GLOBAL KNOWLEDGE ARK

Biology · Evolutionary Processes & Adaptation

Experimental and observed evolution, including resistance

Evolution can be investigated while populations reproduce. Researchers can compare starting material with descendants, record conditions and test alternative explanations. A change observed under one set of conditions is evidence to analyse, not a licence to assume every future population will repeat the same outcome.

The introductory inheritance chapter distinguishes alleles, genotypes and phenotypes. Meiosis explains segregation and crossing over; the DNA chapter explains sequence changes. Fractions and simple multiplication are enough for the constructed examples here. No example represents a measured natural population unless explicitly identified as published research.

Back to the Biology map

By the end, you can

Use lineage history, controls and direct observations to test evolutionary explanations, including the origin and spread of resistance.

01

When a variant arose can change how many descendants it has

In 1943, Salvador Luria and Max Delbrück examined bacterial resistance to a virus, or bacteriophage. Their comparison addressed whether resistance variants arose before the viral challenge or only as an induced response to it. Uneven numbers of resistant descendants among independently growing groups supported the existence of variants before exposure in that studied system.

An early sequence change can be inherited by a large clone, while a later change has fewer opportunities to produce descendants before the comparison. Therefore, counting resistant individuals is not the same as counting independent mutation events. One event can give many resistant descendants.

The original lineage diagram illustrates that logic without reproducing the experiment. It follows four rounds of equal binary division from one starting cell and assumes faithful inheritance, no death and only one assigned variant event. An event in one cell at generation 1 leaves three further divisions, yielding eight variant descendants. An event in one cell at generation 3 leaves one division, yielding two.

The historical result does not establish a constant mutation rate under every condition or prove that all resistance variants always arise before any exposure. Mutation mechanisms can depend on conditions, as the preceding lesson explains. The test separated specific hypotheses in a particular system.

One early origin can leave more descendants

One early origin can leave more descendantsTwo complete constructed four-generation binary trees differ only in the assigned generation when one S cell changes to R.Early eventOne assigned event at generation 1Generation01234SRSRRSSRRRRSSSSRRRRRRRRSSSSSSSS8 R, 8 S; one assigned mutation eventLate eventOne assigned event at generation 3Generation01234SSSSSSSRSSSSSSSRRSSSSSSSSSSSSSS2 R, 14 S; one assigned mutation eventVariant descendantsOther descendants
R and S are abstract heritable variants, not named strains or a microbial growth protocol. Equal binary division, faithful inheritance, no death and exactly one assigned event in each tree; no real mutation rate or challenge outcome is estimated.
Read the diagram in words

Two complete constructed four-generation binary trees differ only in the assigned generation when one S cell changes to R.

Early event

Cell identityGenerationInherited variant
early-g0-i00S
early-g1-i01R
early-g1-i11S
early-g2-i02R
early-g2-i12R
early-g2-i22S
early-g2-i32S
early-g3-i03R
early-g3-i13R
early-g3-i23R
early-g3-i33R
early-g3-i43S
early-g3-i53S
early-g3-i63S
early-g3-i73S
early-g4-i04R
early-g4-i14R
early-g4-i24R
early-g4-i34R
early-g4-i44R
early-g4-i54R
early-g4-i64R
early-g4-i74R
early-g4-i84S
early-g4-i94S
early-g4-i104S
early-g4-i114S
early-g4-i124S
early-g4-i134S
early-g4-i144S
early-g4-i154S
Parent cellDaughter cell
early-g0-i0early-g1-i0
early-g0-i0early-g1-i1
early-g1-i0early-g2-i0
early-g1-i0early-g2-i1
early-g1-i1early-g2-i2
early-g1-i1early-g2-i3
early-g2-i0early-g3-i0
early-g2-i0early-g3-i1
early-g2-i1early-g3-i2
early-g2-i1early-g3-i3
early-g2-i2early-g3-i4
early-g2-i2early-g3-i5
early-g2-i3early-g3-i6
early-g2-i3early-g3-i7
early-g3-i0early-g4-i0
early-g3-i0early-g4-i1
early-g3-i1early-g4-i2
early-g3-i1early-g4-i3
early-g3-i2early-g4-i4
early-g3-i2early-g4-i5
early-g3-i3early-g4-i6
early-g3-i3early-g4-i7
early-g3-i4early-g4-i8
early-g3-i4early-g4-i9
early-g3-i5early-g4-i10
early-g3-i5early-g4-i11
early-g3-i6early-g4-i12
early-g3-i6early-g4-i13
early-g3-i7early-g4-i14
early-g3-i7early-g4-i15

One assigned event at generation 1: early-g1-i0. 8 R, 8 S; one assigned mutation event

