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Biology · Fungal Biology & Symbioses

Spores, sexual and asexual cycles and mating systems

Two compatible hyphal regions join, and their nuclei occupy a shared compartment. Has a diploid nucleus formed? Counting the nuclei first prevents a common mistake: two separate haploid nuclei are a different organisation from one nucleus containing two chromosome sets.

Phylogeny, Classification & the Tree of Life introduces clades and the difference between similarity and ancestry. Energy, Enzymes & Metabolism supplies enzyme action, fermentation and separate material and energy accounts. Meiosis & Patterns of Inheritance supplies chromosome sets and DNA-copy accounting for the optional nuclear-cycle calculation. Qualitative explanations come before those calculations; the model's units and counting rules are introduced where they are used. Cells, Membranes & Organisation is a useful return path for membrane boundaries, and Microbial Eukaryotes & Algal Diversity introduces photosynthetic partners.

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By the end, you can

Trace cell joining, nuclear fusion and meiosis separately, and interpret spores and mating types within a stated life cycle.

01

A spore needs a life-cycle context

A fungal spore is a propagule that can contribute to dispersal or establishment. Sexual cycles and asexual reproduction can both produce spores. Some fungal spores are moved passively; some fungal lineages have motile zoospores. The word spore therefore does not specify whether meiosis occurred, whether mating is required next, or how long the stage survives.

Asexual production can involve mitotic division followed by packaging or separation of propagules. Sexual reproduction involves a route that combines nuclear contributions and includes meiosis. To classify a particular stage, trace the events before and after it. A small round object is insufficient evidence of its reproductive origin.

Fungal spores are also different from the bacterial endospores introduced earlier. The bacterial survival structure develops within an existing cell and is not a general mechanism for multiplying that cell into numerous offspring. A shared name should prompt comparison of mechanisms rather than an assumption of equivalence.

02

Plasmogamy precedes karyogamy in the selected route

Plasmogamy is the joining of cells or cytoplasm. Karyogamy is fusion of nuclei. Those events can be separated in time. A dikaryotic compartment has two separate haploid nuclei, commonly represented as n + n. Some fungi maintain such an organisation for an extended phase; other fungal routes differ. The selected route is an accounting model, not the timetable of all fungi.

Here n means one complete assigned set of two chromosomes. Each initial unreplicated haploid nucleus has two chromosomes, two chromatids and 1C of DNA. After plasmogamy, a shared compartment contains two such nuclei: four chromosomes and 2C in total, still distributed between separate nuclei. After karyogamy, one diploid nucleus contains four chromosomes, four chromatids and 2C.

Replication changes the copies of DNA rather than the number of complete chromosome sets. The diploid nucleus then has four chromosomes, eight chromatids and 4C. At completion of meiosis I, each of two nuclei is haploid with two replicated chromosomes, four chromatids and 2C. At completion of meiosis II, each of four nuclei has two chromosomes, two chromatids and 1C. These are completed stages, with no anaphase centromere-counting ambiguity.

