Laboratory experiments of speciation

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A simplification of an allopatric speciation experiment where two lines of fruit flies are raised on maltose and starch media Drosophila speciation experiment.svg
A simplification of an allopatric speciation experiment where two lines of fruit flies are raised on maltose and starch media

Laboratory experiments of speciation have been conducted for all four modes of speciation: allopatric, peripatric, parapatric, and sympatric; and various other processes involving speciation: hybridization, reinforcement, founder effects, among others. Most of the experiments have been done on flies, in particular Drosophila fruit flies. [1] However, more recent studies have tested yeasts, fungi, and even viruses.

Contents

It has been suggested that laboratory experiments are not conducive to vicariant speciation events (allopatric and peripatric) due to their small population sizes and limited generations. [2] Most estimates from studies of nature indicate that speciation takes hundreds of thousands to millions of years. [3] On the other hand, many species are thought to have speciated faster and more recently, such as the European flounders ( Platichthys flesus ) that spawn in pelagic and demersal zones—having allopatrically speciated in under 3000 generations. [4]

Table of experiments

Six publications have attempted to compile, review, and analyze the experimental research on speciation:

  1. John Ringo, David Wood, Robert Rockwell, and Harold Dowse in 1985; [5]
  2. William R. Rice and Ellen E. Hostert in 1993; [6]
  3. Ann-Britt Florin and Anders Ödeen in 2002; [2]
  4. Mark Kirkpatrick and Virginie Ravigné in 2002; [7]
  5. Jerry A. Coyne and H. Allen Orr in 2004; [1] and
  6. James D. Fry in 2009. [8]

The table summarizes the studies and data reviewed in these publications. It also references several contemporary experiments and is non-exhaustive. In the table, multiple numbers separated by semi-colons in the generations column indicate that multiple experiments were conducted. The replications (in parentheses) indicates the number of populations used in the experiments—i.e. how many times the experiment was replicated. Various types of selection have been imposed on experimental populations and are indicated by the selection type column. Negative or positive results of each experiment are provided by the reproductive isolation column. Pre-zygotic reproductive isolation means that the reproducing individuals in the populations were unable to produce offspring (effectively a positive result). Post-zygotic isolation means that the reproducing individuals were able to produce offspring but they were either sterile or inviable (a positive result as well). Negative results are indicated by "none"—that is, the experiments did not result in reproductive isolation.

Laboratory experiments of speciation [1] [6] [7] [2] [8]
Species TraitGenerations (replications) [duration]Tested Selection typeStudied genetic drift Reproductive isolation ReferenceYear
Drosophila melanogaster Escape response 18 Vicariant, reinforcement, parapatric/

sympatric

Indirect; divergentYesPre-zygoticGrant & Mettler [9] 1969
D. melanogaster Locomotion 112VicariantIndirect; divergentNoPre-zygoticBurnet & Connolly [10] 1974
D. melanogasterTemperature, humidity 70–130VicariantIndirect; divergentYesPre-zygoticKilias et al. [11] 1980
D. melanogaster DDT adaptation600 [25 years, +15 years]VicariantDirectNoPre-zygoticBoake et al. [12] 2003
D. melanogaster17, 9, 9, 1, 1, 7, 7, 7, 7Vicariant; parapatric/

sympatric

Direct, divergentPre-zygotic in vicariance; none with gene flowBarker & Karlsson [13] 1974
D. melanogaster40; 50ReinforcementDirect; divergentPre-zygoticCrossley [14] 1974
D. melanogasterLocomotion45VicariantDirect; divergentNoNonevan Dijken & Scharloo [15] [16] 1979
D. melanogasterReinforcementDirect; divergentPre-zygoticWallace [17] 1953
D. melanogaster36; 31ReinforcementDirect; divergentPre-zygoticKnight [18] 1956
D. melanogaster EDTA adaptation25, 25, 25, 14Semi-allopatric, reinforcementIndirect; divergentNoPost-zygoticRobertson [19] [20] 1966
D. melanogaster25 (8)Vicariant; reinforcement; parapatric; sympatricDirectNoneHostert [21] 1997
D. melanogasterAbdominal chaeta

