegans}}'' || Genetic screening || {{w|Sydney Brenner}} conducts a groundbreaking genetic screen and creates the first genetic map for ''{{w|Caenorhabditis elegans}}'' (''C. elegans''), isolating notable mutants such as Dumpy, Squat, Long, Blistered, and Roller. His genetic map identifies over 100 loci, and he estimates the genome to contain about 2,000 essential genes. As a nematode, ''C. elegans'' serves as a vital model organism in genetics due to its short lifespan, rapid generation time, and hermaphroditic nature, allowing for efficient isolation and characterization of mutants. Brenner's research also reveals the significance of a "nonsense" codon in mRNA, crucial for protein synthesis.<ref>{{cite web |title=Sydney Brenner |url=https://www.cshl.edu/personal-collections/sydney-brenner/ |website=Cold Spring Harbor Laboratory |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Essential genes |url=https://www.ncbi.nlm.nih.gov/books/NBK19771/ |author=Kenneth Kemphues |website=NCBI Bookshelf |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=The Ancestral Caenorhabditis elegans Cuticle Suppresses rol-1 |url=https://academic.oup.com/g3journal/article/10/7/2385/6026277 |author=Luke M Noble, Asif Miah, Taniya Kaur, Matthew V Rockman |journal=G3: Genes|volume=10 |issue=7 |pages=2385–2395 |date=01 July 2020 |doi=10.1534/g3.120.401336 |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=What Can We Learn About Human Disease from the Nematode C. elegans? |author=Javier Apfeld, Scott Alper |journal=Methods in Molecular Biology |volume=1706 |pages=53–75 |year=2018 |doi=10.1007/978-1-4939-7471-9_4 |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC6391162}}</ref><ref>{{cite journal |title=The art and design of genetic screens: Caenorhabditis elegans |author=Erik M Jorgensen, Susan E Mango |journal=Nature Reviews Genetics |volume=3 |issue=5 |pages=356–369 |year=2002 |doi=10.1038/nrg794 |url=https://pubmed.ncbi.nlm.nih.gov/11988761/}}</ref><ref>{{cite web |title=Essential genes |url=https://www.ncbi.nlm.nih.gov/books/NBK20086/ |author=Kenneth Kemphues |website=NCBI Bookshelf |accessdate=16 October 2024}}</ref> || {{w|United Kingdom}}
 |-
-| 1966 || || || ''Homo sapiens'': First edition of Mendelian Inheritance in Man ||
+| 1980 || ''{{w|Drosophila melanogaster}}'' || Embryonic mutagenesis || In their study published in ''[[w:Nature (journal)|Nature]]'', {{w|Christiane Nüsslein-Volhard}} and Eric Wieschaus investigate embryonic lethal mutants of ''{{w|Drosophila melanogaster}}'', identifying 15 loci that significantly alter the segmental pattern of the larva. This comprehensive screening aims to uncover genes involved in segmentation, revealing that the segmentation process encompasses multiple levels of spatial organization: the egg as a developmental unit, a repeat unit spanning two segments, and the individual segment itself. Their findings not only provide insights into the genetic architecture governing body segmentation in ''{{w|Drosophila}}'' but also lays a foundation for understanding developmental biology. This work contributes to the broader use of ''Drosophila'' as a model organism in genetics, influencing research on gene function and developmental processes across various species.<ref>{{cite journal |title=Mutations affecting segment number and polarity in Drosophila |author=C Nüsslein-Volhard, E Wieschaus |journal=Nature |date=1980-10-30 |volume=287 |issue=5785 |pages=795-801 |doi=10.1038/287795a0 |PMID=6776413 |accessdate=16 October 2024}}</ref> || {{w|Germany}}
+|-