Nuclear boundaries change the count's meaning

Nuclear boundaries change the count's meaningChromosome and DNA accounts follow a selected fungal sexual route, with separate optional postmeiotic and asexual branches.Selected sexual routeOne selected unreplicated haploid nucleusNuclear boundaries: 1 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 2 · DNA per nucleus (C): 11Total chromosomes: 2 · Total chromatids: 2 ·Total DNA (C): 1A second compatible haploid contributionjoins; nuclei remain separateAfter plasmogamy: two separate haploid nucleiNuclear boundaries: 2 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 2 · DNA per nucleus (C): 112Total chromosomes: 4 · Total chromatids: 4 ·Total DNA (C): 2Two nuclei fuseAfter karyogamy: one diploid nucleusNuclear boundaries: 1 · Sets per nucleus: 2Chromosomes per nucleus: 4 · Chromatids pernucleus: 4 · DNA per nucleus (C): 21Total chromosomes: 4 · Total chromatids: 4 ·Total DNA (C): 2DNA copies; chromosome sets remain twoAfter premeiotic DNA replicationNuclear boundaries: 1 · Sets per nucleus: 2Chromosomes per nucleus: 4 · Chromatids pernucleus: 8 · DNA per nucleus (C): 41Total chromosomes: 4 · Total chromatids: 8 ·Total DNA (C): 4Homologues separate into two nucleiCompletion of meiosis INuclear boundaries: 2 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 4 · DNA per nucleus (C): 212Total chromosomes: 4 · Total chromatids: 8 ·Total DNA (C): 4Sisters separate into four nucleiCompletion of meiosis IINuclear boundaries: 4 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 2 · DNA per nucleus (C): 11234Total chromosomes: 8 · Total chromatids: 8 ·Total DNA (C): 4Optional postmeiotic routeOptional copying before mitosisOptional DNA replication in four haploidnucleiNuclear boundaries: 4 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 4 · DNA per nucleus (C): 21234Total chromosomes: 8 · Total chromatids: 16 ·Total DNA (C): 8Optional eight haploid nuclear productsOptional completed mitoses: eight haploidnucleiNuclear boundaries: 8 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 2 · DNA per nucleus (C): 112345678Total chromosomes: 16 · Total chromatids: 16· Total DNA (C): 8Separate asexual routeOne selected unreplicated haploid nucleusNuclear boundaries: 1 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 2 · DNA per nucleus (C): 11Total chromosomes: 2 · Total chromatids: 2 ·Total DNA (C): 1Copying in the asexual branchAsexual branch: replication in one haploidnucleusNuclear boundaries: 1 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 4 · DNA per nucleus (C): 21Total chromosomes: 2 · Total chromatids: 4 ·Total DNA (C): 2Two haploid nuclear productsAsexual branch: two haploid daughter nucleiNuclear boundaries: 2 · Sets per nucleus: 1Chromosomes per nucleus: 2 · Chromatids pernucleus: 2 · DNA per nucleus (C): 112Total chromosomes: 4 · Total chromatids: 4 ·Total DNA (C): 2One complete chromosome set per nucleusn + n: two separate nuclei2n: two sets within one nucleusCentromere-counted chromosomeTwo sister chromatids in one chromosomeTotals across the displayed nucleiNuclear boundaryAssigned glyph positions; no physical scaleNuclear products; final spore packagingunspecified1C: DNA of one unreplicated haploid nucleus
Two separate haploid nuclei can have the same total DNA as one unreplicated diploid nucleus. The diagram counts nuclear products before final spore packaging.
Read the diagram in words

The starting nucleus is n with 2 chromosomes, 2 chromatids and 1C. Plasmogamy produces two separate n nuclei, together 4 chromosomes and 2C. Karyogamy produces one 2n nucleus with 4 chromosomes and 2C. Replication gives that nucleus 4 chromosomes, 8 chromatids and 4C. After meiosis I each of two nuclei is n with 2 chromosomes, 4 chromatids and 2C. After meiosis II each of four nuclei is n with 2 chromosomes, 2 chromatids and 1C. An optional copying-and-mitosis branch produces eight n, 1C nuclei; a separate haploid mitotic branch produces two n, 1C nuclei.