number

21–31VicariantDirectYesNoneSantibanez & Waddington [22] 1958
D. melanogaster Sternopleural chaeta number32Vicariant, reinforcement, parapatric/

sympatric

DirectNoNoneBarker & Cummins [23] 1969
D. melanogaster Phototaxis, geotaxis 20VicariantNoNoneMarkow [24] [25] 1975; 1981
D. melanogasterPeripatricYesRundle et al. [26] 1998
D. melanogasterVicariant; peripatricYesMooers et al. [27] 1999
D. melanogaster12ReinforcementDivergentPre-zygoticThoday & Gibson [28] 1962
D. melanogasterNoneThoday & Gibson [29] [30] 1970; 1971
D. melanogaster16ReinforcementIndirectNoneSpiess & Wilke [31] 1954
D. melanogasterReinforcementDirect; divergentPre-zygoticEhrman [32] [33] [34] [35] 1971; 1973; 1979; 1983
D. melanogasterSternopleural chaeta number5; 27; 27; 1; 1; 1; 1; 1Parapatric/

sympatric

NoneChabora [36] 1968
D. melanogasterNoneScharloo [37] 1967
D. melanogaster1, 1Coyne & Grant [38] 1972
D. melanogaster25Rice [39] 1985
D. melanogaster25DisruptivePre-zygoticRice & Salt [40] 1988
D. melanogaster35; 35SympatricPre-zygoticRice & Salt [41] 1990
D. melanogaster NaCl and CuSO4 levels in food[3 years in allopatry, 1 in sympatry]Allopatric; reinforcement; sympatricPre-zygotic in allopatry, none in sympatryWallace [42] 1982
D. melanogasterReinforcementEhrman et al. [43] 1991
D. melanogasterReinforcementFukatami & Moriwaki [44] 1970
Drosophila simulans Scutellar bristles, development speed, wing width; desiccation resistance, fecundity, ethanol resistance; courtship display, re-mating speed, lek behavior; pupation height, clumped egg laying, general activity[3 years]Vicariant; peripatricYesPost-zygoticRingo et al. [5] 1985
Drosophila paulistorum 131; 131ReinforcementDirectPre-zygoticDobzhansky et al. [45] 1976
D. paulistorum[5 years]VicariantDobzhansky and Pavlovsky [46] 1966
Drosophila willistoni pH adaptation34–122VicariantIndirect; divergentNoPre-zygoticKalisz & Cordeiro [47] 1980
Drosophila pseudoobscura Carbohydrate source12VicariantIndirectYesPre-zygoticDodd [48] 1989
D. pseudoobscuraTemperature adaptation25–60VicariantDirectEhrman [49] [50] [51] [52] [53] 1964;

1969

D. pseudoobscuraPhototaxis, geotaxis5–11VicariantIndirectNoPre-zygoticdel Solar [54] 1966
D. pseudoobscuraVicariant; peripatricPre-zygoticPowell [55] [56] 1978; 1985
D. pseudoobscuraPeripatric; vicariantYesGaliana et al. [57] 1993
D. pseudoobscuraTemperature photoperiod; food37 (78) [33–34 months]VicariantDivergentYesNoneRundle [58] 2003
D. pseudoobscura &

Drosophila persimilis

22; 16; 9ReinforcementDirect; divergentPre-zygoticKoopman [59] 1950
D. pseudoobscura &