+| 1981 || ''Danio rerio'' || Clonal production || American molecular biologist {{w|George Streisinger}} et al. publish a pivotal study in ''[[w:Nature (journal)|Nature]]'' detailing the production of clones of homozygous diploid zebrafish (''danio rerio''). This research demonstrates that large-scale production of homozygous diploid zebrafish can be achieved through simple physical treatments. By cloning individual homozygotes, the researchers established a reliable method for generating genetically identical fish, which is crucial for conducting genetic analyses in vertebrates. The availability of these cloned homozygous zebrafish not only advances the understanding of genetic inheritance and development in this model organism but also paves the way for further research in {{w|genetics}}, {{w|developmental biology}}, and {{w|toxicology}}, solidifying zebrafish's role as a valuable tool in scientific inquiry.<ref>{{cite journal |title=Production of clones of homozygous diploid zebra fish (Brachydanio rerio) |author=G Streisinger, C Walker, N Dower, D Knauber, F Singer |journal=Nature |volume=291 |pages=293 |date=February 1981 |doi=10.1038/291293a0 |url=https://pubmed.ncbi.nlm.nih.gov/7248006/ |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Zebrafish (Danio rerio) |url=https://www.researchgate.net/figure/Zebrafish-Danio-rerio-https-enwikipediaorg-wiki-Zebrafish-George-Streisinger_fig1_328415952 |accessdate=16 October 2024}}</ref> || {{w|United States}}
 |-
-| 1974 || || || ''C. elegans'': Important genetics publication. ||
+| 1984 || ''{{w|Arabidopsis thaliana}}'' || Genome estimation || Leutwiler et al. utilize reassociation kinetics and quantitative gel blot hybridization to estimate the genome size of ''{{w|Arabidopsis thaliana}}'' at approximately 70 megabases (Mb). However, other techniques, including Feulgen photometry and flow cytometry, suggest a range of 0.085–0.215 picograms (pg) for the genome size. The actual genome size of ''A. thaliana'' is closer to 135 Mb, consisting of five pairs of chromosomes, making it the smallest genome among flowering plants. The complete genome sequence would be published in 2000, revealing around 25,498 predicted genes. Variations in genome size among organisms can be attributed to differences in the amplification, deletion, and divergence of repetitive sequences, highlighting the complexity of genomic structure and evolution.<ref>{{cite web |title=Arabidopsis thaliana |url=https://www.sciencedirect.com/topics/biochemistry-genetics-and-molecular-biology/arabidopsis-thaliana |website=ScienceDirect |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Annotation for Arabidopsis thaliana |url=https://plants.ensembl.org/Arabidopsis_thaliana/Info/Annotation |website=Ensembl Plants |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Differences in Genome Size Between Closely Related Species: The Drosophila melanogaster Species Subgroup |author=Matthieu Boulesteix, Michèle Weiss, Christian Biémont |journal=Molecular Biology and Evolution |volume=23 |issue=1 |pages=162–167 |date=January 2006 |doi=10.1093/molbev/msj012 |url=https://academic.oup.com/mbe/article/23/1/162/1193365 |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Comparisons with Caenorhabditis (∼100 Mb) and Drosophila (∼175 Mb) Using Flow Cytometry Show Genome Size in Arabidopsis to be ∼157 Mb and thus ∼25 % Larger than the Arabidopsis Genome Initiative Estimate of ∼125 Mb |author=MICHAEL D BENNETT, ILIA J LEITCH, H JAMES PRICE, J SPENCER JOHNSTON |journal=Annals of Botany |volume=91 |issue=5 |pages=547–557 |date=April 2003 |doi=10.1093/aob/mcg057 |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC4242247/ |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Genome Size Variation among Accessions of Arabidopsis thaliana |author=HEIKE SCHMUTHS, ARMIN MEISTER, RALF HORRES, KONRAD BACHMANN |journal=Annals of Botany |volume=93 |issue=3 |pages=317–321 |date=March 2004 |doi=10.1093/aob/mch037 |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC4242198/ |accessdate=16 October 2024}}</ref> || {{w|United States}}
 |-
-| 1980 || || || ''D. melanogaster'': NüssleinVolhard and Wieschaus isolate developmental mutants. ||