Per nucleus
Completed nuclear stageNuclear boundariesSets per nucleusChromosomes per nucleusChromatids per nucleusDNA per nucleus (C)
One selected unreplicated haploid nucleus11221
After plasmogamy: two separate haploid nuclei21221
After karyogamy: one diploid nucleus12442
After premeiotic DNA replication12484
Completion of meiosis I21242
Completion of meiosis II41221
Optional DNA replication in four haploid nuclei41242
Optional completed mitoses: eight haploid nuclei81221
Asexual branch: replication in one haploid nucleus11242
Asexual branch: two haploid daughter nuclei21221
Counts across all displayed nuclei
Completed nuclear stageTotal chromosomesTotal chromatidsTotal DNA (C)
One selected unreplicated haploid nucleus221
After plasmogamy: two separate haploid nuclei442
After karyogamy: one diploid nucleus442
After premeiotic DNA replication484
Completion of meiosis I484
Completion of meiosis II884
Optional DNA replication in four haploid nuclei8168
Optional completed mitoses: eight haploid nuclei16168
Asexual branch: replication in one haploid nucleus242
Asexual branch: two haploid daughter nuclei442
Nuclear boundary
Completed nuclear stageNucleus identityDrawing xDrawing yDrawing radiusSets per nucleus
One selected unreplicated haploid nucleushaploid-start-n1001.11
After plasmogamy: two separate haploid nucleidikaryon-n1001.11
After plasmogamy: two separate haploid nucleidikaryon-n2301.11
After karyogamy: one diploid nucleusdiploid-n1001.12
After premeiotic DNA replicationreplicated-diploid-n1001.12
Completion of meiosis Imeiosis-i-n1001.11
Completion of meiosis Imeiosis-i-n2301.11
Completion of meiosis IImeiosis-ii-n1001.11
Completion of meiosis IImeiosis-ii-n2301.11
Completion of meiosis IImeiosis-ii-n3601.11
Completion of meiosis IImeiosis-ii-n4901.11
Optional DNA replication in four haploid nucleipostmeiotic-copy-n1001.11
Optional DNA replication in four haploid nucleipostmeiotic-copy-n2301.11
Optional DNA replication in four haploid nucleipostmeiotic-copy-n3601.11
Optional DNA replication in four haploid nucleipostmeiotic-copy-n4901.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n1001.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n2301.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n3601.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n4901.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n5031.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n6331.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n7631.11
Optional completed mitoses: eight haploid nucleipostmeiotic-mitosis-n8931.11
Asexual branch: replication in one haploid nucleusasexual-copy-n1001.11
Asexual branch: two haploid daughter nucleiasexual-mitosis-n1001.11
Asexual branch: two haploid daughter nucleiasexual-mitosis-n2301.11
Centromere-counted chromosome
Nucleus identityModel identityChromatid copiesCentromere x within nucleusCentromere y within nucleusChromatid half-lengthSister half-widthChromosome representation
haploid-start-n1haploid-start-n1-chr11-0.2100.30.08One chromatid
haploid-start-n1haploid-start-n1-chr210.2100.30.08One chromatid
dikaryon-n1dikaryon-n1-chr11-0.2100.30.08One chromatid
dikaryon-n1dikaryon-n1-chr210.2100.30.08One chromatid
dikaryon-n2dikaryon-n2-chr11-0.2100.30.08One chromatid
dikaryon-n2dikaryon-n2-chr210.2100.30.08One chromatid
diploid-n1diploid-n1-chr11-0.6300.30.08One chromatid
diploid-n1diploid-n1-chr21-0.2100.30.08One chromatid
diploid-n1diploid-n1-chr310.2100.30.08One chromatid
diploid-n1diploid-n1-chr410.6300.30.08One chromatid
replicated-diploid-n1replicated-diploid-n1-chr12-0.6300.30.08Two sister chromatids
replicated-diploid-n1replicated-diploid-n1-chr22-0.2100.30.08Two sister chromatids
replicated-diploid-n1replicated-diploid-n1-chr320.2100.30.08Two sister chromatids
replicated-diploid-n1replicated-diploid-n1-chr420.6300.30.08Two sister chromatids
meiosis-i-n1meiosis-i-n1-chr12-0.2100.30.08Two sister chromatids
meiosis-i-n1meiosis-i-n1-chr220.2100.30.08Two sister chromatids
meiosis-i-n2meiosis-i-n2-chr12-0.2100.30.08Two sister chromatids
meiosis-i-n2meiosis-i-n2-chr220.2100.30.08Two sister chromatids
meiosis-ii-n1meiosis-ii-n1-chr11-0.2100.30.08One chromatid
meiosis-ii-n1meiosis-ii-n1-chr210.2100.30.08One chromatid
meiosis-ii-n2meiosis-ii-n2-chr11-0.2100.30.08One chromatid
meiosis-ii-n2meiosis-ii-n2-chr210.2100.30.08One chromatid
meiosis-ii-n3meiosis-ii-n3-chr11-0.2100.30.08One chromatid
