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Experimental evolution is the use of laboratory experiments or controlled field manipulations to explore evolutionary dynamics.
Evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
may be observed in the laboratory as populations adapt to new environmental conditions by
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
.
Adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
can arise in experimental evolution in two different ways. One is via an individual organism gaining a novel beneficial
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
. The other is from
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
frequency change in standing genetic variation already present in a population of organisms. Other evolutionary forces outside of mutation and natural selection can also play a role or be incorporated into experimental evolution studies, such as
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
and
gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
. The organism used is decided by the experimenter, based on the hypothesis to be tested. Many
generation A generation is all of the people born and living at about the same time, regarded collectively. It also is "the average period, generally considered to be about 20–⁠30 years, during which children are born and grow up, become adults, and b ...
s are required for adaptive mutation to occur, and experimental evolution via mutation is carried out in
virus A virus is a submicroscopic infectious agent that replicates only inside the living Cell (biology), cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Viruses are ...
es or
unicellular A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
organisms with rapid generation times, such as
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
and asexual clonal
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
. Polymorphic populations of asexual or sexual
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
, and
multicellular A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...
eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
s like
Drosophila ''Drosophila'' (), from Ancient Greek δρόσος (''drósos''), meaning "dew", and φίλος (''phílos''), meaning "loving", is a genus of fly, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or p ...
, can adapt to new environments through allele frequency change in standing genetic variation. Organisms with longer generations times, although costly, can be used in experimental evolution. Laboratory studies with foxes and with
rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
(see below) have shown that notable adaptations can occur within as few as 10–20 generations and experiments with wild guppies have observed adaptations within comparable numbers of generations. More recently, experimentally evolved individuals or populations are often analyzed using
whole genome sequencing Whole genome sequencing (WGS), also known as full genome sequencing or just genome sequencing, is the process of determining the entirety of the DNA sequence of an organism's genome at a single time. This entails sequencing all of an organism's ...
, an approach known as Evolve and Resequence (E&R). E&R can identify mutations that lead to adaptation in clonal individuals or identify alleles that changed in frequency in polymorphic populations, by comparing the sequences of individuals/populations before and after adaptation. The sequence data makes it possible to pinpoint the site in a
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
sequence that a mutation/allele frequency change occurred to bring about adaptation. The nature of the adaptation and functional follow up studies can shed insight into what effect the mutation/allele has on
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
.


History


Domestication and breeding

Unwittingly, humans have carried out evolution experiments for as long as they have been domesticating plants and animals.
Selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant m ...
of plants and animals has led to varieties that differ dramatically from their original wild-type ancestors. Examples are the
cabbage Cabbage, comprising several cultivars of '' Brassica oleracea'', is a leafy green, red (purple), or white (pale green) biennial plant grown as an annual vegetable crop for its dense-leaved heads. It is descended from the wild cabbage ( ''B.& ...
varieties,
maize Maize (; ''Zea mays''), also known as corn in North American English, is a tall stout grass that produces cereal grain. It was domesticated by indigenous peoples in southern Mexico about 9,000 years ago from wild teosinte. Native American ...
, or the large number of different dog breeds. The power of human breeding to create varieties with extreme differences from a single species was already recognized by
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
. In fact, he started out his book '' The Origin of Species'' with a chapter on variation in domestic animals. In this chapter, Darwin discussed in particular the pigeon.


Early

One of the first to carry out a controlled evolution experiment was William Dallinger. In the late 19th century, he cultivated small
unicellular organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
s in a custom-built incubator over a time period of seven years (1880–1886). Dallinger slowly increased the temperature of the incubator from an initial 60 °F up to 158 °F. The early cultures had shown clear signs of distress at a temperature of 73 °F, and were certainly not capable of surviving at 158 °F. The organisms Dallinger had in his incubator at the end of the experiment, on the other hand, were perfectly fine at 158 °F. However, these organisms would no longer grow at the initial 60 °F. Dallinger concluded that he had found evidence for Darwinian adaptation in his incubator, and that the organisms had adapted to live in a high-temperature environment. Dallinger's incubator was accidentally destroyed in 1886, and Dallinger could not continue this line of research. From the 1880s to 1980, experimental evolution was intermittently practiced by a variety of evolutionary biologists, including the highly influential
Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (; ; January 25, 1900 – December 18, 1975) was a Russian-born American geneticist and evolutionary biologist. He was a central figure in the field of evolutionary biology for his work in shaping the modern ...
. Like other experimental research in evolutionary biology during this period, much of this work lacked extensive replication and was carried out only for relatively short periods of evolutionary time.