+| 1996 (April) || ''{{w|Saccharomyces cerevisiae}}'' || {{w|Genome sequencing}} || The complete genome of ''{{w|Saccharomyces cerevisiae}}'' (baker’s yeast) strain S288C is sequenced, making it the first eukaryotic genome to be fully mapped. This significant milestone involves over 600 scientists from North America, Japan, and Europe, who collaboratively sequenced its 12 million base pairs across 16 chromosomes. The ''S. cerevisiae'' genome contains about 6,000 genes, with roughly one-third having human counterparts. This achievement underscores ''S. cerevisiae'''s role as a model organism in eukaryotic genomics, proving invaluable for studying fundamental processes, such as cell cycle regulation, which is crucial for cancer research. Additionally, this unicellular fungus is widely used in the food and beverage industry.<ref>{{cite web |title=1996 Release: Yeast Genome Sequenced |url=https://www.genome.gov/10000510/1996-release-yeast-genome-sequenced |website=Genome.gov |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Online Education Kit: 1996 Yeast Genome Sequenced |url=https://www.genome.gov/25520379/online-education-kit-1996-yeast-genome-sequenced |website=Genome.gov |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Yeast: one cell, one reference sequence, many genomes? |author=Szymanski, Erika; Vermeulen, Niki; Wong, Mark |journal=New Genetics and Society |volume=38 |issue=4 |pages=430-450 |year=2019 |doi=10.1080/14636778.2019.1677150 |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Introducing Saccharomyces cerevisiae: The Best Known Yeast in the World |url=https://quadram.ac.uk/blogs/introducing-saccharomyces-cerevisiae-the-best-known-yeast-in-the-world/ |website=Quadram Institute |accessdate=16 October 2024}}</ref><ref>{{cite journal |title=Saccharomyces cerevisiae and its industrial applications |author=Maria Parapouli, Anastasios Vasileiadis, Amalia-Sofia Afendra, Efstathios Hatziloukas |journal=AIMS Microbiology |date=11 February 2020 |volume=6 |issue=1 |pages=1–31 |doi=10.3934/microbiol.2020001 |accessdate=16 October 2024}}</ref> || {{w|North America}}, {{w|Japan}}, {{w|Europe}}
 |-
-| 1981 || || || ''D. rerio'': Clonal propagation method published. ||
+| 1996 || ''Danio rerio'' || Mutant screening || A large-scale screen for developmental mutants in ''Danio rerio'' ({{w|zebrafish}}) is led by researchers {{w|Christiane Nüsslein-Volhard}} in Tübingen, Germany, and Wolfgang Driever in Boston, USA. This extensive effort results in thousands of mutant lines exhibiting defects in major organ systems and embryo patterning, with the findings published in a special issue of the journal [[w:Development (journal)|Development]], which would remain its largest issue to date. The success of this screen would inspire many research centers to continue exploring mutant genetics. The zebrafish would emerge as a prominent model organism in vertebrate developmental biology due to its small size, low maintenance cost, and the transparency of its early embryos, which facilitates the screening of morphological defects.<ref>{{cite journal |title=Large-scale mutagenesis in the zebrafish: in search of genes controlling development in a vertebrate |author=Mary C. Mullins, Matthias Hammerschmidt, Pascal Haffter, Christiane Nüsslein-Volhard |journal=Current Biology |year=2000 |url=https://www.cell.com/current-biology/abstract/S0960-9822(00)00048-8 |accessdate=2024-10-16}}</ref><ref>{{cite journal |title=Zebrafish Make a Big Splash |author=Judith S Eisen |journal=Cell |volume=87 |issue=6 |pages=969–977 |year=1996 |url=https://www.cell.com/fulltext/S0092-8674(00)81792-4 |accessdate=2024-10-16}}</ref><ref>{{cite journal |title=Zebrafish as a Developmental Model Organism for Pediatric Research |author=Matthew B Veldman, Shuo Lin |journal=Pediatric Research |volume=64 |pages=470–476 |year=2008 |url=https://www.nature.com/articles/pr2008227 |accessdate=2024-10-16}}</ref><ref>{{cite journal |title=The State of the Art of the Zebrafish Model for Toxicology and Toxicologic Pathology Research—Advantages and Current Limitations |author=Jan M Spitsbergen, Michael L Kent |journal=Environmental Health Perspectives |volume=114 |issue=4 |pages=545–552 |year=2006 |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC1909756 |accessdate=2024-10-16}}</ref> || {{w|Germany}}, {{w|United States}}
+|-