meiosis-ii-n3meiosis-ii-n3-chr210.2100.30.08One chromatid
meiosis-ii-n4meiosis-ii-n4-chr11-0.2100.30.08One chromatid
meiosis-ii-n4meiosis-ii-n4-chr210.2100.30.08One chromatid
postmeiotic-copy-n1postmeiotic-copy-n1-chr12-0.2100.30.08Two sister chromatids
postmeiotic-copy-n1postmeiotic-copy-n1-chr220.2100.30.08Two sister chromatids
postmeiotic-copy-n2postmeiotic-copy-n2-chr12-0.2100.30.08Two sister chromatids
postmeiotic-copy-n2postmeiotic-copy-n2-chr220.2100.30.08Two sister chromatids
postmeiotic-copy-n3postmeiotic-copy-n3-chr12-0.2100.30.08Two sister chromatids
postmeiotic-copy-n3postmeiotic-copy-n3-chr220.2100.30.08Two sister chromatids
postmeiotic-copy-n4postmeiotic-copy-n4-chr12-0.2100.30.08Two sister chromatids
postmeiotic-copy-n4postmeiotic-copy-n4-chr220.2100.30.08Two sister chromatids
postmeiotic-mitosis-n1postmeiotic-mitosis-n1-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n1postmeiotic-mitosis-n1-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n2postmeiotic-mitosis-n2-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n2postmeiotic-mitosis-n2-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n3postmeiotic-mitosis-n3-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n3postmeiotic-mitosis-n3-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n4postmeiotic-mitosis-n4-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n4postmeiotic-mitosis-n4-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n5postmeiotic-mitosis-n5-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n5postmeiotic-mitosis-n5-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n6postmeiotic-mitosis-n6-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n6postmeiotic-mitosis-n6-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n7postmeiotic-mitosis-n7-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n7postmeiotic-mitosis-n7-chr210.2100.30.08One chromatid
postmeiotic-mitosis-n8postmeiotic-mitosis-n8-chr11-0.2100.30.08One chromatid
postmeiotic-mitosis-n8postmeiotic-mitosis-n8-chr210.2100.30.08One chromatid
asexual-copy-n1asexual-copy-n1-chr12-0.2100.30.08Two sister chromatids
asexual-copy-n1asexual-copy-n1-chr220.2100.30.08Two sister chromatids
asexual-mitosis-n1asexual-mitosis-n1-chr11-0.2100.30.08One chromatid
asexual-mitosis-n1asexual-mitosis-n1-chr210.2100.30.08One chromatid
asexual-mitosis-n2asexual-mitosis-n2-chr11-0.2100.30.08One chromatid
asexual-mitosis-n2asexual-mitosis-n2-chr210.2100.30.08One chromatid
Nuclear boundaries change the count's meaning
FromToRelationship
One selected unreplicated haploid nucleusAfter plasmogamy: two separate haploid nucleiA second compatible haploid contribution joins; nuclei remain separate
After plasmogamy: two separate haploid nucleiAfter karyogamy: one diploid nucleusTwo nuclei fuse
After karyogamy: one diploid nucleusAfter premeiotic DNA replicationDNA copies; chromosome sets remain two
After premeiotic DNA replicationCompletion of meiosis IHomologues separate into two nuclei
Completion of meiosis ICompletion of meiosis IISisters separate into four nuclei
Completion of meiosis IIOptional DNA replication in four haploid nucleiOptional copying before mitosis
Optional DNA replication in four haploid nucleiOptional completed mitoses: eight haploid nucleiOptional eight haploid nuclear products
One selected unreplicated haploid nucleusAsexual branch: replication in one haploid nucleusCopying in the asexual branch
Asexual branch: replication in one haploid nucleusAsexual branch: two haploid daughter nucleiTwo haploid nuclear products
Selected route
Selected routeCompleted nuclear stage
Selected sexual routeOne selected unreplicated haploid nucleus
Selected sexual routeAfter plasmogamy: two separate haploid nuclei
Selected sexual routeAfter karyogamy: one diploid nucleus
Selected sexual routeAfter premeiotic DNA replication
Selected sexual routeCompletion of meiosis I
Selected sexual routeCompletion of meiosis II
Optional postmeiotic routeOptional DNA replication in four haploid nuclei
Optional postmeiotic routeOptional completed mitoses: eight haploid nuclei
Separate asexual routeOne selected unreplicated haploid nucleus
Separate asexual routeAsexual branch: replication in one haploid nucleus
Separate asexual routeAsexual branch: two haploid daughter nuclei
1C: DNA of one unreplicated haploid nucleus
Assigned quantityValue
Chromosomes in the assigned haploid set2

