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Evolution

"It is not the strongest of the species that survives... but the one most responsive to change ." — apocryphally attributed to Darwin (actually Megginson...

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"It is not the strongest of the species that survives... but the one most responsive to change ." — apocryphally attributed to Darwin (actually Megginson, 1963), but the spirit is right. Key sections include: Evolution One law. Four billion years.; One algorithm , three ingredients.; Charles Darwin, 1809–1882.; Five forces that shape allele frequencies.; Lines of convergent evidence.; One universal ancestor.; How one becomes two.; Who built the theory.; Evolution in equations.; Of life , in years..

Key sections

  • 01Evolution One law. Four billion years.
  • 02One algorithm , three ingredients.
  • 03Charles Darwin, 1809–1882.
  • 04Five forces that shape allele frequencies.
  • 05Lines of convergent evidence.
  • 06One universal ancestor.
  • 07How one becomes two.
  • 08Who built the theory.
  • 09Evolution in equations.
  • 10Of life , in years.
  • 11The Big Five.
  • 12The hominin branch.
  • 13And the extended one.
  • 14Where the field moves.
  • 15Still open.
  • 16Watch & read.
  • 17Endless forms most beautiful.

Topics covered

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Slide outline
  1. 01Evolution One law. Four billion years.
  2. 02One algorithm , three ingredients.
  3. 03Charles Darwin, 1809–1882.
  4. 04Five forces that shape allele frequencies.
  5. 05Lines of convergent evidence.
  6. 06One universal ancestor.
  7. 07How one becomes two.
  8. 08Who built the theory.
  9. 09Evolution in equations.
  10. 10Of life , in years.
  11. 11The Big Five.
  12. 12The hominin branch.
  13. 13And the extended one.
  14. 14Where the field moves.
  15. 15Still open.
  16. 16Watch & read.
  17. 17Endless forms most beautiful.
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Slide 01

One algorithm, three ingredients.

  • P. 01 / 17 — the principle
  • Charles Darwin's central idea was almost embarrassingly simple. Wherever there is variation, inheritance, and differential reproduction, populations must change over generations.
  • Variation
  • Individuals differ. Mutations, recombination, and developmental noise generate the raw material on which selection acts.
  • Inheritance
  • Offspring resemble parents more than they resemble strangers. Mendelian particles, since 1900; genes, since the 1920s.
  • Differential reproduction
  • Some variants leave more offspring than others. The world cannot support all that is born.
  • Given those three, evolution is mathematically inevitable.
Slide 02

Charles Darwin, 1809–1882.

  • P. 02 / 17 — Darwin
  • Failed in medicine at Edinburgh, took holy orders at Cambridge instead. In 1831, at 22, sailed as gentleman naturalist on HMS Beagle's five-year survey voyage. The experience — fossils in Patagonia, finches and tortoises on the Galápagos, coral atolls in the Pacific — convinced him species are not fixed.
  • He read Malthus on population in 1838 and the puzzle's last piece fell into place. He sat on the theory for twenty years, terrified of the consequences. In 1858 a letter from Alfred Russel Wallace, working independently in Indonesia, forced his hand. They presented jointly at the Linnean Society. On the Origin of Species followed in November 1859 — sold out in a day.
  • Galápagos finches
  • — sketch after Darwin, 1845
Slide 03

Five forces that shape allele frequencies.

  • P. 03 / 17 — mechanisms
  • Mutation
  • Random changes to DNA. Rate ~10⁻⁸–10⁻⁹ per base per generation in mammals.
  • Selection
  • Differential survival/reproduction. Natural, sexual, kin, group. Directional, stabilizing, disruptive.
  • Drift
  • Random changes due to finite population. Dominates when Ne is small or selection is weak.
  • Gene flow
  • Migration mixes populations. Counteracts local adaptation.
  • Non-random mating
  • Inbreeding, assortative mating. Changes genotype frequencies even without changing alleles.
  • Δq ≈ spq(q−q̄) under selection · σ²q = pq/2Ne per gen drift
Slide 04

Lines of convergent evidence.

  • P. 04 / 17 — evidence
  • Fossil record. Smooth morphological transitions: Tiktaalik (375 Ma) between fish and tetrapods; Archaeopteryx (150 Ma) between dinosaurs and birds; Pakicetus, Ambulocetus toward whales.
  • Comparative anatomy. Homologous limbs (human arm = whale fin = bat wing). Vestigial structures (whale pelvis, human appendix).
  • Embryology. Pharyngeal arches in all vertebrate embryos. Hox-gene patterning conserved across the animal kingdom.
  • Biogeography. Marsupial radiation in Australia; Galápagos endemism. Distributions trace plate movement and colonization history.
  • Genetics. Universal genetic code. Pseudogenes shared with relatives. Endogenous retroviruses at identical loci in primates.
  • Direct observation. Antibiotic resistance, peppered moths, Italian wall lizards on Pod Mrčaru in 36 yr (Herrel et al., 2008).
  • Pentadactyl limb — homology
  • Same bones, four jobs.
Slide 05

One universal ancestor.