+| 1997 || ''{{w|Escherichia coli}}'' || {{w|Genome sequencing}} || The complete genome sequence of ''{{w|Escherichia coli}}'' is published, providing significant insights into one of the most extensively studied bacteria in molecular biology. ''E. coli'' resides in the lower intestinal tract of animals and is typically harmless, making it an ideal model organism for biochemists and geneticists. By this time, researchers had long used ''E. coli'' to investigate fundamental biochemical processes and {{w|gene regulation}}. The sequenced genome, comprising approximately 4.6 million base pairs and around 4,000 genes, enables scientists to deepen their understanding of its biology. Moreover, comparing non-pathogenic strains with pathogenic ones can aid in developing treatments and preventive strategies for illnesses such as food poisoning.<ref>{{cite web |title=Online Education Kit: 1997 E. coli Genome Sequenced |url=https://www.genome.gov/25520386/online-education-kit-1997-e-coli-genome-sequenced |website=Genome.gov |accessdate=2024-10-16}}</ref> || {{w|United States}}
 |-
-| 1984 || || || ''A. thaliana'': Leutwiler et al. determine genome size. ||
+| 1998 || ''{{w|Caenorhabditis elegans}}'' || {{w|Genome sequencing}} || The genome of ''{{w|Caenorhabditis elegans}}'', a small soil-dwelling nematode, is fully sequenced and published in ''[[w:Science (journal)|Science]]'', making it the first multicellular organism to have its genome completed. This effort, led by the ''C. elegans'' Sequencing Consortium—a collaboration between the Genome Sequencing Center in St. Louis and the Sanger Centre in Hinxton—demonstrates the feasibility of high-throughput sequencing techniques, crucial for the {{w|Human Genome Project}}. With an accuracy of one error per 10,000 bases, the ''C. elegans'' genome becomes a valuable resource for gene discovery. Its use as a model organism provides insights into genetic function and developmental biology, further influencing biomedical research.<ref>{{cite journal |title=Arabidopsis Genome Initiative |url=https://pubmed.ncbi.nlm.nih.gov/10098407/ |journal=Nature |accessdate=2024-10-16}}</ref><ref>{{cite web |title=The Pilot Project for the Human Genome Project: Sequencing C. elegans |url=https://www.yourgenome.org/theme/the-pilot-project-for-the-human-genome-project-sequencing-ic-elegans-i/ |website=Your Genome |accessdate=2024-10-16}}</ref><ref>{{cite web |title=Online Education Kit: 1998 Genome of Roundworm C. elegans Sequenced |url=https://www.genome.gov/25520394/online-education-kit-1998-genome-of-roundworm-c-elegans-sequenced |website=Genome.gov |accessdate=2024-10-16}}</ref> || {{w|United States}}, {{w|United Kingdom}}
 |-
-| 1986 || || || ''D. rerio'': Important genetics publication. ||
+| 2000 (December) || ''{{w|Arabidopsis thaliana}}'' || {{w|Genome sequencing}} || The genome of ''{{w|Arabidopsis thaliana}}'', a small flowering plant, is fully sequenced, marking a historic milestone as the first complete genome of a flowering plant, and launching the era of plant genomics. This achievement provides an essential genetic reference that would be freely accessible to scientists, revolutionizing plant science by enabling in-depth studies on plant genetics, growth, and development. The global collaboration involves major institutions such as Stanford Genome Technology Center and Cold Spring Harbor Laboratory. The ''A. thaliana'' genome spans around 125 megabases, contains roughly 25,500 genes, and features about 35% unique genes, with evidence of ancient polyploidy in large segmental duplications.<ref>{{cite journal |title=Publication of the complete genome sequence: Importance for comparative genomics and pan-genomes |url=https://www.sciencedirect.com/science/article/abs/pii/S1360138521002818 |journal=Current Opinion in Genetics & Development |date=2021 |accessdate=2024-10-16}}</ref><ref>{{cite web |title=The Human Genome: December 2000 Update |url=https://www.nsf.gov/pubs/2002/bio0202/genome.htm |website=National Science Foundation |accessdate=2024-10-16}}</ref><ref>{{cite journal |title=Twenty Years Ago: The Arabidopsis Genome Sequencing Project |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC8226293/ |journal=Proceedings of the National Academy of Sciences |accessdate=2024-10-16}}</ref> || {{w|United States}}