Modern

Experimental evolution has been used in various formats to understand underlying evolutionary processes in a controlled system. Experimental evolution has been performed on multicellular and unicellular eukaryotes, prokaryotes, and viruses. Similar works have also been performed by directed evolution of individual
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
,
ribozyme Ribozymes (ribonucleic acid enzymes) are RNA molecules that have the ability to Catalysis, catalyze specific biochemical reactions, including RNA splicing in gene expression, similar to the action of protein enzymes. The 1982 discovery of ribozy ...
and replicator genes.


Aphids

In the 1950s, the Soviet biologist Georgy Shaposhnikov conducted
experiments An experiment is a procedure carried out to support or refute a hypothesis, or determine the efficacy or likelihood of something previously untried. Experiments provide insight into Causality, cause-and-effect by demonstrating what outcome o ...
on aphids of the Dysaphis genus. By transferring them to plants normally nearly or completely unsuitable for them, he had forced populations of parthenogenetic descendants to adapt to the new food source to the point of reproductive isolation from the regular populations of the same species.


Fruit flies

One of the first of a new wave of experiments using this strategy was the laboratory "evolutionary radiation" of ''
Drosophila melanogaster ''Drosophila melanogaster'' is a species of fly (an insect of the Order (biology), order Diptera) in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly", "pomace fly" ...
'' populations that Michael R. Rose started in February, 1980. This system started with ten populations, five cultured at later ages, and five cultured at early ages. Since then more than 200 different populations have been created in this laboratory radiation, with selection targeting multiple characters. Some of these highly differentiated populations have also been selected "backward" or "in reverse," by returning experimental populations to their ancestral culture regime. Hundreds of people have worked with these populations over the better part of three decades. Much of this work is summarized in the papers collected in the book ''Methuselah Flies''. The early experiments in flies were limited to studying phenotypes but the molecular mechanisms, i.e., changes in DNA that facilitated such changes, could not be identified. This changed with genomics technology. Subsequently, Thomas Turner coined the term Evolve and Resequence (E&R) and several studies used E&R approach with mixed success. One of the more interesting experimental evolution studies was conducted by Gabriel Haddad's group at UC San Diego, where Haddad and colleagues evolved flies to adapt to low oxygen environments, also known as hypoxia. After 200 generations, they used E&R approach to identify genomic regions that were selected by natural selection in the hypoxia adapted flies. More recent experiments are following up E&R predictions with RNAseq and genetic crosses. Such efforts in combining E&R with experimental validations should be powerful in identifying genes that regulate adaptation in flies. Much recently the experimental evolution in flies have taken the course to address the molecular mechanisms and in doing so it might pave way to understand physiology of an organism better and thus redefine disease therapeutics.


Microbes

Many microbial species have short
generation time In population biology and demography Demography () is the statistical study of human populations: their size, composition (e.g., ethnic group, age), and how they change through the interplay of fertility (births), mortality (deaths), and mi ...
s, easily sequenced genomes, and well-understood biology. They are therefore commonly used for experimental evolution studies. The bacterial species most commonly used for experimental evolution include ''P. fluorescens'', ''Pseudomonas aeruginosa'', ''Enterococcus faecalis'' and ''E. coli'' (see below), while the Yeast ''S. cerevisiae'' has been used as a model for the study of eukaryotic evolution.


