Timeline of model organisms

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This is a timeline of 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. Throughout history, these organisms have been 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:

  • How did early dissections of animals and humans shape the use of model organisms in biological research?
    • Sort the full timeline by "Event type" and look for the group of rows with value "Early development".
    • You will see a summary of early milestones in the use of model organisms, all contributing to the development of anatomy and biological research methodologies.
  • How did early advancements in diverse sciences contribute to the foundation of model organism research?
    • Sort the full timeline by "Event type" and look for the group of rows with value "Scientific bakground".
    • You will see some key milestones in early biological research, having shaped taxonomy, microbiology, and comparative anatomy, influencing modern scientific approaches and model organism research.
  • What are some key events in the introduction of model organisms for genetic and biological research throughout history?
  • How have model organisms contributed to our understanding of genetics?
    • Sort the full timeline by "Event type" and look for the group of rows with value "Genetic research".
    • You will see a number of significant advancements in genetic research through the study of model organisms.
  • What are some key books that have advanced our understanding of genetic research using model organisms?
    • Sort the full timeline by "Event type" and look for the group of rows with value "Literature".
    • You will see some key publications and their contributions to genetic research, such as books that advanced understanding of model organisms in genetics, drug discovery, human genetic disorders, and emerging model organisms, with their influence on scientific progress.
  • What are some key milestones in the sequencing of genomes of model organisms, and how have they contributed to advancements in genetics and biomedical research?
    • You will see a summary of significant milestones in genome sequencing of model organisms, such as Drosophila melanogaster, Saccharomyces cerevisiae, and others. Each entry describes the organism, the year of sequencing, key findings, and their contributions to genetics and biomedical research.

Big picture

Time period Development summary More details
Ancient times – 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 – 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.[2] This period also sees early embryological studies using frogs and chickens, as researchers begin to explore developmental biology in a more structured way.[3]
Mid-20th century – late 20th century Rise of molecular biology The mid-20th century brings revolutionary advancements in molecular biology, with model organisms playing central roles.[4] Bacteria like Escherichia coli becomes vital for studying DNA replication, gene expression, and molecular genetics,[5] while Caenorhabditis elegans provides insights into developmental biology and programmed cell death.Cite error: Invalid parameter in <ref> tag 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.[6]
Late 20th century – 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.[7][8] New technologies like CRISPR-Cas9 revolutionize gene editing, allowing scientists to modify genes with precision in organisms like zebrafish and mice.[9][10][11][12] 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.[13][14] 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.

Overview of principal model organisms

Model organism First pioneering use Key atributes Target organisms
Escherichia coli (E. coli) Fast growth, simple genome, easy genetic manipulation, well-understood biochemistry.[15] Suitable for biochemical and genetic research, ease of cultivation, and simpler structure compared to eukaryotes.
Saccharomyces cerevisiae (yeast) 1950s–1960s (genetic studies)[16]; 1970s (cell cycle research)[17] Eukaryotic, unicellular, simple to culture, genetic tools available, shares cellular machinery with animals Eukaryotes, especially unicellular fungi

Full timeline

Inclusion criteria

We include:

  • Significant, foundational introductions of model organisms that later became widely adopted for research in genetics, biology, or other fields.
  • Key discoveries, technological advances, and influential research that established organisms as model systems or significantly advanced scientific understanding.
  • Major publications that significantly advanced the use of model organisms in genetics, research, and drug discovery, as well as foundational works.

We exclude:

  • Findings that were later disproven, irreproducible, or had minimal impact on the organism's use in research, as well as peripheral studies that didn’t directly contribute to model organism status.
  • Publications that are less impactful, cover niche topics, or do not significantly influence the broader field.
  • Organizational foundations.