D. persimilis

18 (4)DirectPre-zygoticKessler [60] 1966
Drosophila mojavensis 12DirectPre-zygoticKoepfer [61] 1987
D. mojavensisDevelopment time13DivergentYesNoneEtges [62] 1998
Drosophila adiastola PeripatricYesPre-zygoticArita & Kaneshiro [63] 1974
Drosophila silvestris PeripatricYesAhearn [64] 1980
Musca domestica Geotaxis38VicariantIndirectNoPre-zygoticSoans et al. [65] 1974
M. domesticaGeotaxis16VicariantDirect; divergentNoPre-zygoticHurd & Eisenburg [66] 1975
M. domesticaPeripatricYesMeffert & Bryant [67] 1991
M. domesticaRegan et al. [68] 2003
Bactrocera cucurbitae Development time40–51DivergentYesPre-zygoticMiyatake & Shimizu [69] 1999
Zea mays 6; 6ReinforcementDirect; divergentPre-zygoticPaterniani [70] 1969
Drosophila grimshawi PeripatricJones, Widemo, & Arrendal [71] N/A
Saccharomyces cerevisiae Leu & Murry [72] 2006
D. melanogasterReinforcementHarper & Lambert [73] 1983
Tribolium castaneum Pupa l weight15 (6)DisruptiveHalliburton & Gall [74] 1983
D. melanogasterGeotaxisDivergentLofdahl et al. [75] 1992
D. pseudoobscura[10 years]Moya et al. [76] 1995
Neurospora DivergentDettman et al. [77] 2008
S. cerevisiae500DivergentDettman et al. [78] 2007
Sepsis cynipsea 35Martin & Hosken [79] 2003
D. melanogasterWigby & Chapman [80] 2006
D. pseudoobscuraSexual conflict48–52 (4; 4; 4)Bacigalupe et al. [81] 2007
D. serrataRundle et al. [82] 2005
Drosophila serrata & D. birchiiMate recognition9 (3; 3)ReinforcementNaturalPre-zygoticHiggie et al. [83] 2000
Enterobacteria phage λ Escherichia coli receptor exploitation35 cycles (6)Vicariant, sympatricPre-zygoticMeyer et al. [84] 2016
Tetranychus urticae Resistance to host plant toxinOvermeer [85] 1966
T. urticaeResistance to host plant toxinFry [86] 1999
Helianthus annus × H. petiolaris and H. anomalusHybridRieseburg et al. [87] 1996
S. cerevisiaeGreig et al. [88] 2002
D. melanogaster Life history Ghosh & Joshi [89] 2012
Drosophila subobscura Mate behaviorBárbaro et al. [90] 2015
Digital organisms~42,000; ~850 (20)EcologicalPost-zygoticAnderson & Harmon [91] 2014
Schizosaccharomyces pombe Complete reproductive isolationSeike et al. [92] 2015
D. pseudoobscuraCourtship song130Debelle et al. [93] 2014
Callosobruchus maculatus 40 (16)Debelle et al. [94] 2010

See also

Related Research Articles

<i>Drosophila</i> Genus of flies

Drosophila is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit. They should not be confused with the Tephritidae, a related family, which are also called fruit flies ; tephritids feed primarily on unripe or ripe fruit, with many species being regarded as destructive agricultural pests, especially the Mediterranean fruit fly.

Microevolution is the change in allele frequencies that occurs over time within a population. This change is due to four different processes: mutation, selection, gene flow and genetic drift. This change happens over a relatively short amount of time compared to the changes termed macroevolution.

Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within lineages. Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book On the Origin of Species. He also identified sexual selection as a likely mechanism, but found it problematic.

<span class="mw-page-title-main">Theodosius Dobzhansky</span> Russian-American geneticist and evolutionary biologist (1900–1975)

Theodosius Grigorievich Dobzhansky was a prominent Russian and American geneticist and evolutionary biologist. He was a central figure in the field of evolutionary biology for his work in shaping the modern synthesis. Born in the Russian Empire, Dobzhansky emigrated to the United States in 1927, aged 27.

Allopatric speciation – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from each other to an extent that prevents or interferes with gene flow.