One complete chromosome set per nucleus

n + n: two separate nuclei

2n: two sets within one nucleus

Centromere-counted chromosome

Two sister chromatids in one chromosome

Totals across the displayed nuclei

Nuclear boundary

Assigned glyph positions; no physical scale

Nuclear products; final spore packaging unspecified

Assigned set of two chromosomes, completed nuclear stages and selected routes; no organelle DNA, whole-mycelium nuclear count, real genome size, anaphase counts, crossover pattern or universal spore packaging is represented.

03

Count nuclear divisions before counting spores

The four haploid nuclei produced by this meiosis are not automatically four final spores in every fungal route. A subsequent mitotic division can produce eight haploid nuclei before packaging. Other routes have different packaging or survival. The model includes an optional postmeiotic branch to show why a count of meiotic products and a count of released spores answer different questions.

If four 1C haploid nuclei each replicate, the total DNA rises from 4C to 8C. Their mitotic divisions can then yield eight 1C haploid nuclei. The total remains 8C after division. Producing eight nuclei without the preceding DNA copying would fail the material account. This option is not an extra round of meiosis and does not halve ploidy again.

The separate asexual branch begins with one unreplicated haploid nucleus. Replication raises its DNA from 1C to 2C; mitosis produces two nuclei, each haploid and 1C. Mutation can occur during asexual lineages, so mitotic reproduction does not guarantee forever-identical descendants. It does avoid the particular joining and meiotic-segregation route represented in the sexual branch.

04

Mating type is a compatibility system

Mating types often regulate recognition and compatibility among fungal partners. They do not imply a universal pair of visibly male and female fungal bodies. Systems vary, including the number of compatibility factors and whether partners are needed in the same way. Compatibility under one rule is also not a complete species-delimitation test.

The four-isolate model assigns A and B to type I, and C and D to type II. Its explicit rule allows only unlike types, excludes self-pairs and counts unordered pairs once. The compatible pairs are A–C, A–D, B–C and B–D. These are possibilities, not observed mating frequencies. A–B and C–D fail this particular rule.

There are six unordered distinct pairs among four isolates: 4 × 3 ÷ 2. Four meet the rule and two do not. That fraction, 4/6, describes the list of eligible pairs. It becomes a probability only after specifying a sampling rule, such as choosing uniformly from those six pairs. It is not a universal fungal mating success rate.