Late event

Cell identityGenerationInherited variant
late-g0-i00S
late-g1-i01S
late-g1-i11S
late-g2-i02S
late-g2-i12S
late-g2-i22S
late-g2-i32S
late-g3-i03R
late-g3-i13S
late-g3-i23S
late-g3-i33S
late-g3-i43S
late-g3-i53S
late-g3-i63S
late-g3-i73S
late-g4-i04R
late-g4-i14R
late-g4-i24S
late-g4-i34S
late-g4-i44S
late-g4-i54S
late-g4-i64S
late-g4-i74S
late-g4-i84S
late-g4-i94S
late-g4-i104S
late-g4-i114S
late-g4-i124S
late-g4-i134S
late-g4-i144S
late-g4-i154S
Parent cellDaughter cell
late-g0-i0late-g1-i0
late-g0-i0late-g1-i1
late-g1-i0late-g2-i0
late-g1-i0late-g2-i1
late-g1-i1late-g2-i2
late-g1-i1late-g2-i3
late-g2-i0late-g3-i0
late-g2-i0late-g3-i1
late-g2-i1late-g3-i2
late-g2-i1late-g3-i3
late-g2-i2late-g3-i4
late-g2-i2late-g3-i5
late-g2-i3late-g3-i6
late-g2-i3late-g3-i7
late-g3-i0late-g4-i0
late-g3-i0late-g4-i1
late-g3-i1late-g4-i2
late-g3-i1late-g4-i3
late-g3-i2late-g4-i4
late-g3-i2late-g4-i5
late-g3-i3late-g4-i6
late-g3-i3late-g4-i7
late-g3-i4late-g4-i8
late-g3-i4late-g4-i9
late-g3-i5late-g4-i10
late-g3-i5late-g4-i11
late-g3-i6late-g4-i12
late-g3-i6late-g4-i13
late-g3-i7late-g4-i14
late-g3-i7late-g4-i15

One assigned event at generation 3: late-g3-i0. 2 R, 14 S; one assigned mutation event

02

Resistance has an origin and a subsequent population history

Antimicrobial resistance concerns microbes with reduced susceptibility to a relevant antimicrobial; it does not mean the patient’s body has become resistant. In antibiotic comparisons, resistant bacteria can grow under conditions that inhibit susceptible alternatives. Genetic changes can alter a target, transport or other cellular functions, and resistance-associated material can also be acquired from other microbes.

A selective condition can increase the representation of heritable resistant variants because their descendants contribute more than susceptible alternatives under that condition. That enrichment need not create the original change. Moving resistant microbes or transferring resistance-associated genetic material adds routes of spread beyond a local lineage’s reproduction.

A surviving cell is not, by that observation alone, proof of a new heritable resistance mutation. Temporary physiological states, different exposure and other explanations may affect survival. Establishing heritability and the relevant mechanism requires appropriate evidence. This chapter supplies evolutionary reasoning, not treatment decisions or a procedure for growing resistant microbes.

A resistant lineage can originate, expand and move

A resistant lineage can originate, expand and moveDistinct pathways show a sequence change in one microbe, inheritance by descendants, and genetic material entering a recipient from another microbe. A selective context changes reproductive representation rather than reaching back to choose the original change.Geneticmaterial entersanother microbeStarting microbeInitial geneticmaterialMicrobe with aninherited variantChanged geneticmaterialDescendants inheritthe variantVariant indescendantsAnother microbialsourceResistance-associatedmaterialRecipient acquiringinherited materialAcquired materialChanged reproductiverepresentationInherited variationin a populationRelevant selectivecontextDifferentreproductivecontributions underspecified conditionsVariant arisesInherited during reproductionGenetic material enters another microbeContributes descendantsContributes descendants if inheritedSelection affects representation
Conceptual cells and inherited material; no organism, plasmid, drug, exposure, transfer procedure or measured success rate is specified. Survival by itself does not prove a heritable mutation.
Read the diagram in words

Distinct pathways show a sequence change in one microbe, inheritance by descendants, and genetic material entering a recipient from another microbe. A selective context changes reproductive representation rather than reaching back to choose the original change.

Selected entityInherited material or context
Starting microbeInitial genetic material
Microbe with an inherited variantChanged genetic material
Descendants inherit the variantVariant in descendants
Another microbial sourceResistance-associated material
Recipient acquiring inherited materialAcquired material
Changed reproductive representationInherited variation in a population
Relevant selective contextDifferent reproductive contributions under specified conditions
SourceDestinationRelationship
Starting microbeMicrobe with an inherited variantVariant arises
Microbe with an inherited variantDescendants inherit the variantInherited during reproduction
Another microbial sourceRecipient acquiring inherited materialGenetic material enters another microbe
Descendants inherit the variantChanged reproductive representationContributes descendants
Recipient acquiring inherited materialChanged reproductive representationContributes descendants if inherited
Relevant selective contextChanged reproductive representationSelection affects representation
03

Replay experiments can test the role of history

A 2008 study by Zachary Blount, Christina Borland and Richard Lenski examined the emergence of citrate use in an experimental Escherichia coli lineage under oxygen-containing conditions. Comparisons using stored earlier material investigated whether previous evolutionary changes affected the later outcome. The results supported a role for historical genetic background in that studied innovation.

The point is not that a population consciously prepared for a future trait. Earlier variants arose and spread under their own conditions; they could change the consequences or accessibility of later changes. Replaying from different earlier backgrounds tests that dependence more directly than inventing a story after seeing the endpoint.