 |-
-| 1996 || || || ''S. cerevisiae'': Genome sequenced. ||
+| 2000 || ''{{w|Drosophila melanogaster}}'' || {{w|Genome sequencing}} || The genome of the fruit fly ''{{w|Drosophila melanogaster}}'' is sequenced in a groundbreaking effort published in the March 24, 2000 issue of ''[[w:Science (journal)|Science]]''. This project, a collaboration between {{w|Celera Genomics}} and the Drosophila Genome Projects, marks the first successful application of the whole genome shotgun (WGS) method in a multicellular organism. Researchers sequence approximately 97 to 98 percent of the genome, capturing nearly all of the estimated 13,600 genes. This achievement is significant in genetic research, establishing a precedent for future genome projects. Further improvements and annotations would since be made by the Berkeley Drosophila Genome Project and FlyBase.<ref>{{cite web |title=Drosophila Genome Sequenced |url=https://doe-humangenomeproject.ornl.gov/drosophila-genome-sequenced |website=DOE Human Genome Project |accessdate=29 September 2024}}</ref><ref>{{cite journal |title=Title of the Article |author=Author(s) |journal=Journal Name |year=2006 |volume= |issue= |pages= |doi=10.1186/gb-2006-7-1-r10 |url=https://pubmed.ncbi.nlm.nih.gov/16339363/#:~:text=Abstract,be%20solved%20in%20the%20future.}}</ref><ref>{{cite web |title=Drosophila Genome Sequence Completed |url=https://www.hhmi.org/news/drosophila-genome-sequence-completed |website=HHMI |date=2000 |accessdate=2024-10-16}}</ref> || {{w|United States}}
 |-
-| 1996 || || || ''D. rerio'': Large-scale screen for developmental mutants. ||
+| 2001 || {{w|Human}} || {{w|Genome sequencing}} || The International Human Genome Sequencing Consortium publishes a draft sequence of the human genome in ''[[w:Nature (journal)|Nature]]'', marking a landmark achievement in genomics. This collaborative project involves 16 genome centers worldwide, which coordinated efforts through regular meetings and phone conferences. Following a policy of rapid data release, the consortium made the assembled genome sequence publicly available within 24 hours. The draft reveals key insights: the human genome spans 2.85 billion bases, with DNA sequences that are 99.9% identical between individuals. It includes approximately 22,300 protein-coding genes, numerous segmental duplications, and over 3 million single nucleotide polymorphisms (SNPs). The Human Genome Project lays a crucial foundation for biomedical research, enabling advancements in disease research and personalized medicine.<ref>{{cite journal |last=International Human Genome Sequencing Consortium |title=Finishing the euchromatic sequence of the human genome |journal=Nature |date=2004-10-21 |volume=431 |issue=7011 |pages=931-945 |doi=10.1038/nature03001 |pmid=15496913 |url=https://pubmed.ncbi.nlm.nih.gov/15496913/ |accessdate=2024-10-16}}</ref><ref>{{cite journal |last=International Human Genome Sequencing Consortium |title=Initial sequencing and analysis of the human genome |journal=Nature |date=2001-02-01 |volume=409 |pages=860–921 |url=https://www.nature.com/articles/35057062 |accessdate=2024-10-16}}</ref><ref>{{cite web |title=KEGG Homo sapiens Genome Database |url=https://www.genome.jp/dbget-bin/www_bget?hsa |website=KEGG |accessdate=16 October 2024}}</ref> || {{w|United States}}
 |-
-| 1997 || || || ''E. coli'': Genome sequenced. ||
+| 2002 || || {{w|Genome sequencing}} || The Mouse Genome Sequencing Consortium—including the {{w|Broad Institute}}, {{w|Washington University}}, and the {{w|Sanger Institute}}—publish a draft of the mouse genome, using the C57BL/6J strain, the most widely studied inbred strain. This draft revealed around 24,500 protein-coding genes across 19 autosomal pairs, plus X and Y sex chromosomes. The mouse genome, about 14% shorter than the human genome, aligns with large human chromosomal segments, and 75% of mouse genes have direct human counterparts. The similarities between mouse and human genomes make the mouse an essential model for studying human disease, with its short lifecycle and rapid breeding facilitating genetic studies on a large scale.