Biology:Experimental evolution

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Short description: Use of laboratory and field experiments to explore evolutionary dynamics
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Evolutionary biology
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Darwin's finches by John Gould
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Experimental evolution is the use of laboratory experiments or controlled field manipulations to explore evolutionary dynamics.[1] Evolution may be observed in the laboratory as individuals/populations adapt to new environmental conditions by natural selection.

There are two different ways in which adaptation can arise in experimental evolution. One is via an individual organism gaining a novel beneficial mutation.[2] The other is from allele frequency change in standing genetic variation already present in a population of organisms.[2] 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 and gene flow.[3]

The organism used is decided by the experimenter, based on the hypothesis to be tested. Many generations are required for adaptive mutation to occur, and experimental evolution via mutation is carried out in viruses or unicellular organisms with rapid generation times, such as bacteria and asexual clonal yeast.[1][4][5] Polymorphic populations of asexual or sexual yeast,[2] and multicellular eukaryotes like Drosophila, can adapt to new environments through allele frequency change in standing genetic variation.[3] Organisms with longer generations times, although costly, can be used in experimental evolution. Laboratory studies with foxes[6] and with rodents (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.[7]

More recently, experimentally evolved individuals or populations are often analyzed using whole genome sequencing,[8][9] an approach known as Evolve and Resequence (E&R).[10] 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.[2] The sequence data makes it possible to pinpoint the site in a DNA sequence that a mutation/allele frequency change occurred to bring about adaptation.[10][9][2] The nature of the adaptation and functional follow up studies can shed insight into what effect the mutation/allele has on phenotype.

History

Domestication and breeding

This Chihuahua mix and Great Dane show the wide range of dog breed sizes created using artificial selection.

Unwittingly, humans have carried out evolution experiments for as long as they have been domesticating plants and animals. Selective breeding of plants and animals has led to varieties that differ dramatically from their original wild-type ancestors. Examples are the cabbage varieties, maize, 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. 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.

Altogether at least a score of pigeons might be chosen, which if shown to an ornithologist, and he were told that they were wild birds, would certainly, I think, be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail in the same genus; more especially as in each of these breeds several truly-inherited sub-breeds, or species as he might have called them, could be shown him.

(...) I am fully convinced that the common opinion of naturalists is correct, namely, that all have descended from the rock-pigeon (Columba livia), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects.

– Charles Darwin, The Origin of Species

Early

Drawing of the incubator used by Dallinger in his evolution experiments.

One of the first to carry out a controlled evolution experiment was William Dallinger. In the late 19th century, he cultivated small unicellular organisms 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.[11][12]

From the 1880s to 1980, experimental evolution was intermittently practiced by a variety of evolutionary biologists, including the highly influential Theodosius Dobzhansky. 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.[13]

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[14] and unicellular[15] eukaryotes, prokaryotes,[16] and viruses.[17] Similar works have also been performed by directed evolution of individual enzyme,[18][19] ribozyme[20] and replicator[21][22] genes.

Aphids

поколения=generations, Смертность=mortality

In the 1950s, the Soviet biologist Georgy Shaposhnikov conducted experiments 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.[23]

Fruit flies

One of the first of a new wave of experiments using this strategy was the laboratory "evolutionary radiation" of Drosophila melanogaster populations that Michael R. Rose started in February, 1980.[24] 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.[25]

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.[26] Subsequently, Thomas Turner coined the term Evolve and Resequence (E&R)[10] and several studies used E&R approach with mixed success.[27][28] 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.[29] After 200 generations, they used E&R approach to identify genomic regions that were selected by natural selection in the hypoxia adapted flies.[30] More recent experiments are following up E&R predictions with RNAseq[31] and genetic crosses.[9] Such efforts in combining E&R with experimental validations should be powerful in identifying genes that regulate adaptation in flies.