Timeline

Year Species Event type Details Location
384–322 BC Early development Ancient Greek philosopher Aristotle conducts extensive anatomical studies on various animals, pioneering the field of comparative biology. Through detailed dissections and observations, he classifies organisms based on shared characteristics, distinguishing between vertebrates and invertebrates. His work explores physiological structures, reproduction, and adaptation. Aristotle’s empirical approach would influence later biological thought, laying the groundwork for taxonomy and evolutionary studies.[18] Ancient Greece
325-250 BC Homo sapiens Early development Greek physician Herophilos (later known as the "Father of Anatomy") is credited as the first to systematically dissect and analyze the human body, laying the foundation for future anatomical studies. Arguably most early work in dissection is attributed to Herophilus. He develops techniques for conducting human dissections and provides detailed, accurate descriptions of human organs. Additionally, Herophilus categorizes the human body into functional systems, greatly advancing the understanding of anatomy and influencing medical knowledge for centuries.[19] Ancient Greece
129–216 CE Apes, pigs, goats, etc Early development Galen, a Greek physician in the Roman Empire, extensively uses animals, particularly apes, pigs, and goats, as model organisms to study anatomy, physiology, and pharmacology. Due to limited access to human cadavers, he performs dissections and vivisections on animals, believing that their anatomy could provide insights into human biology. Galen’s work contributes significantly to early medical knowledge, particularly in understanding the circulatory and nervous systems. His findings would form the foundation of medical teachings in Europe for over a millennium. However, his reliance on animal dissections leads to anatomical inaccuracies, which would be later corrected during the Renaissance.[20][21] Roman Empire
1543 Homo sapiens Early development Flemish physician Andreas Vesalius publishes De Humani Corporis Fabrica Libri Septem, a groundbreaking work on human anatomy. Vesalius revolutionizes the study of anatomy by using human cadavers for dissection, challenging previous reliance on animal models, such as apes and pigs, which had been used by earlier anatomists like Galen. His detailed and accurate illustrations of the human body corrects many anatomical errors and lays the foundation for modern anatomy. Vesalius's emphasis on direct observation of human specimens marks a significant shift in scientific methodology, promoting the importance of using model organisms that closely resemble the species being studied.[22] Belgium
1674 Scientific background Dutch microbiologist and microscopist Antonie van Leeuwenhoek becomes the first to observe red blood cells and protozoa using his handcrafted microscopes. Two years later, in 1676, at the age of 44, he identifies bacteria and sperm cells from an animal’s testes. His discoveries revolutionize microbiology by revealing microscopic life forms previously unknown to science. These early observations lay the groundwork for the study of model organisms, as protozoa, bacteria, and spermatozoa would become fundamental subjects in biological research.[23] Netherlands
1735 Scientific background Swedish biologist and physician Carl Linnaeus publishes Systema Naturae, a seminal work that lays the foundation for modern taxonomy. This publication introduces binomial nomenclature, a system of naming species using two Latin names: the genus and the species. This standardized method allows for clear identification and classification of organisms, replacing previous, more chaotic naming systems. Linnaeus' work would revolutionize biology, providing a universal framework for naming and organizing life on Earth, which would remain in use until today.[24] Sweden
1800 Scientific background Karl Friedrich Burdach coins the terms "morphology" and "biology," marking significant contributions to the study of anatomy and natural sciences. He is influenced by Romantic philosophy and Naturphilosophie, advancing neuroanatomy, particularly through research on the brain's evolutionary history. Burdach's focus includes the anatomy of the brain and the 5th and 7th cranial nerves. His contributions would shape the fields of biology, physiology, and medical education, influencing future developments in the sciences.[25] Germany
1822 Danio rerio Model introduction 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.[26] United Kingdom
1833 Chlamydomonas reinhardtii Model introduction 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.[27] Kingdom of Prussia
1853 Pisum sativum Genetic research Austrian biologist 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 counts and analyzes his results, identifying consistent patterns of trait transmission. His findings lead 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.[28][29][30][31] In 1900, Mendel's laws would be rediscovered by Hugo de Vries, Carl Correns, and Erich von Tschermak.[32][33] Austria
1856 Rattus norvegicus Model standardization The first documented scientific study involving rats is published in France, marking a pivotal moment in the history of biomedical research. The study focuses on the effects of adrenalectomy (removal of adrenal glands) in rats and represents the earliest formal use of Rattus norvegicus as a laboratory model organism. This event lays the groundwork for the rat’s emergence as a preferred species in experimental biology due to its physiological similarity to humans, ease of handling, and rapid breeding. Over time, rats became widely used in controlled laboratory settings, influencing diverse fields such as physiology, toxicology, and behavioral science.[34] France