Four compatible pairs under one rule

Four compatible pairs under one ruleConnections compare all six unordered distinct pairs of four isolates, with only unlike assigned types eligible.ABCDIsolate A · IIsolate B · IIsolate C · IIIsolate D · IIEligible under the assigned ruleIneligible under the assigned ruleA–B · Ineligible under the assigned ruleA–C · Eligible under the assigned ruleA–D · Eligible under the assigned ruleB–C · Eligible under the assigned ruleB–D · Eligible under the assigned ruleC–D · Ineligible under the assigned ruleEligible pair count: 4 / 6Probability assumes uniform choice among sixpairsEligibility is not observed reproductive success
The eligibility rule is supplied. The graph distinguishes possible partners from measured mating success.
Read the diagram in words

A and B have type I; C and D have type II. A–C, A–D, B–C and B–D are eligible. A–B and C–D are ineligible. Uniform choice among six unordered distinct pairs gives four eligible choices, or 2/3.

Four compatible pairs under one rule
Assigned isolateAssigned mating typeDrawing xDrawing y
Isolate AI00
Isolate BI03
Isolate CII50
Isolate DII53
Four compatible pairs under one rule
Unordered distinct pairEligibility under the rule
A–BIneligible under the assigned rule
A–CEligible under the assigned rule
A–DEligible under the assigned rule
B–CEligible under the assigned rule
B–DEligible under the assigned rule
C–DIneligible under the assigned rule
Four compatible pairs under one rule
Assigned quantityValue
Only unlike types eligibleYes
Self-pairs allowedNo
Pairs counted without orderYes
Eligible pair count4
Total distinct pair count6
Uniform-choice eligibility probability0.6666666666666666

Assigned type I

Assigned type II

Eligible under the assigned rule

Ineligible under the assigned rule

Probability assumes uniform choice among six pairs

Eligibility is not observed reproductive success

Original compatibility rule with two assigned types and no self-pairs; the rule is not a universal fungal system, observed success rate or complete species-delimitation test.

Worked example

Same total DNA, different organisation

Compare the n + n compartment immediately after joining with the diploid nucleus immediately after nuclear fusion. Both have 2C of DNA in this model, and both contain four chromosome copies in total. The first has two nuclear boundaries, each enclosing one set. The second has one nuclear boundary enclosing both sets.

A statement that 'the cell has 2C' therefore cannot identify ploidy without nuclear organisation and replication stage. A replicated haploid nucleus would also contain 2C, but have two chromosomes and four chromatids. Specify the counted unit, chromosome sets and DNA-copy stage together.

Check your understanding

  1. Why is an n + n compartment not the same as a 2n nucleus?

    Show reasoning

    The n + n compartment retains two separate haploid nuclei; the 2n nucleus contains both complete sets within one nuclear boundary after karyogamy. Their total DNA can be equal while their nuclear organisation differs.

  2. At completion of meiosis I in the model, what does each nucleus contain?

    Show reasoning

    Each is haploid, with two replicated chromosomes, four chromatids and 2C. Homologues have separated; sister chromatids remain together at this completed stage. Meiosis II then produces nuclei with two chromosomes, two chromatids and 1C each.

  3. Uniformly choose one of the six unordered distinct pairs in the compatibility model. What fraction is eligible, and what does that establish?

    Show reasoning

    Four of six pairs are eligible, giving 2/3 under the specified uniform sampling rule. It establishes compatibility by the assigned rule, not actual mating success, reproductive output or a universal rule for species boundaries.

What to carry forward

Follow cytoplasm, nuclear boundaries, chromosome sets and DNA copies separately; interpret a spore or mating type within its specific reproductive route.

References

Resources supporting this lesson’s explanations and examples.

  1. An Overview of the Function and Maintenance of Sexual Reproduction in Dikaryotic Fungi (Frontiers in Microbiology 9:503 (2018); doi:10.3389/fmicb.2018.00503)R. M. Wallen and Michael H. PerlinSpecialist reviewpmc.ncbi.nlm.nih.govRead source
  2. Fungal evolution: diversity, taxonomy and phylogeny of the Fungi (Biological Reviews 94(6):2101–2137 (2019); doi:10.1111/brv.12550)Miguel A. Naranjo-Ortiz and Toni GabaldónSpecialist reviewonlinelibrary.wiley.comRead source