Microbes

Many microbial species have short generation times, 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,[32] Pseudomonas aeruginosa,[33] Enterococcus faecalis [34] and E. coli (see below), while the Yeast S. cerevisiae has been used as a model for the study of eukaryotic evolution.[35]

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. On February 24, 1988, Lenski started growing twelve lineages of E. coli under identical growth conditions.[36][37] When one of the populations evolved the ability to aerobically metabolize citrate from the growth medium and showed greatly increased growth,[38] 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.[citation needed] 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.[39]

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.[40]

Laboratory house mice

Mouse from the Garland selection experiment with attached running wheel and its rotation counter.

In 1998, Theodore Garland, Jr. and colleagues started a long-term experiment that involves selective breeding of mice for high voluntary activity levels on running wheels.[41] This experiment also continues to this day (> 90 generations). 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.

Female mouse with her litter, from the Garland selection experiment.

The HR mice exhibit an elevated maximal aerobic capacity when tested on a motorized treadmill. They also exhibit alterations in motivation and the reward system of the brain. Pharmacological studies point to alterations in dopamine function and the endocannabinoid system.[42] The High Runner lines have been proposed as a model to study human attention-deficit hyperactivity disorder (ADHD), 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 (Poland) started a multidirectional selection on a non-laboratory rodent, the bank vole Myodes (= Clethrionomys) glareolus.[43] 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 and thermogenic capacity increased in the Aerobic lines.[44][45] 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”).[46]

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.[47] 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 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[48] or the budding yeast Saccharomyces cerevisiae[49] degrade (lose function) during laboratory evolution. With appropriate selection, mechanisms underlying the evolutionary regain of lost biological function can be studied.[50] Experimental evolution of mammalian cells harboring synthetic gene circuits[51] reveals the role of cellular heterogeneity in the evolution of drug resistance, with implications for chemotherapy resistance of cancer cells.

Other examples

Stickleback 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.[52]

Microbial cells [53] and recently mammalian cells [54] 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 in the classroom. A number of exercises involving bacteria and yeast teach concepts ranging from the evolution of resistance[55] to the evolution of multicellularity.[56] 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.[57]