<ref>{{cite web |title=UniProt: Homo sapiens (Human) Proteome |url=https://www.uniprot.org/proteomes/UP000000589 |website=UniProt |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Mouse Genome Project |url=https://www.broadinstitute.org/mouse/mouse-genome-project |website=Broad Institute |accessdate=16 October 2024}}</ref><ref>{{cite web |title=Online Education Kit: 2002 Mouse Genome Sequenced |url=https://www.genome.gov/25520486/online-education-kit-2002-mouse-genome-sequenced |website=Genome.gov |accessdate=16 October 2024}}</ref> || {{w|United States}}
 |-
-| 1998 || || || ''C. elegans'': Genome sequenced. ||
+| 2003 || ''{{w|Neurospora crassa}}'' || {{w|Genome sequencing}} || The complete genome sequence of ''{{w|Neurospora crassa}}'' is reported, revealing a genome size of 43 million base pairs that encodes approximately 10,000 genes organized into seven linkage groups. The sequencing is conducted using whole genome shotgun sequencing and paired-end sequencing methods. Notably, ''N. crassa'' features a G + C content of about 50% and exhibits minimal repetitive DNA, making it a valuable model organism for studying genetics, gene regulation, and metabolic processes. This comprehensive genomic information enhances the understanding of fungal biology and facilitates comparative studies across various organisms.<ref>{{cite web |title=Neurospora crassa |url=https://www.sciencedirect.com/topics/biochemistry-genetics-and-molecular-biology/neurospora-crassa |website=sciencedirect.com |accessdate=19 October 2024}}</ref> || {{w|United States}}
 |-
-| 2000 || || || ''A. thaliana'': Genome sequenced. ||
+| 2003 || ''{{w|Nothobranchius furzeri}}'' || Model introduction || ''{{w|Nothobranchius furzeri}}'' (Turquoise killifish) is introduced as model organism for aging and longevity research due to its short lifespan and vulnerability to age-related diseases. With a maximum survival time of under 12 weeks in laboratory conditions, ''N. furzeri'' exhibits a typical age-dependent increase in mortality. This species' short lifespan, small size, and ability to be propagated in captivity make it ideal for studying the biology of aging. Researchers, including Stefano Valdesalici and Alessandro Cellerino, highlighted its potential in evolutionary studies related to senescence and the effects of natural selection on lifespan.<ref>{{cite journal |title=Extremely short lifespan in the annual fish Nothobranchius furzeri |url=https://royalsocietypublishing.org/doi/10.1098/rsbl.2003.0048 |author=Stefano Valdesalici, Alessandro Cellerino |journal=Biology Letters |date=7 November 2003 |doi=10.1098/rsbl.2003.0048 |accessdate=24 October 2024}}</ref>
 |-
-| 2000 || || || D. melanogaster: Genome sequenced. ||
+| 2004 || || Literature || Pamela M. Carroll and Kevin Fitzgerald publish ''Model Organisms in Drug Discovery'', which explores the use of model organisms like ''Drosophila'' (fruit flies), ''C. elegans'' worms, yeast, mice, and zebrafish in drug discovery and biomedical research. These organisms share many biological pathways with humans, making them invaluable for studying diseases and testing therapeutic compounds. The book highlights how advances in {{w|bioinformatics}}, {{w|proteomics}}, and automation technologies have enhanced the use of these models on an industrial scale. Leading experts discuss real-life applications of model organisms in therapeutic areas, their role in drug discovery, and their potential future impact.<ref>{{cite book |title=Model Organisms in Drug Discovery |url=https://books.google.com.ar/books/about/Model_Organisms_in_Drug_Discovery.html?id=WZAylt3It-kC&source=kp_book_description&redir_esc=y |publisher=John Wiley & Sons |year=2012 |accessdate=20 October 2024}}</ref> || {{w|United States}}
 |-
-| 2001 || || || ''H. sapiens'': Genome sequenced. ||