Even carefully controlled experimental populations are finite and have specific environments. Repeated populations need not reach identical outcomes. Their results can reveal mechanisms and constraints, while extrapolation to other species, habitats or clinical situations needs additional evidence. The diagram represents the comparison logic, not the study’s measured success frequencies.

Stored earlier material can test historical dependence

Stored earlier material can test historical dependenceA source lineage changes through time; retained samples of different earlier backgrounds separately seed repeated comparisons under the same specified assay context.Earlierbackground ALater backgroundBSubsequent sourcelineageRetained AmaterialRetained BmaterialIndependentcomparisons fromAIndependentcomparisons fromBSame specified assay context; compare inherited outcomes
Logic of historical comparisons, not the exact branches, number of replays or outcome rates of the 2008 study. An earlier background is not guaranteed to produce or prevent the innovation.
Read the diagram in words

A source lineage changes through time; retained samples of different earlier backgrounds separately seed repeated comparisons under the same specified assay context.

SourceDestinationRelationship
Earlier background ALater background BInherited changes over time
Later background BSubsequent source lineageFurther lineage history
Earlier background ARetained A materialRetain material
Later background BRetained B materialRetain material
Retained A materialIndependent comparisons from AStart comparisons from A
Retained B materialIndependent comparisons from BStart comparisons from B

Same specified assay context; compare inherited outcomes

04

Design comparisons that distinguish explanations

For a field population, record what changed, what was sampled and how inherited variation was identified. Compare environments, reproductive outcomes and possible movement between populations. For an experimental population, ancestral material and repeated independent lineages can help separate inherited change from immediate environmental effects.

An increase at a locus, a changed character and a reproductive advantage are related observations that still answer different questions. Evidence becomes more persuasive when those observations form a tested causal account and plausible alternatives are checked. A negative result can also narrow an explanation: failure to repeat an outcome may reveal dependence on history or conditions.

Worked example

Count events separately from descendants

In the constructed early-event tree, one variant cell at generation 1 becomes 2, then 4, then 8 variant cells by generation 4. In the late-event tree, one variant cell at generation 3 becomes 2 by generation 4. Each full tree ends with 16 cells: eight variants and eight other cells, or two variants and fourteen others.

Both trees contain one assigned mutation event. The final variant frequencies are 8/16 = 0.50 and 2/16 = 0.125. Their difference follows from timing under this model, without a difference in division rate or an extra mutation. These small deterministic trees are not estimates of real mutation rates or the distribution in Luria and Delbrück’s data.

Check your understanding

  1. Does eight resistant descendants necessarily mean eight independent resistance mutations?

    Show reasoning

    No. A single inherited change can expand through a clone. Lineage relationships and the timing of changes help distinguish the number of origins from the number of descendants.

  2. Why compare an evolved phenotype with ancestral material under the same assay conditions?

    Show reasoning

    It helps separate an inherited difference from a difference caused merely by the immediate environment or measurement. Appropriate replication and other controls are still needed; the comparison is not sufficient for every mechanism claim.

  3. If only one of several experimental lineages develops a new trait, has the evidence for evolution failed?

    Show reasoning

    No. The observed inherited change remains evidence. Different outcomes can reveal the influence of mutation supply, chance and history. A mechanism does not have to guarantee the same endpoint in every finite lineage to operate.

What to carry forward

Observed evolution is tested through inherited differences, reproductive histories and explicit comparisons; origins, enrichment and spread are separate parts of a resistance explanation.

References

Resources supporting this lesson’s explanations and examples.

  1. Mutations of Bacteria from Virus Sensitivity to Virus Resistance (Genetics 28(6), 491–511, 1943; DOI 10.1093/genetics/28.6.491)Salvador E. Luria and Max DelbrückOriginal researchpmc.ncbi.nlm.nih.govRead source
  2. Salvador Luria and Max Delbrück on Random Mutation and Fluctuation Tests (Genetics 202(2), 367–368, 2016; DOI 10.1534/genetics.115.186163)Andrew MurrayAuthor historical perspectiveacademic.oup.comRead source
  3. Historical contingency and the evolution of a key innovation in an experimental population of Escherichia coli (PNAS 105(23), 7899–7906, 2008; DOI 10.1073/pnas.0803151105)Zachary D. Blount, Christina Z. Borland and Richard E. LenskiOriginal researchpmc.ncbi.nlm.nih.govRead source
  4. About Antimicrobial Resistance (Institutional explanation dated 2025-01-31; inspected 2026-10-12)United States Centers for Disease Control and PreventionInstitutional resourcecdc.govRead source
  5. Mutation as a Stress Response and the Regulation of Evolvability (Critical Reviews in Biochemistry and Molecular Biology 42(5), 399–435, 2007; DOI 10.1080/10409230701648502)Rodrigo S. Galhardo, P. J. Hastings and Susan M. RosenbergAuthor scientific reviewpmc.ncbi.nlm.nih.govRead source
  6. Definitions and guidelines for research on antibiotic persistence (Nature Reviews Microbiology 17, 441–448, 2019; DOI 10.1038/s41579-019-0196-3)Nathalie Q. Balaban and international coauthorsAuthor consensus statementnature.comRead source