<span class="mw-page-title-main">Haldane's rule</span> Observation in evolutionary biology

Haldane's rule is an observation about the early stage of speciation, formulated in 1922 by the British evolutionary biologist J. B. S. Haldane, that states that if — in a species hybrid — only one sex is inviable or sterile, that sex is more likely to be the heterogametic sex. The heterogametic sex is the one with two different sex chromosomes; in therian mammals, for example, this is the male.

<span class="mw-page-title-main">Peripatric speciation</span> Speciation in which a new species is formed from an isolated smaller peripheral population

Peripatric speciation is a mode of speciation in which a new species is formed from an isolated peripheral population. Since peripatric speciation resembles allopatric speciation, in that populations are isolated and prevented from exchanging genes, it can often be difficult to distinguish between them. Nevertheless, the primary characteristic of peripatric speciation proposes that one of the populations is much smaller than the other. The terms peripatric and peripatry are often used in biogeography, referring to organisms whose ranges are closely adjacent but do not overlap, being separated where these organisms do not occur—for example on an oceanic island compared to the mainland. Such organisms are usually closely related ; their distribution being the result of peripatric speciation.

Experimental evolution is the use of laboratory experiments or controlled field manipulations to explore evolutionary dynamics. Evolution may be observed in the laboratory as individuals/populations adapt to new environmental conditions by natural selection.

<i>Genetics and the Origin of Species</i> 1937 book by Theodosius Dobzhansky

Genetics and the Origin of Species is a 1937 book by the Ukrainian-American evolutionary biologist Theodosius Dobzhansky. It is regarded as one of the most important works of modern synthesis and was one of the earliest. The book popularized the work of population genetics to other biologists and influenced their appreciation for the genetic basis of evolution. In his book, Dobzhansky applied the theoretical work of Sewall Wright (1889–1988) to the study of natural populations, allowing him to address evolutionary problems in a novel way during his time. Dobzhansky implements theories of mutation, natural selection, and speciation throughout his book to explain the habits of populations and the resulting effects on their genetic behavior. The book explains evolution in depth as a process over time that accounts for the diversity of all life on Earth. The study of evolution was present, but greatly neglected at the time. Dobzhansky illustrates that evolution regarding the origin and nature of species during this time in history was deemed mysterious, but had expanding potential for progress to be made in its field.

<span class="mw-page-title-main">Disruptive selection</span> Natural selection for extreme trait values over intermediate ones

In evolutionary biology, disruptive selection, also called diversifying selection, describes changes in population genetics in which extreme values for a trait are favored over intermediate values. In this case, the variance of the trait increases and the population is divided into two distinct groups. In this more individuals acquire peripheral character value at both ends of the distribution curve.

<span class="mw-page-title-main">Parapatric speciation</span> Speciation within a population where subpopulations are reproductively isolated

In parapatric speciation, two subpopulations of a species evolve reproductive isolation from one another while continuing to exchange genes. This mode of speciation has three distinguishing characteristics: 1) mating occurs non-randomly, 2) gene flow occurs unequally, and 3) populations exist in either continuous or discontinuous geographic ranges. This distribution pattern may be the result of unequal dispersal, incomplete geographical barriers, or divergent expressions of behavior, among other things. Parapatric speciation predicts that hybrid zones will often exist at the junction between the two populations.

The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring are sterile. These barriers maintain the integrity of a species by reducing gene flow between related species.

<i>Drosophila pseudoobscura</i> Species of fly

Drosophila pseudoobscura is a species of fruit fly, used extensively in lab studies of speciation. It is native to western North America.

<span class="mw-page-title-main">Ecological speciation</span>

Ecological speciation is a form of speciation arising from reproductive isolation that occurs due to an ecological factor that reduces or eliminates gene flow between two populations of a species. Ecological factors can include changes in the environmental conditions in which a species experiences, such as behavioral changes involving predation, predator avoidance, pollinator attraction, and foraging; as well as changes in mate choice due to sexual selection or communication systems. Ecologically-driven reproductive isolation under divergent natural selection leads to the formation of new species. This has been documented in many cases in nature and has been a major focus of research on speciation for the past few decades.