+| 2007 || ''{{w|Chlamydomonas reinhardtii}}'' || {{w|Genome sequencing}} || The complete nuclear genome sequence of ''{{w|Chlamydomonas reinhardtii}}'' is published. This organism diverged from land plants over 1 billion years ago and serves as a model for studying chloroplast-based photosynthesis and the structure and function of eukaryotic flagella, which are absent in land plants. The ∼120-megabase genome is sequenced, and comparative phylogenomic analyses identify genes encoding uncharacterized proteins likely related to chloroplast and flagellar function. This research enhances our understanding of the ancestral eukaryotic cell, uncovers novel genes associated with photosynthesis and flagellar functions, and links ciliopathy with flagellar composition and function.<ref>{{cite journal |last=Merchant |first=Sabeeha S. |last2=Prochnik |first2=Simon E. |last3=Vallon |first3=Olivier |last4=Harris |first4=Elizabeth H. |last5=Karpowicz |first5=Steven J. |last6=Witman |first6=George B. |last7=Terry |first7=Astrid |last8=Salamov |first8=Asaf |last9=Fritz-Laylin |first9=Lillian K. |last10=Maréchal-Drouard |first10=Laurence |last11=Marshall |first11=Wallace F. |last12=Qu |first12=Liang-Hu |last13=Nelson |first13=David R. |last14=Sanderfoot |first14=Anton A. |last15=Spalding |first15=Martin H. |last16=Kapitonov |first16=Vladimir V. |last17=Ren |first17=Qinghu |last18=Ferris |first18=Patrick |last19=Lindquist |first19=Erika |last20=Shapiro |first20=Harris |last21=Lucas |first21=Susan M. |last22=Grimwood |first22=Jane |last23=Schmutz |first23=Jeremy |last24=Cardol |first24=Pierre |last25=Cerutti |first25=Heriberto |last26=Chanfreau |first26=Guillaume |last27=Chen |first27=Chun-Long |last28=Cognat |first28=Valérie |last29=Croft |first29=Martin T. |last30=Dent |first30=Rachel |last31=Dutcher |first31=Susan |last32=Fernández |first32=Emilio |last33=Fukuzawa |first33=Hideya |last34=González-Ballester |first34=David |last35=González-Halphen |first35=Diego |last36=Hallmann |first36=Armin |last37=Hanikenne |first37=Marc |last38=Hippler |first38=Michael |last39=Inwood |first39=William |last40=Jabbari |first40=Kamel |last41=Kalanon |first41=Ming |last42=Kuras |first42=Richard |last43=Lefebvre |first43=Paul A. |last44=Lemaire |first44=Stéphane D. |last45=Lobanov |first45=Alexey V. |last46=Lohr |first46=Martin |last47=Manuell |first47=Andrea |last48=Meier |first48=Iris |last49=Mets |first49=Laurens |last50=Mittag |first50=Maria |last51=Mittelmeier |first51=Telsa |last52=Moroney |first52=James V. |last53=Moseley |first53=Jeffrey |last54=Napoli |first54=Carolyn |last55=Nedelcu |first55=Aurora M. |last56=Niyogi |first56=Krishna |last57=Novoselov |first57=Sergey V. |last58=Paulsen |first58=Ian T. |last59=Pazour |first59=Greg |last60=Purton |first60=Saul |last61=Ral |first61=Jean-Philippe |last62=Riaño-Pachón |first62=Diego Mauricio |last63=Riekhof |first63=Wayne |last64=Rymarquis |first64=Linda |last65=Schroda |first65=Michael |last66=Stern |first66=David |last67=Umen |first67=James |last68=Willows |first68=Robert |last69=Wilson |first69=Nedra |last70=Zimmer |first70=Sara Lana |last71=Allmer |first71=Jens |last72=Balk |first72=Janneke |last73=Bisova |first73=Katerina |last74=Chen |first74=Chong-Jian |last75=Elias |first75=Marek |last76=Gendler |first76=Karla |last77=Hauser |first77=Charles |last78=Lamb |first78=Mary Rose |last79=Ledford |first79=Heidi |last80=Long |first80=Joanne C. |last81=Minagawa |first81=Jun |last82=Page |first82=M. Dudley |last83=Pan |first83=Junmin |last84=Pootakham |first84=Wirulda |last85=Roje |first85=Sanja |last86=Rose |first86=Annkatrin |last87=Stahlberg |first87=Eric |last88=Terauchi |first88=Aimee M. |last89=Yang |first89=Pinfen |last90=Ball |first90=Steven |last91=Bowler |first91=Chris |last92=Dieckmann |first92=Carol L. |last93=Gladyshev |first93=Vadim N. |last94=Green |first94=Pamela |last95=Jorgensen |first95=Richard |last96=Mayfield |first96=Stephen |last97=Grossman |first97=Arthur R. |date=October 12, 2007 |title=The Chlamydomonas Genome Reveals the Evolution of Key Animal and Plant Functions |journal=Science |volume=318 |issue=5848 |pages=245–250 |doi=10.1126/science.1143609 |pmid=17932292}}</ref> ||
 |-