Lenski's ''E. coli'' experiment

One of the most widely known examples of laboratory bacterial evolution is the long-term ''E.coli'' experiment of
Richard Lenski Richard E. Lenski (born 1956) is an American evolutionary biologist who is the John A. Hannah Distinguished Professor of Microbial Ecology at Michigan State University. He is a member of the National Academy of Sciences and a MacArthur Fellow. ...
. On February 24, 1988, Lenski started growing twelve lineages of '' E. coli'' under identical growth conditions. When one of the populations evolved the ability to aerobically metabolize citrate from the growth medium and showed greatly increased growth, this provided a dramatic observation of evolution in action. The experiment continues to this day, and is now the longest-running (in terms of generations) controlled evolution experiment ever undertaken. Since the inception of the experiment, the bacteria have grown for more than 60,000 generations. Lenski and colleagues regularly publish updates on the status of the experiments.


''Leishmania donovani''

Bussotti and collaborators isolated amastigotes from '' Leishmania donovani'' and cultured them in vitro for 3800 generations (36 weeks). The culture of these parasites showed how they adapted to in vitro conditions by compensating for the loss of a NIMA-related kinase, important for the correct progression of mitosis, by increasing the expression of another orthologous kinase as the culture generations progressed. Furthermore, it was observed how ''L. donovani'' has been adapted to in vitro culture by reducing the expression of 23 transcripts related to flagellar biogenesis and increasing the expression of ribosomal protein clusters and non-coding RNAs such as nucleolar small RNAs. Flagella are considered less necessary by the parasite in in vitro culture and therefore the progression of generations leads to their elimination, causing an energy saving due to lower motility so that proliferation and growth rate in culture is higher. The amplified snoRNAs also lead to increased ribosomal biosynthesis, increased protein biosynthesis and thus increased growth rate of the culture. These adaptations observed over generations of parasites are governed by copy number variations (CNV) and epistatic interactions between affected genes, and allow us to justify ''Leishmania'' genomic instability through its post-transcriptional regulation of gene expression.


High Runner laboratory house mice

In 1993, Theodore Garland, Jr. and colleagues started a long-term experiment that involves selective breeding of
mice A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
for high voluntary activity levels on running wheels. This experiment also continues to this day (> 105
generation A generation is all of the people born and living at about the same time, regarded collectively. It also is "the average period, generally considered to be about 20–⁠30 years, during which children are born and grow up, become adults, and b ...
s). Mice from the four replicate "High Runner" lines evolved to run almost three times as many running-wheel revolutions per day compared with the four unselected control lines of mice, mainly by running faster than the control mice rather than running for more minutes/day. However, the High Runner lines have evolved in somewhat different ways, with some emphasizing running speed versus duration or vice versa, thus demonstrating "multiple solutions" that seem to be based partly in evolved muscle characteristics. The HR mice have an elevated endurance running ability and maximal aerobic capacity when tested on a motorized treadmill. They also exhibit alterations in
motivation Motivation is an mental state, internal state that propels individuals to engage in goal-directed behavior. It is often understood as a force that explains why people or animals initiate, continue, or terminate a certain behavior at a particul ...
and the
reward system The reward system (the mesocorticolimbic circuit) is a group of neural structures responsible for incentive salience (i.e., "wanting"; desire or craving for a reward and motivation), associative learning (primarily positive reinforcement and c ...
of the
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
.
Pharmacological Pharmacology is the science of drugs and medications, including a substance's origin, composition, pharmacokinetics, pharmacodynamics, therapeutic use, and toxicology. More specifically, it is the study of the interactions that occur between ...
studies point to alterations in
dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. It is an amine synthesized ...
function and the
endocannabinoid system The endocannabinoid system (ECS) is a biological system composed of endocannabinoids, which are neurotransmitters that bind to cannabinoid receptors, and cannabinoid receptor proteins that are expressed throughout the central nervous system ( ...
. The High Runner lines have been proposed as a model to study human attention-deficit hyperactivity disorder (
ADHD Attention deficit hyperactivity disorder (ADHD) is a neurodevelopmental disorder characterised by symptoms of inattention, hyperactivity, impulsivity, and emotional dysregulation that are excessive and pervasive, impairing in multiple ...
), and administration of Ritalin reduces their wheel running approximately to the levels of control mice.