See also


References

  1. ↑ 1.0 1.1 "Experimental Evolution". Nature. http://www.nature.com/subjects/experimental-evolution. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 "Elucidating the molecular architecture of adaptation via evolve and resequence experiments". Nature Reviews. Genetics 16 (10): 567–582. October 2015. doi:10.1038/nrg3937. PMID 26347030. 
  3. ↑ 3.0 3.1 "Experimental evolution". Trends in Ecology & Evolution 27 (10): 547–560. October 2012. doi:10.1016/j.tree.2012.06.001. PMID 22819306. https://serval.unil.ch/resource/serval:BIB_23A48B184D98.P001/REF.pdf. 
  4. ↑ "The Beagle in a bottle". Nature 457 (7231): 824–829. February 2009. doi:10.1038/nature07892. PMID 19212400. Bibcode: 2009Natur.457..824B. 
  5. ↑ "Evolution experiments with microorganisms: the dynamics and genetic bases of adaptation". Nature Reviews. Genetics 4 (6): 457–469. June 2003. doi:10.1038/nrg1088. PMID 12776215. 
  6. ↑ "Early Canid Domestication: The Farm-Fox Experiment: Foxes bred for tamability in a 40-year experiment exhibit remarkable transformations that suggest an interplay between behavioral genetics and development.". American Scientist 87 (2): 160–169. March 1999. doi:10.1511/1999.2.160. http://www.americanscientist.org/issues/feature/1999/2/early-canid-domestication-the-farm-fox-experiment/2. 
  7. ↑ "Evaluation of the Rate of Evolution in Natural Populations of Guppies (Poecilia reticulata)". Science 275 (5308): 1934–1937. March 1997. doi:10.1126/science.275.5308.1934. PMID 9072971. 
  8. ↑ "Genome dynamics during experimental evolution". Nature Reviews. Genetics 14 (12): 827–839. December 2013. doi:10.1038/nrg3564. PMID 24166031. 
  9. ↑ 9.0 9.1 9.2 "Whole-Genome Resequencing of Experimental Populations Reveals Polygenic Basis of Egg-Size Variation in Drosophila melanogaster". Molecular Biology and Evolution 32 (10): 2616–2632. October 2015. doi:10.1093/molbev/msv136. PMID 26044351. 
  10. ↑ 10.0 10.1 10.2 "Population-based resequencing of experimentally evolved populations reveals the genetic basis of body size variation in Drosophila melanogaster". PLOS Genetics 7 (3): e1001336. March 2011. doi:10.1371/journal.pgen.1001336. PMID 21437274. 
  11. ↑ "The Reverend Dr William Henry Dallinger, F.R.S. (1839-1909)". Notes and Records of the Royal Society of London 54 (1): 53–65. January 2000. doi:10.1098/rsnr.2000.0096. PMID 11624308. 
  12. ↑ Darwin Under the Microscope: Witnessing Evolution in Microbes. W. H. Freeman. 2011. pp. 42–43. ISBN 978-0981519494. https://ncse.ngo/files/pub/evolution/Excerpt--lightofevolution.pdf. 
  13. ↑ "An experimental study of interaction between genetic drift and natural selection". Evolution 11 (3): 311–319. 1957. doi:10.2307/2405795. 
  14. ↑ "Aerial performance of Drosophila melanogaster from populations selected for upwind flight ability". The Journal of Experimental Biology 200 (Pt 21): 2747–2755. November 1997. doi:10.1242/jeb.200.21.2747. PMID 9418031. 
  15. ↑ "Experimental evolution of multicellularity". Proceedings of the National Academy of Sciences of the United States of America 109 (5): 1595–1600. January 2012. doi:10.1073/pnas.1115323109. PMID 22307617. Bibcode: 2012PNAS..109.1595R. 
  16. ↑ "Genome evolution and adaptation in a long-term experiment with Escherichia coli". Nature 461 (7268): 1243–1247. October 2009. doi:10.1038/nature08480. PMID 19838166. Bibcode: 2009Natur.461.1243B. 
  17. ↑ "Evolutionary robustness of an optimal phenotype: re-evolution of lysis in a bacteriophage deleted for its lysin gene". Journal of Molecular Evolution 61 (2): 181–191. August 2005. doi:10.1007/s00239-004-0304-4. PMID 16096681. Bibcode: 2005JMolE..61..181H. 
  18. ↑ "In the light of directed evolution: pathways of adaptive protein evolution". Proceedings of the National Academy of Sciences of the United States of America 106 (Suppl 1): 9995–10000. June 2009. doi:10.1073/pnas.0901522106. PMID 19528653. 
  19. ↑ "In vitro evolution goes deep". Proceedings of the National Academy of Sciences of the United States of America 108 (20): 8071–8072. May 2011. doi:10.1073/pnas.1104843108. PMID 21551096. Bibcode: 2011PNAS..108.8071M. 