1857 Saccharomyces cerevisiae Model introduction 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.[35][36] France
1865 Pisum sativum Milestone research Gregor Mendel presents his research on plant hybridization to the Natural History Society of Brno. His findings form the basis for modern genetics by revealing how traits are inherited across generations. Mendel's experiments with pea plants (Pisum sativum) provide critical insights into the principles of inheritance, such as the laws of segregation and independent assortment. These principles are fundamental to genetics research and would influence the use of Pisum sativum as a model organism for studying genetic patterns in plants.[37] Austrian Empire
1885 Frog Milestone research German zoologist Wilhelm Roux publishes his Contributions to the Developmental Mechanics of the Embryo, a foundational work in experimental embryology. Roux introduces the concept of developmental mechanics (Entwicklungsmechanik), which emphasizes studying embryonic development through controlled experiments rather than just observation. In the book, Roux describes his famous frog embryo experiments, where he uses a heated needle to kill one cell of a two-cell embryo. He observes that the surviving cell develops into only half an embryo, supporting the idea of mosaic development—suggesting that embryonic cells have predetermined fates. His work lays the groundwork for modern developmental biology.[38] Germany
1900 Zea mays Milestone research 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.[39] German Empire
1902 Mus musculus Model introduction 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 are instrumental in the development of mouse models for genetic research, which would continue to be crucial in biomedical research and the study of human genetics.[40][41] United States
1909 Drosophila melanogaster Model introduction 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 breeds Drosophila in large quantities, facilitating the analysis of spontaneous mutations and the localization of genes. Morgan's work lays the foundation for understanding the linear arrangement of genes in chromosomes and significantly advances the field of genetics.[42] United States (Columbia University)
1909 Mus musculus (lab strains) Model standardization American scientist C. C. Little develops the first inbred mouse strain, known as DBA, to reduce genetic variability in research. This marks a pivotal moment in standardizing animal models for genetic studies. The success of DBA leads to the creation of many other inbred strains, which would become essential for ensuring reproducibility in mammalian genetic research. By using genetically uniform mice, scientists can reliably compare experimental results across different laboratories, advancing consistency and precision in biomedical research.[43][44] United States
1913 Zea mays Milestone 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.[45] United States
1915 Drosophila melanogaster Literature 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.[46] United States
1927 Neurospora crassa Genetic research 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.[47][48][49] United States
1930 Chlamydomonas Genetic 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.[50] Germany
1935 Saccharomyces cerevisiae Model introduction 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 lays the foundation for subsequent studies in yeast genetics, which would be later pursued by researchers such as Lindegren in the U.S. and Ephrussi in France.[51][52] Denmark
1937 Paramecium Genetic research 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.[53][54][55] United States
1939 Enterobacteria phage T2 Milestone research 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.[56][57][58] United States
1941 Neurospora crassa Milestone research 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.[59][60][61] 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.[62] Germany
1943 Saccharomyces cerevisiae Milestone research 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.[63][64][65] United States
1946 Escherichia coli Milestone research 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.[66][67][68][69][70] United States
1949 Saccharomyces cerevisiae Milestone research 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.[71] France
1950 Zea mays Milestone research 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.[72][73][74][75] United States
1951 Escherichia virus Lambda Milestone research 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.[76] United States
1952 Salmonella typhimurium Milestone research 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.[77][78][79][80][81] United States
1953 Tetrahymena thermophila Genetic research 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.[82] 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.[83] United Kingdom
1954 Neurospora crassa Literature 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.[84] 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.[85] 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.[86][87] United Kingdom
1966 Homo sapiens 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.[88] United States