Multidirectional selection on bank voles

In 2005 Paweł Koteja with Edyta Sadowska and colleagues from the
Jagiellonian University The Jagiellonian University (, UJ) is a public research university in Kraków, Poland. Founded in 1364 by Casimir III the Great, King Casimir III the Great, it is the oldest university in Poland and one of the List of oldest universities in con ...
(Poland) started a multidirectional selection on a non-laboratory rodent, the
bank vole The bank vole (''Clethrionomys glareolus'') is a small vole with red-brown fur and some grey patches, with a tail about half as long as its body. A rodent, it lives in woodland areas and is around in length. The bank vole is found in much of Eu ...
''Myodes (= Clethrionomys) glareolus''. The voles are selected for three distinct traits, which played important roles in the adaptive radiation of terrestrial vertebrates: high maximum rate of aerobic metabolism, predatory propensity, and herbivorous capability. Aerobic lines are selected for the maximum rate of oxygen consumption achieved during swimming at 38°C; Predatory lines – for a short time to catch live crickets; Herbivorous lines – for capability to maintain body mass when fed a low-quality diet “diluted” with dried, powdered grass. Four replicate lines are maintained for each of the three selection directions and another four as unselected Controls. After approximately 20 generations of selective breeding, voles from the Aerobic lines evolved a 60% higher swim-induced metabolic rate than voles from the unselected Control lines. Although the selection protocol does not impose a thermoregulatory burden, both the
basal metabolic rate Basal metabolic rate (BMR) is the rate of energy expenditure per unit time by endothermic animals at rest.. In other words it is the energy required by body organs to perform normal It is reported in energy units per unit time ranging from watt ( ...
and thermogenic capacity increased in the Aerobic lines. Thus, the results have provided some support for the “aerobic capacity model” for the evolution of endothermy in mammals. More than 85% of the Predatory voles capture the crickets, compared to only about 15% of unselected Control voles, and they catch the crickets faster. The increased predatory behavior is associated with a more proactive coping style (“
personality Personality is any person's collection of interrelated behavioral, cognitive, and emotional patterns that comprise a person’s unique adjustment to life. These interrelated patterns are relatively stable, but can change over long time per ...
”). During the test with low-quality diet, the Herbivorous voles lose approximately 2 grams less mass (approximately 10% of the original body mass) than the Control ones. The Herbivorous voles have an altered composition of the bacterial microbiome in their
caecum The cecum ( caecum, ; plural ceca or caeca, ) is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, ...
. Thus, the selection has resulted in evolution of the entire holobiome, and the experiment may offer a laboratory model of hologenome evolution.


Synthetic biology

Synthetic biology Synthetic biology (SynBio) is a multidisciplinary field of science that focuses on living systems and organisms. It applies engineering principles to develop new biological parts, devices, and systems or to redesign existing systems found in nat ...
offers unique opportunities for experimental evolution, facilitating the interpretation of evolutionary changes by inserting genetic modules into host genomes and applying selection specifically targeting such modules. Synthetic biological circuits inserted into the genome of
Escherichia coli ''Escherichia coli'' ( )Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. is a gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Escherichia'' that is commonly fo ...
or the budding yeast
Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungal microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have be ...
degrade (lose function) during laboratory evolution. With appropriate selection, mechanisms underlying the evolutionary regain of lost biological function can be studied. Experimental evolution of mammalian cells harboring synthetic gene circuits reveals the role of cellular heterogeneity in the evolution of drug resistance, with implications for
chemotherapy Chemotherapy (often abbreviated chemo, sometimes CTX and CTx) is the type of cancer treatment that uses one or more anti-cancer drugs (list of chemotherapeutic agents, chemotherapeutic agents or alkylating agents) in a standard chemotherapy re ...
resistance of cancer cells.