  20. ↑ "In vitro evolution suggests multiple origins for the hammerhead ribozyme". Nature 414 (6859): 82–84. November 2001. doi:10.1038/35102081. PMID 11689947. Bibcode: 2001Natur.414...82S. 
  21. ↑ "Evidence for de novo production of self-replicating and environmentally adapted RNA structures by bacteriophage Qbeta replicase". Proceedings of the National Academy of Sciences of the United States of America 72 (1): 162–166. January 1975. doi:10.1073/pnas.72.1.162. PMID 1054493. Bibcode: 1975PNAS...72..162S. 
  22. ↑ "An extracellular Darwinian experiment with a self-duplicating nucleic acid molecule". Proceedings of the National Academy of Sciences of the United States of America 58 (1): 217–224. July 1967. doi:10.1073/pnas.58.1.217. PMID 5231602. Bibcode: 1967PNAS...58..217M. 
  23. ↑ "Origin and breakdown of reproductive isolation and the criterion of the species.". Entomological Review 45: 1–8. 1966. http://rogov.zwz.ru/Macroevolution/epi17.pdf. 
  24. ↑ "Artificial Selection on a Fitness-Component in Drosophila Melanogaster". Evolution; International Journal of Organic Evolution 38 (3): 516–526. May 1984. doi:10.2307/2408701. PMID 28555975. 
  25. ↑ Methuselah Flies. Singapore: World Scientific. 2004. doi:10.1142/5457. ISBN 978-981-238-741-7. 
  26. ↑ "Genome-wide analysis of a long-term evolution experiment with Drosophila". Nature 467 (7315): 587–590. September 2010. doi:10.1038/nature09352. PMID 20844486. Bibcode: 2010Natur.467..587B. 
  27. ↑ "Sequencing pools of individuals - mining genome-wide polymorphism data without big funding". Nature Reviews. Genetics 15 (11): 749–763. November 2014. doi:10.1038/nrg3803. PMID 25246196. 
  28. ↑ "Combining experimental evolution with next-generation sequencing: a powerful tool to study adaptation from standing genetic variation". Heredity 114 (5): 431–440. May 2015. doi:10.1038/hdy.2014.86. PMID 25269380. 
  29. ↑ "Experimental selection for Drosophila survival in extremely low O(2) environment". PLOS ONE 2 (5): e490. May 2007. doi:10.1371/journal.pone.0000490. PMID 17534440. Bibcode: 2007PLoSO...2..490Z. 
  30. ↑ "Experimental selection of hypoxia-tolerant Drosophila melanogaster". Proceedings of the National Academy of Sciences of the United States of America 108 (6): 2349–2354. February 2011. doi:10.1073/pnas.1010643108. PMID 21262834. Bibcode: 2011PNAS..108.2349Z. 
  31. ↑ "Genomic basis of aging and life-history evolution in Drosophila melanogaster". Evolution; International Journal of Organic Evolution 66 (11): 3390–3403. November 2012. doi:10.1111/j.1558-5646.2012.01710.x. PMID 23106705. 
  32. ↑ "Adaptive radiation in a heterogeneous environment". Nature 394 (6688): 69–72. July 1998. doi:10.1038/27900. PMID 9665128. Bibcode: 1998Natur.394...69R. 
  33. ↑ "Reactive oxygen species drive evolution of pro-biofilm variants in pathogens by modulating cyclic-di-GMP levels". Open Biology 6 (11): 160162. November 2016. doi:10.1098/rsob.160162. PMID 27881736. 
  34. ↑ "No collateral antibiotic sensitivity by alternating antibiotic pairs" (in English). The Lancet Microbe 3 (1): e7. 2021-11-15. doi:10.1016/S2666-5247(21)00270-6. ISSN 2666-5247. PMID 35544116. 
  35. ↑ "Adaptive radiation in a heterogeneous environment". Nature 394 (6688): 69–72. July 1998. doi:10.1038/nature12344. PMID 9665128. Bibcode: 2013Natur.500..571L. 
  36. ↑ "Long-Term Experimental Evolution in Escherichia coli. I. Adaptation and Divergence During 2,000 Generations". The American Naturalist 138 (6): 1315–1341. 1991-12-01. doi:10.1086/285289. ISSN 0003-0147. 
  37. ↑ "From Here to Eternity--The Theory and Practice of a Really Long Experiment". PLOS Biology 13 (6): e1002185. June 2015. doi:10.1371/journal.pbio.1002185. PMID 26102073. 
  38. ↑ "Historical contingency and the evolution of a key innovation in an experimental population of Escherichia coli". Proceedings of the National Academy of Sciences of the United States of America 105 (23): 7899–7906. June 2008. doi:10.1073/pnas.0803151105. PMID 18524956. Bibcode: 2008PNAS..105.7899B. 