-| 2002 || || || ''M. musculus'': Genome sequenced ||
+| 2009 || || Literature || David A. Crotty and Alexander Gann publish ''Emerging Model Organisms: A Laboratory Manual'', which explores the growing diversity of model organisms used in molecular, cellular, and {{w|developmental biology}}. With advances in {{w|gene expression}} technology, lower genome sequencing costs, and a rising interest in evolutionary biology, by this time researchers expand beyond traditional models. The book introduces 23 emerging model organisms, such as bats, butterflies, snails, and tomatoes, each offering unique research opportunities.<ref>{{cite book |title=Emerging Model Organisms |url=https://books.google.com.ar/books/about/Emerging_Model_Organisms.html?id=3m3wAAAAMAAJ&source=kp_book_description&redir_esc=y |publisher=Cold Spring Harbor Laboratory Press |year=2009 |accessdate=20 October 2024}}</ref> ||
 |-
-| 2003 || || || ''N. crassa'': Genome sequenced. ||
+| 2019 || || Literature || Daiana S. Avila publishes ''Caenorhabditis Elegans: An Overview and Emerging Roles in Studying Disease'', which examines the use of the nematode ''C. elegans'' as a model organism for studying various diseases. By this time, this nonpathogenic organism had been instrumental in uncovering genetic mechanisms linked to cancer, aging, and nervous system development. The book covers disease models in ''C. elegans'' for aging, metabolic syndrome, neurodegenerative diseases, and chemically induced neurodegeneration. It reviews the creation of transgenic animals mimicking human diseases, advances in molecular understanding, and potential treatments, highlighting both the strengths and limitations of these models.<ref>{{cite book |title=Caenorhabditis Elegans: An Overview of Emerging Systems for Studying Development |url=https://www.amazon.com/Caenorhabditis-Elegans-Overview-Emerging-Studying/dp/1536149462 |publisher=Nova Science Publishers |year=2018 |accessdate=20 October 2024}}</ref> ||
 |-
 |}
+== Numerical and visual data ==
+=== Google trends ===
+The image below shows Google Trends data for model organism (tipic) from January 2004 to October 2024, as well as interest by region.<ref>{{cite web |title=Google Trends: "model organism" |url=https://trends.google.com/trends/explore?date=all&q=%2Fm%2F04xb_&hl=en |website=Google Trends |accessdate=20 October 2024}}</ref>
+[[File:Model-organism-google-trends1.png|thumb|center|900px]]
+=== Google Ngram Viewer ===
+The chart below shows Google Ngram Viewer data for "model organism" from 1800 to 2022.<ref>{{cite web |title=Google Ngram Viewer: "model organism" |url=https://books.google.com/ngrams/graph?content=model+organism&year_start=1800&year_end=2022&corpus=en&smoothing=3# |website=Google Books Ngram Viewer |accessdate=20 October 2024}}</ref>
+[[File:Model-organism-ngram-viewer.png|thumb|center|900px]]
+=== Wikipedia views ===
+The chart below shows pageviews of the English Wikipedia article {{w|Model organism}}, from July 2015 to October 2024, when the screenshot was taken.<ref>{{cite web |title=Pageviews for Model organism on Wikipedia |url=https://wikipediaviews.org/displayviewsformultiplemonths.php?page=Model+organism&allmonths=allmonths-api&language=en&drilldowns[0]=mobile-app&drilldowns[1]=desktop-spider&drilldowns[2]=mobile-web-spider |website=wikipediaviews.org |accessdate=20 October 2024}}</ref>
+[[File:Model-organisms-wikipedia-views.png|thumb|center|600px]]
 ==Meta information on the timeline==
@@ Line 111: / Line 147: @@
 ===How the timeline was built===
-The initial version of the timeline was written by [[User:FIXME|FIXME]].
+The initial version of the timeline was written by [[User:Sebastian|Sebastian]].
 {{funding info}} is available.
@@ Line 122: / Line 158: @@
 ===What the timeline is still missing===
+* Books
+* Recognition. Nobel
+* Species discovery dates
 * Davis, R. H. (2004). The age of model organisms. Nature Reviews Genetics, 5(1), 69–76. doi:10.1038/nrg1250
 * A column for kingdom

Difference between revisions of "Timeline of model organisms"

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