<span class="mw-page-title-main">Reinforcement (speciation)</span> Process of increasing reproductive isolation

Reinforcement is a process of speciation where natural selection increases the reproductive isolation between two populations of species. This occurs as a result of selection acting against the production of hybrid individuals of low fitness. The idea was originally developed by Alfred Russel Wallace and is sometimes referred to as the Wallace effect. The modern concept of reinforcement originates from Theodosius Dobzhansky. He envisioned a species separated allopatrically, where during secondary contact the two populations mate, producing hybrids with lower fitness. Natural selection results from the hybrid's inability to produce viable offspring; thus members of one species who do not mate with members of the other have greater reproductive success. This favors the evolution of greater prezygotic isolation. Reinforcement is one of the few cases in which selection can favor an increase in prezygotic isolation, influencing the process of speciation directly. This aspect has been particularly appealing among evolutionary biologists.

<span class="mw-page-title-main">History of speciation</span> Aspect of history

The scientific study of speciation — how species evolve to become new species — began around the time of Charles Darwin in the middle of the 19th century. Many naturalists at the time recognized the relationship between biogeography and the evolution of species. The 20th century saw the growth of the field of speciation, with major contributors such as Ernst Mayr researching and documenting species' geographic patterns and relationships. The field grew in prominence with the modern evolutionary synthesis in the early part of that century. Since then, research on speciation has expanded immensely.

<span class="mw-page-title-main">Evidence for speciation by reinforcement</span> Overview article

Reinforcement is a process within speciation where natural selection increases the reproductive isolation between two populations of species by reducing the production of hybrids. Evidence for speciation by reinforcement has been gathered since the 1990s, and along with data from comparative studies and laboratory experiments, has overcome many of the objections to the theory. Differences in behavior or biology that inhibit formation of hybrid zygotes are termed prezygotic isolation. Reinforcement can be shown to be occurring by measuring the strength of prezygotic isolation in a sympatric population in comparison to an allopatric population of the same species. Comparative studies of this allow for determining large-scale patterns in nature across various taxa. Mating patterns in hybrid zones can also be used to detect reinforcement. Reproductive character displacement is seen as a result of reinforcement, so many of the cases in nature express this pattern in sympatry. Reinforcement's prevalence is unknown, but the patterns of reproductive character displacement are found across numerous taxa, and is considered to be a common occurrence in nature. Studies of reinforcement in nature often prove difficult, as alternative explanations for the detected patterns can be asserted. Nevertheless, empirical evidence exists for reinforcement occurring across various taxa and its role in precipitating speciation is conclusive.

Maria R. Servedio is a Canadian-American professor at the University of North Carolina at Chapel Hill. Her research spans a wide range of topics in evolutionary biology from sexual selection to evolution of behavior. She largely approaches these topics using mathematical models. Her current research interests include speciation and reinforcement, mate choice, and learning with a particular focus on evolutionary mechanisms that promote premating (prezygotic) isolation. Through integrative approaches and collaborations, she uses mathematical models along with experimental, genetic, and comparative techniques to draw conclusions on how evolution occurs. She has published extensively on these topics and has more than 50 peer-reviewed articles. She served as Vice President in 2018 of the American Society of Naturalists, and has been elected to serve as President in 2023.

Allochronic speciation is a form of speciation arising from reproductive isolation that occurs due to a change in breeding time that reduces or eliminates gene flow between two populations of a species. The term allochrony is used to describe the general ecological phenomenon of the differences in phenology that arise between two or more species—speciation caused by allochrony is effectively allochronic speciation.