Other examples

Stickleback The sticklebacks are a family of ray-finned fishes, the Gasterosteidae which have a Holarctic distribution in fresh, brackish and marine waters. They were thought to be related to the pipefish and seahorses but are now thought to be more close ...
fish have both marine and freshwater species, the freshwater species evolving since the last ice age. Freshwater species can survive colder temperatures. Scientists tested to see if they could reproduce this evolution of cold-tolerance by keeping marine sticklebacks in cold freshwater. It took the marine sticklebacks only three generations to evolve to match the 2.5 degree Celsius improvement in cold-tolerance found in wild freshwater sticklebacks. Microbial cells and recently mammalian cells are evolved under nutrient limiting conditions to study their metabolic response and engineer cells for useful characteristics.


For teaching

Because of their rapid generation times microbes offer an opportunity to study
microevolution Microevolution is the change in allele frequencies that occurs over time within a population. This change is due to four different processes: mutation, selection ( natural and artificial), gene flow and genetic drift. This change happens over ...
in the classroom. A number of exercises involving bacteria and yeast teach concepts ranging from the evolution of resistance to the evolution of multicellularity. With the advent of next-generation sequencing technology it has become possible for students to conduct an evolutionary experiment, sequence the evolved genomes, and to analyze and interpret the results.


For industrial use

Laboratory synthetic evolution, not necessarily directed, is also used to develop new phenotypes in organisms used for biomanufacturing. Natural mutations can be complemented with methods that directly generate phenotypic diversity. With yeast, the generation of new phenotypes can be accelerated by manipulation at the transcription level: by stochastically introducing artificial transcription factors, globally altering the transcription machinery, CRISPR interference and activation. Also with yeast, transposons, loxP sequences, and highly error-prone orthogonal DNAP-DNA plasmid pairs can be used to accelerate the generation of mutations.


See also

*
Artificial selection Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ...
* Bacteriophage experimental evolution * Directed evolution *
Domestication Domestication is a multi-generational Mutualism (biology), mutualistic relationship in which an animal species, such as humans or leafcutter ants, takes over control and care of another species, such as sheep or fungi, to obtain from them a st ...
*
Evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary biolo ...
*
Evolutionary physiology Evolutionary physiology is the study of the biological evolution of physiological structures and processes; that is, the manner in which the functional characteristics of organisms have responded to natural selection or sexual selection or change ...
*
Genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
* Genomics of domestication * Laboratory experiments of speciation *
Quantitative genetics Quantitative genetics is the study of quantitative traits, which are phenotypes that vary continuously—such as height or mass—as opposed to phenotypes and gene-products that are Categorical variable, discretely identifiable—such as eye-col ...
*
Selection limits A selection limit is a term from animal breeding and quantitative genetics that refers to a cessation of progress even when continued directional selection is being applied to a trait, such as body size. In other words, a breeder or scientist i ...
*
Selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant m ...
* Tame Silver Fox


References


Further reading

* * * * * * * * * * *


External links


E. coli Long-term Experimental Evolution Project Site
, Lenski lab,
Michigan State University Michigan State University (Michigan State or MSU) is a public university, public Land-grant university, land-grant research university in East Lansing, Michigan, United States. It was founded in 1855 as the Agricultural College of the State o ...
*
movie
illustrating the dramatic differences in wheel-running behavior.
Experimental Evolution Publications by Ted Garland: Artificial Selection for High Voluntary Wheel-Running Behavior in House Mice
— a detailed list of publications.

— a list of laboratories that study experimental evolution.
Network for Experimental Research on Evolution
University of California The University of California (UC) is a public university, public Land-grant university, land-grant research university, research university system in the U.S. state of California. Headquartered in Oakland, California, Oakland, the system is co ...
. * {{cite news , last1=Nicholls , first1=Henry , title=My little zebra: The secrets of domestication , url=https://www.newscientist.com/article/mg20427281-500-my-little-zebra-the-secrets-of-domestication/ , work=New Scientist , date=30 September 2009 * Inquiry-basedbr>middle school lesson plan: "Born to Run: Artificial Selection Lab"

Digital Evolution for Education software
Evolutionary biology Biology experiments