  • P. 05 / 17 — the tree
  • A schematic, not to scale or completeness.
  • All living organisms share the same genetic code, the same handedness of biological molecules, and large blocks of homologous genes. The simplest explanation: they share an ancestor. The Last Universal Common Ancestor (LUCA) lived ~3.8 Gya, perhaps near hydrothermal vents.
Slide 06

How one becomes two.

  • P. 06 / 17 — speciation
  • Allopatric
  • Geographic isolation. Most common mode. Galápagos finches; Lake Baikal seals; Hawaiian silverswords.
  • Sympatric
  • Divergence in same place — niche partitioning, host shift. Cichlid fish in African rift lakes (~500 species in Lake Malawi alone).
  • Parapatric
  • Adjacent ranges with limited gene flow. Ring species, e.g. Larus gulls around the Arctic.
  • Reproductive isolating mechanisms: prezygotic (mating time, place, behavior, gamete incompatibility) and postzygotic (hybrid inviability, sterility — see the mule).
Slide 07

Plate VII

  • Cabinet of curiosities. The 19th-century museum was the engine of comparative zoology.
Slide 08

Who built the theory.

  • P. 08 / 17 — key figures
  • Lamarck
  • 1744–1829
  • First systematic theory of transmutation. Inheritance of acquired characters — mostly wrong, but he raised the question.
  • Darwin
  • 1809–82
  • Natural selection. Origin of Species, 1859. Descent of Man, 1871.
  • Wallace
  • 1823–1913
  • Co-discoverer of natural selection. Biogeography founder.
  • Mendel
  • 1822–84
  • Particulate inheritance. Critical to the modern synthesis.
  • Fisher · Haldane · Wright
  • 1920s–30s
  • Founders of population genetics. Reconciled Darwin with Mendel.
  • Dobzhansky
  • 1900–75
  • Modern Synthesis architect. "Nothing in biology makes sense except in the light of evolution."
  • Mayr
  • 1904–2005
  • Biological species concept. Systematics and the Origin of Species, 1942.
  • Hamilton
  • 1936–2000
  • Inclusive fitness, kin selection (1964). Hamilton's rule: rB > C.
Slide 09

Evolution in equations.

  • P. 09 / 17 — population genetics
  • Hardy–Weinberg
  • p2 + 2pq + q2 = 1
  • Genotype frequencies in absence of evolution. Deviation diagnoses force at work.
  • Hamilton's Rule
  • rB > C
  • Altruism evolves when relatedness × benefit to relative exceeds cost to actor.
  • Fisher's Theorem
  • Δw̄ = VA / w̄
  • Rate of increase in mean fitness equals additive genetic variance, scaled.
  • Selection coefficient
  • Δq = spq(p−q) ÷ ...
  • Allele frequency change per generation under selection of strength s.
  • Drift variance
  • σ²q = pq/2Ne
  • Random walk in finite population. Effective N ≪ census N usually.
  • Price equation
  • Δz̄ = Cov(w, z)/w̄ + E(wΔz)/w̄
  • George Price (1970): unifies all evolutionary change.
Slide 10

Of life, in years.

  • P. 10 / 17 — timeline
  • ~3.8 GyaEarliest chemical signatures of life — graphite in Greenland metasediments.
  • ~3.5 GyaStromatolites; cyanobacteria.
  • ~2.4 GyaGreat Oxygenation Event.
  • ~2.0 GyaEukaryotic origin via endosymbiosis (mitochondria).
  • ~1.5 GyaSexual reproduction.
  • ~570 MyaEdiacaran biota — first multicellular animals.
  • ~540 MyaCambrian explosion — most modern phyla appear.
  • ~440 MyaPlants colonize land.
  • ~375 MyaTetrapods: Tiktaalik et al.
  • ~330 MyaAmniotes — life freed from water for reproduction.
  • ~250 MyaPermian–Triassic extinction. ~96% of marine species lost.
  • ~150 MyaBirds branch from theropod dinosaurs.
  • ~66 MyaK–Pg extinction. Mammals radiate.
  • ~6 MyaHominin–chimpanzee LCA.
  • ~300 kyaAnatomically modern Homo sapiens.
  • ~12 kyaDomestication of plants and animals.
Slide 11

The Big Five.