1972 Scientific background The development of recombinant DNA technology in the United States revolutionizes genetics by allowing scientists to manipulate and combine DNA from different organisms. This breakthrough lays the foundation for creating genetically modified organisms (GMOs) and advanced research in molecular biology. The technology greatly enhances the use of model organisms, particularly bacteria and mice, for studying genetics, gene expression, and disease mechanisms. It provides new tools for exploring gene functions, advancing medical research, and improving therapeutic strategies, further solidifying model organisms' crucial role in scientific discovery and biomedical progress.[89][90] United States
1974 Caenorhabditis elegans Milestone research 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.[91][92][93][94][95][96] United Kingdom
1976 Mus musculus Drug screening The National Cancer Institute (NCI) launches the Developmental Therapeutics Program (DTP), formalizing the use of inbred and immunocompromised mice—particularly athymic nude mice—for systematic in vivo screening of anti-cancer compounds. This initiative institutionalizes the laboratory mouse as the gold standard for oncology drug evaluation. The program's tumor panel includes both murine and human tumor models, enabling more predictive assessments of clinical efficacy. The DTP significantly expands drug screening capacity and leads to the discovery of key agents like paclitaxel. It marks a turning point in preclinical cancer research by anchoring mouse-based models in federal drug development protocols.[97][98][99] United States
1980 Drosophila melanogaster Milestone research 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.[100] Germany
1981 Danio rerio Milestone research 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.[101][102] United States
1991 Caenorhabditis elegans Drug screening Caenorhabditis elegans is first systematically used in pharmacological assays to study neuromuscular function, marking a foundational moment in its role as a model for neuroactive drug screening. Researchers employ compounds such as levamisole, a nicotinic acetylcholine receptor agonist, and aldicarb, an acetylcholinesterase inhibitor, to investigate synaptic transmission at the neuromuscular junction. These studies revealed that behavioral responses to these agents could be used to identify mutants with altered neurotransmission, establishing C. elegans as a powerful system for linking drug effects to genetic pathways and neuronal function.[103][104]
1984 Arabidopsis thaliana Genetic research 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.[105][106][107][108][109] United States
1984 Danio rerio Drug screening Scientists begin using zebrafish embryos to study developmental toxicity due to their transparent bodies and external fertilization, which allow for direct observation of embryonic development. This approach enables high-throughput chemical screening, making it possible to quickly and efficiently test large numbers of compounds for harmful effects on development. Compared to traditional mammalian models, zebrafish offer a faster, cost-effective alternative, enhancing the ability to identify potential teratogens and toxic substances. The model would since become a valuable tool in toxicology and drug safety assessment.[110][111][112] United States
1996 (April) Saccharomyces cerevisiae Milestone research 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.[113][114][115][116][117] North America, Japan, Europe
1996 Danio rerio Milestone research 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, with the zebrafish emerging 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.[118][119][120][121] Germany, United States
1997 Escherichia coli Milestone research 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.[122] United States
1998 Caenorhabditis elegans Milestone research 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.[123][124][125] United States, United Kingdom
2000 (December) Arabidopsis thaliana Milestone research 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.[126][127][128] United States
2000 Drosophila melanogaster Milestone research 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.[129][130][131] United States
2001 Human Milestone research 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.[132][133][134] United States
2002 Milestone research 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.[135][136][137] United States
2002 Mouse (B6-Ins2Akita/+) Model introduction A study introduces B6-Ins2Akita/+ mice as a new model for studying islet transplantation in insulin-dependent diabetes. Unlike traditional diabetic rodent models that rely on toxic chemicals to induce hyperglycemia, these mice develop diabetes spontaneously due to a mutation that impairs insulin secretion. The mice are insulin-sensitive and respond well to islet transplantation. Transplants with syngeneic islets reverse hyperglycemia within 72 hours, and euglycemia is maintained until the graft is removed. Allogeneic grafts are rejected, demonstrating immune competence. This model avoids beta-cell autoimmunity and may be valuable for testing human islets and beta-cell replacement therapies, including stem cell-based approaches.[138] United States
2003 Neurospora crassa Milestone research 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.[139] 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.[140]
