Abstract
LONG BEFORE it was given such a name, experimental evolution took place during the serial passages of pathogenic viruses and bacteria necessary to develop live attenuated vaccines (Kawecki et al. 2012). The field of experimental evolution as we know it now, however, has developed rapidly and dramatically during the past three decades. Probably the best known experimental evolution study is Richard Lenski’s long-term evolution experiment (LTEE) at Michigan State University. Starting with twelve replicate E. coli cultures in February 1988, the experiment reached an astonishing 66,000 generations in November 2016, and (of course) continues. Using a strain that lacks a mechanism for genetic exchange, any changes seen in the twelve lines is a reflection of mutations alone, under the action of selection.
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Notes
- 1.
Lenski’s lab typically freezes samples of every 500th generation at −80 °C. Unfrozen “fossil” samples are allowed one day to thaw and acclimate to experimental conditions before being used for comparisons of fitness, cell size, or other assays.
- 2.
In clonal interference, competition occurs among organisms with different beneficial mutations, which makes it more difficult for the beneficial mutations to spread throughout the population; this is a particular issue in asexual populations, which do not undergo genetic recombination, rendering the beneficial mutations incapable of “joining forces”.
- 3.
In diminishing-returns epistasis, “the marginal improvement from a beneficial mutation declines with increasing fitness” (Wiser et al. 2013).
- 4.
The DM25 medium contained 139 μM glucose, serving as a carbon source for non-Cit+ E. coli.
- 5.
When establishing the LTEE in 1988, Lenski and colleagues used twelve E. coli strains derived from a single clone, identical except for their ability to utilize arabinose. Six strains had a mutation that allowed them to utilize arabinose (strains Ara+1, Ara+2, etc.…), and six did not (Ara-1, Ara-2, etc.…) This mutation had no effect on the strains under the conditions of the LTEE, which did not include arabinose in the DM25 medium, but did allow the strains to be distinguished from each other when grown on tetrazolium-arabinose plates, where the Ara− cells make red colonies and the Ara+ cells make colonies that appear white.
- 6.
Cit+ cells never completely dominated the population; about 1% of the population retained the Cit− phenotype when Cit+ and Cit− cells were grown together. As Blount et al. (2008) observed, “[a]lthough the Cit+ cells continued to use glucose, they did not drive the Cit− subpopulation extinct because the Cit− cells were superior competitors for glucose. Thus, the overall diversity increased as one population gave rise evolutionarily to an ecological community with two members, one a resource specialist and the other a generalist.”
- 7.
Ara-3 was not one of the populations that had evolved a hypermutator phenotype!
- 8.
Note that “older” means later in the lineage, not “more ancestral”.
- 9.
Blount et al. note that “[c]omparative studies have shown that gene duplications have an important creative role in evolution by generating redundancies that allow neo-functionalization… Our findings highlight the less-appreciated capacity of duplications to produce new functions by promoter capture events that change gene regulatory networks.”
- 10.
Relative fitness was measured as the ratio of doublings of the competing genotypes, using a method described by Lenski and colleagues in the context of the LTEE (Lenski et al. 1991).
- 11.
Even colonies without cheaters collapse eventually, when the mat becomes too heavy to remain at the air-broth interface and sinks below the surface, where all the cells perish due to lack of oxygen. Colonies with cheaters collapse faster than purely cooperative colonies.
- 12.
Gregory Velicer was a graduate student, and later a postdoc, in Lenski’s laboratory before moving to Indiana University and later leading his own research laboratories at the Max Planck Institute in Tübingen and at the ETH in Zurich.
- 13.
Note how this essentially mimics a reverse of ZoBell’s “bottle effect” (Chap. 9).
- 14.
Recall from Chap. 11 that C. reinhardtii reproduces by multiple fission, and offspring do not break out from the parent immediately.
- 15.
See Chap. 11.
- 16.
A similar process appears to be at work in higher-level collectivities. In eusocial insects, there can be several orders of magnitude difference in the lifetime of workers compared to queens, and this has been suggested to result from oxidative stress on the workers’ metabolism (Corona et al. 2005; Remolina and Hughes 2008; Goldsby et al. 2014a). Young and Robinson (1983) suggested that the increased metabolic load imposed by foraging behavior in workers increases oxidative stress. The relative roles of RNA and DNA, which have significantly different levels of stability with respect to mutation, may represent a molecular division of labor “ensuring both high fidelity transmission of hereditary information and the execution of critical chemical work” (Goldsby et al. 2014a).
- 17.
In these experiments, colonies could only perform up to a total of five tasks.
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Bahar, S. (2018). Experimental Evolution. In: The Essential Tension. The Frontiers Collection. Springer, Dordrecht. https://doi.org/10.1007/978-94-024-1054-9_12
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