In biology, parallel speciation is a type of speciation where there is repeated evolution of reproductively isolating traits via the same mechanisms occurring between separate yet closely related species inhabiting different environments. This leads to a circumstance where independently evolved lineages have developed reproductive isolation from their ancestral lineage, but not from other independent lineages that inhabit similar environments. In order for parallel speciation to be confirmed, there is a set of three requirements that has been established that must be met: there must be phylogenetic independence between the separate populations inhabiting similar environments to ensure that the traits responsible for reproductive isolation evolved separately, there must be reproductive isolation not only between the ancestral population and the descendent population, but also between descendent populations that inhabit dissimilar environments, and descendent populations that inhabit similar environments must not be reproductively isolated from one another. To determine if natural selection specifically is the cause of parallel speciation, a fourth requirement has been established that includes identifying and testing an adaptive mechanism, which eliminates the possibility of a genetic factor such as polyploidy being the responsible agent.

References

  1. 1 2 3 Coyne, Jerry A.; Orr, H. Allen (2004), Speciation, Sinauer Associates, pp. 1–545, ISBN   978-0-87893-091-3
  2. 1 2 3 Florin, Ann-Britt & Ödeen, Anders (2002), "Laboratory environments are not conducive for allopatric speciation", Journal of Evolutionary Biology, 15 (1): 10–19, doi: 10.1046/j.1420-9101.2002.00356.x {{citation}}: CS1 maint: multiple names: authors list (link)
  3. ↑ Coyne, Jerry A.; Orr, H. Allen (1997), ""Patterns of Speciation in Drosophila" Revisited", Evolution, 51 (1): 295–303, doi: 10.1111/j.1558-5646.1997.tb02412.x , PMID   28568795
  4. ↑ Momigliano, Paolo; Jokinen, Henri; Fraimout, Antoine; Florin, Ann-Britt; Norkko, Alf; Merilä, Juha (2017), "Extraordinarily rapid speciation in a marine fish" (PDF), PNAS, 114 (23): 6074–6079, Bibcode:2017PNAS..114.6074M, doi: 10.1073/pnas.1615109114 , PMC   5468626 , PMID   28533412
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  23. ↑ Barker, J. S. F. & Cummins, L. J. (1969), "The effect of selection for sternopleural bristle number in mating behaviour in Drosophila melanogaster", Genetics, 61 (3): 713–719, doi:10.1093/genetics/61.3.713, PMC   1212235 , PMID   17248436 {{citation}}: CS1 maint: multiple names: authors list (link)
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  25. ↑ Markow, Therese Ann (1981), "Mating preferences are not predictive of the direction of evolution in experimental populations of Drosophila", Science, 213 (4514): 1405–1407, Bibcode:1981Sci...213.1405M, doi:10.1126/science.213.4514.1405, PMID   17732575, S2CID   15497733
  26. ↑ Rundle, H. D., Mooers, Arne Ø. & Whitlock, Michael C. (1998), "Single founder-flush events and the evolution of reproductive isolation", Evolution, 52 (6): 1850–1855, doi:10.1111/j.1558-5646.1998.tb02263.x, PMID   28565304, S2CID   24502821 {{citation}}: CS1 maint: multiple names: authors list (link)
  27. ↑ Mooers, Arne Ø., Rundle, Howard D. & Whitlock, Michael C. (1999), "The effects of selection and bottlenecks on male mating success in peripheral isolates", American Naturalist, 153 (4): 437–444, doi:10.1086/303186, PMID   29586617, S2CID   4411105 {{citation}}: CS1 maint: multiple names: authors list (link)
  28. ↑ Thoday, J. M. & Gibson, J. B. (1962), "Isolation by disruptive selection", Nature, 193 (4821): 1164–1166, Bibcode:1962Natur.193.1164T, doi:10.1038/1931164a0, PMID   13920720, S2CID   5156234 {{citation}}: CS1 maint: multiple names: authors list (link)
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