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.[141] United States
2006 Danio rerio Drug screening Large-scale chemical screens in zebrafish identify small molecules that modulate vertebrate development. Zebrafish are recognized as a cost-effective model for in vivo drug discovery and toxicology. Germany, United States
2007 Chlamydomonas reinhardtii Milestone research 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.[142]
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.[143]
2011 Mus musculus Protocol evaluation The U.S. Food and Drug Administration (FDA) endorses the use of genetically modified mouse models that express humanized drug-metabolizing enzymes. This strategy addresses limitations in traditional mouse models, which often fail to accurately predict human responses due to species-specific differences in drug metabolism. By incorporating human enzymes into mice, researchers can more reliably simulate human pharmacokinetics and assess drug-drug interactions during preclinical testing. These humanized models improve the translation of laboratory findings to clinical outcomes, enhancing the drug development process and supporting safer, more effective therapies for humans.[144] United States
2011 Mus musculus, Danio rerio Protocol evaluation The Knockout Mouse Phenotyping Project (KOMP2) is launched as a trans-NIH and Common Fund initiative to systematically study the function of every protein-coding gene shared by humans and mice. Building on the earlier Knockout Mouse Project, KOMP2 expands efforts using embryonic stem cells and CRISPR to produce and phenotype over 5,500 null mutant mouse lines. The project prioritizes poorly understood genes (the “dark genome”) and supports research into gene function, disease, and therapy. KOMP2 collaborates with major NIH programs, enabling discoveries across pediatric disease, druggability, and undiagnosed disorders, with data used in over 7,000 scientific publications.[145] United States
2012 Danio rerio Protocol evaluation This study outlines a high-throughput screening protocol based on in situ hybridization (ISH) that leverages zebrafish embryos as a model organism for early-stage drug discovery. Zebrafish serve as an effective vertebrate model due to their conserved developmental pathways, optical transparency, and ability to absorb small molecules directly from water, enabling whole-organism assessment of compound activity. The protocol involves embryo collection, chemical treatment, fixation, and ISH staining to visualize gene expression. By using zebrafish, researchers can investigate biological effects in a living vertebrate system, allowing for the identification of compounds—such as dmPGE2 and leflunomide—that have proceeded to clinical evaluation. This organism-based method enhances the physiological relevance of screening and improves detection of toxicity and pathway-specific responses that may be overlooked in cell-based systems.[146] United States
2016 Nothobranchius furzeri Drug screening African turquoise killifish (Nothobranchius furzeri) emerges as a powerful vertebrate model for ageing research due to its exceptionally short lifespan (4–9 months) and physiological similarities to humans. It exhibits key ageing traits—such as neurodegeneration, cancer, and telomere attrition—making it suitable for studying age-related diseases. The species is genetically tractable with efficient tools like CRISPR/Cas9, enabling rapid generation of transgenic lines. Its life cycle allows fast, large-scale screening of ageing-related genes and interventions, including dietary restriction and drug testing. As a bridge between short-lived invertebrate models and long-lived mammals, it offers unique advantages for uncovering mechanisms of vertebrate ageing.[147] Germany, South Korea
2017 Drosophila melanogaster Drug screening Scientists increasingly use Drosophila melanogaster (fruit flies) as models to study rare and undiagnosed human diseases, especially in pediatric patients. The fruit fly's well-understood genetics, short life cycle, and ease of genetic manipulation makes it ideal for rapidly screening large numbers of potential drug therapies. This approach allows researchers to model human disease mutations in flies and identify promising treatment candidates more efficiently than traditional methods. The strategy would contribute to accelerating drug discovery and expanding the understanding of rare genetic conditions using a cost-effective and scalable model organism.[148][149][150][151] United States
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.[152]

Numerical and visual data

The image below shows Google Trends data for model organism (tipic) from January 2004 to October 2024, as well as interest by region.[153]

Google Ngram Viewer

The chart below shows Google Ngram Viewer data for "model organism" from 1800 to 2022.[154]

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.[155]

Meta information on the timeline

How the timeline was built

The initial version of the timeline was written by Sebastian.

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See also

References

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  2. ↑ Elof Axel Carlson (July–September 2013). "How fruit flies came to launch the chromosome theory of heredity". Mutation Research/Reviews in Mutation Research. 753 (1): 1–6. doi:10.1016/j.mrrev.2013.03.001. Retrieved 7 February 2025.
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