  39. ↑ "E. coli Long-term Experimental Evolution Project Site". Michigan State University. http://myxo.css.msu.edu/ecoli/. 
  40. ↑ Bussotti, Giovanni; Piel, Laura; Pescher, Pascale; Domagalska, Malgorzata A.; Rajan, K. Shanmugha; Cohen-Chalamish, Smadar; Doniger, Tirza; Hiregange, Disha-Gajanan et al. (21 December 2021). "Genome instability drives epistatic adaptation in the human pathogen Leishmania" (in en). Proceedings of the National Academy of Sciences 118 (51): e2113744118. doi:10.1073/pnas.2113744118. ISSN 0027-8424. PMID 34903666. Bibcode: 2021PNAS..11813744B. 
  41. ↑ "Artificial selection for increased wheel-running behavior in house mice". Behavior Genetics 28 (3): 227–237. May 1998. doi:10.1023/a:1021479331779. PMID 9670598. http://www.biology.ucr.edu/people/faculty/Garland/SwEA98SE.pdf. 
  42. ↑ "Differential response to a selective cannabinoid receptor antagonist (SR141716: rimonabant) in female mice from lines selectively bred for high voluntary wheel-running behaviour". Behavioural Pharmacology 19 (8): 812–820. December 2008. doi:10.1097/FBP.0b013e32831c3b6b. PMID 19020416. 
  43. ↑ "Laboratory model of adaptive radiation: a selection experiment in the bank vole". Physiological and Biochemical Zoology 81 (5): 627–640. 2008. doi:10.1086/590164. PMID 18781839. 
  44. ↑ "Evolution of basal metabolic rate in bank voles from a multidirectional selection experiment". Proceedings. Biological Sciences 282 (1806): 20150025. May 2015. doi:10.1098/rspb.2015.0025. PMID 25876844. 
  45. ↑ "The effect of chlorpyrifos on thermogenic capacity of bank voles selected for increased aerobic exercise metabolism". Chemosphere 149: 383–390. April 2016. doi:10.1016/j.chemosphere.2015.12.120. PMID 26878110. Bibcode: 2016Chmsp.149..383D. 
  46. ↑ "Experimental evolution of personality traits: open-field exploration in bank voles from a multidirectional selection experiment". Current Zoology 65 (4): 375–384. August 2019. doi:10.1093/cz/zoy068. PMID 31413710. 
  47. ↑ "Experimental Evolution on a Wild Mammal Species Results in Modifications of Gut Microbial Communities". Frontiers in Microbiology 7: 634. 2016. doi:10.3389/fmicb.2016.00634. PMID 27199960. 
  48. ↑ "Designing and engineering evolutionary robust genetic circuits". Journal of Biological Engineering 4: 12. November 2010. doi:10.1186/1754-1611-4-12. PMID 21040586. 
  49. ↑ "Stress-response balance drives the evolution of a network module and its host genome". Molecular Systems Biology 11 (8): 827. August 2015. doi:10.15252/msb.20156185. PMID 26324468. 
  50. ↑ "Evolutionary regain of lost gene circuit function". Proceedings of the National Academy of Sciences 116 (50): 25162–25171. December 2019. doi:10.1073/pnas.1912257116. PMID 31754027. 
  51. ↑ "Role of network-mediated stochasticity in mammalian drug resistance". Nature Communications 10 (1): 2766. June 2019. doi:10.1038/s41467-019-10330-w. PMID 31235692. 
  52. ↑ "Rapid evolution of cold tolerance in stickleback". Proceedings. Biological Sciences 278 (1703): 233–238. January 2011. doi:10.1098/rspb.2010.0923. PMID 20685715. 
  53. ↑ "Adaptive laboratory evolution -- principles and applications for biotechnology". Microbial Cell Factories 12 (1): 64. July 2013. doi:10.1186/1475-2859-12-64. PMID 23815749. 
  54. ↑ "CHO Cells adapted to inorganic phosphate limitation show higher growth and higher pyruvate carboxylase flux in phosphate replete conditions". Biotechnology Progress 33 (3): 749–758. May 2017. doi:10.1002/btpr.2450. PMID 28220676. 
  55. ↑ "Bacteriophage as instructional organisms in introductory biology labs". Bacteriophage 4 (1): e27336. January 2014. doi:10.4161/bact.27336. PMID 24478938. 
  56. ↑ "A Novel Laboratory Activity for Teaching about the Evolution of Multicellularity". The American Biology Teacher 76 (2): 81–87. 2014. doi:10.1525/abt.2014.76.2.3. ISSN 0002-7685. 
  57. ↑ "Using experimental evolution and next-generation sequencing to teach bench and bioinformatic skills". PeerJ PrePrints (3): e1674. 2015. doi:10.7287/peerj.preprints.1356v1. https://peerj.com/preprints/1356/?td=wk. 

Further reading

External links



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