  • P. 11 / 17 — mass extinctions
  • Ordovician–Silurian, ~445 Mya. ~85 % of species. Glaciation, sea-level fall. Marine.
  • Late Devonian, ~375 Mya. ~75 %. Multiple pulses, anoxia.
  • Permian–Triassic, ~252 Mya. ~96 % marine, ~70 % terrestrial. Siberian Traps volcanism. "The Great Dying."
  • Triassic–Jurassic, ~201 Mya. ~80 %. Volcanism (CAMP), ocean acidification.
  • Cretaceous–Paleogene, ~66 Mya. ~76 %. Chicxulub impact (Yucatán) plus Deccan Traps.
  • Sixth. The Holocene/Anthropocene biodiversity crisis: extinction rates 100–1,000× background. ~1 in 4 mammals threatened. Defaunation of large vertebrates.
  • Genus diversity through time
Slide 12

Slide 12

  • P. 12 / 17 — pull quote
  • "There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one..."— Charles Darwin, On the Origin of Species, 1859
Slide 13

The hominin branch.

  • P. 13 / 17 — human evolution
  • Humans share ~98.7 % of nuclear DNA with chimpanzees, ~96 % with gorillas, ~93 % with orangutans. Our split from chimpanzees was ~6 Mya, in Africa. The fossil record since then is rich and tangled — at least a dozen recognized hominin species, several coexisting.
  • Sahelanthropus tchadensis — ~7 Mya · Chad
  • Ardipithecus ramidus — ~4.4 Mya · Ethiopia
  • Australopithecus afarensis — Lucy, ~3.2 Mya
  • Homo habilis — ~2.4 Mya · Olduvai Gorge
  • H. erectus — first hominin out of Africa, ~1.9 Mya
  • H. neanderthalensis — Eurasia, ~400–40 kya
  • H. denisovensis — known mostly from genome
  • H. sapiens — ~300 kya · 1.5–4% Neanderthal admixture in non-Africans
  • Skull volume over time
Slide 14

And the extended one.

  • P. 14 / 17 — modern synthesis
  • The Modern Synthesis (1930s–40s) — Fisher, Haldane, Wright, Dobzhansky, Mayr, Simpson, Stebbins — fused Darwin with Mendelian genetics. Evolution is changes in allele frequencies; speciation is the splitting of gene pools.
  • Since the 1990s, an Extended Evolutionary Synthesis argues for adding: developmental constraints (evo-devo), epigenetic inheritance, niche construction, multilevel selection, plasticity-led evolution. The debate is more about emphasis than mechanism.
  • Evo-devo. Hox gene homology; deep conservation of body plan toolkits.
  • Niche construction. Beavers build dams that select for further damming.
  • Genetic assimilation. Plasticity → fixation (Waddington's flies).
  • Multilevel selection. Selection at gene, individual, group levels (D.S. Wilson, Sober).
  • Horizontal gene transfer. Pervasive in microbes; reticulate trees.
  • Endosymbiosis. Mitochondria, chloroplasts as discrete acquisition events.
Slide 15

Where the field moves.

  • P. 15 / 17 — frontier & open
  • Ancient DNA
  • Svante Pääbo's lab (Nobel 2022) sequenced Neanderthal & Denisovan genomes. Pleistocene mammoth DNA > 1 My old recovered (2021).
  • Experimental evolution
  • Lenski's E. coli LTEE, ~80,000 generations and counting. Citrate utilization evolved at gen ~31,500.
  • De-extinction
  • Mammoth, thylacine, dodo proxies under engineering. Bringing functional ecologies back, not "Jurassic Park".
  • Phylogenomics
  • Whole-genome trees resolve historic disputes; reveal incomplete lineage sorting and introgression.
  • Cultural evolution
  • Henrich; Boyd & Richerson. Dual-inheritance models for human cooperation, language change.
  • Major transitions
  • Maynard Smith & Szathmáry's framework: levels of selection re-organize. From RNA to cells to multicellularity to societies.
  • Still open.
  • What was the chemistry of the origin of life?
  • Why did sex evolve and persist? (The two-fold cost remains a puzzle.)
  • How important is genetic drift vs. selection in shaping genomes?
  • What governs evolvability — the rate at which lineages can adapt?
  • Do mass extinctions follow predictable patterns, or are they contingent?
  • Is the human-cognition transition unique, or repeatable elsewhere?
Slide 16

Watch & read.

  • P. 16 / 17 — go deeper
  • PBS Eons — Deep Time Series
  • Plus David Attenborough's "Life on Earth" (1979) — still unmatched.
  • Watch ↗
  • References
  • Darwin — On the Origin of Species (1859)
  • Dawkins — The Selfish Gene (1976)
  • Gould — The Structure of Evolutionary Theory (2002)
  • Coyne — Why Evolution Is True (2009)
  • Reich — Who We Are and How We Got Here (2018)
  • Maynard Smith & Szathmáry — The Major Transitions in Evolution (1995)
  • Lenski LTEE — > 80 papers, see Nature 2009
Slide 17

Endless forms most beautiful.

  • P. 17 / 17 — coda
  • "Have, and are being, evolved." — closing words, Origin of Species.
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