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A progression of “Nothing in biology makes sense except in the light of .” EVOLUTION (Theodosius Dobzhansky, 1973)

4thC BCE 9th-13thC 18th-19thC 1859 1860s 1880-1920 1900-1960 1960s-present 2000-present onwards Middle-Eastern With the Enlightenment in Europe, science ’s 'On the origin of Gregor Mendel proposes the unit of The eclipse of : The modern synthesis and its consequences: The flowering of evolution: Evolution as universal explanation: Why sex? Functional genomics: scholars outline looks for naturalistic explanations for the ' is first published, becoming the heredity now called the gene – but Evolution largely accepted, Theorists formulate mathematical descriptions Theories proliferate as scientists Evolutionary concepts expand into the Evolutionary Science mines the genome Ancient evolution-like diversity of life. Evolution is first mentioned. cornerstone of evolutionary theory. doesn't make the link with evolution. but no-one sure yet exactly of evolution combining ideas from both Darwin dig deeper into evolutionary mechanisms. study of behaviour with implications for biology turns its for answers to a wide range philosophers theories centuries how it works. and Mendel. understanding human nature. collective head of questions. speculate on before Darwin – to this fascinating the origin of life. but lack evidence. question.

Epigenetics The study of inherited, but non-genetic, Embryo changes to the Mendel’s peas Recapitulation observations Hox gene expression of Crossing white- Ontogeny recapitu- “The embryo of a paradox genes. Lamarck and purple- lates phylogeny: an vertebrate is already may have been organism’s devel- a vertebrate” – so more right produces 3 times opmental path to are very different, ontogeny does not yet some of our than we thought. as many purple adult-hood retraces recapitulate plants. Why? the evolutionary genes are phylogeny. remarkably similar. Genomics of Because heredity path of its species. (Karl Ernst von Baer) (Ernst Haeckel) Struggle for units occur in Except Although Part of the adaptation dominant- Hopeful Punctuated existence Is adaptation mainly recessive pairs. monster theory equilibrium due to in “Environmental Where present – Mendelian Macromutation New species arise Evolution proceeds genes very similar Earth is old Lamarckian 75% of time – the inheritance due to large Ploidy Evo-devo among species, or organisms dominant purple theory Evidence from evolution Offspring inherit mutations – New species can with long periods of synthesis due to genes that to develop new Variation emerges rocks shows the Giraffes stretch their one genetic variant ’hopeful monsters’ form when stasis between The great diversity characteristics to earth is more than recessive white via mutations. Large necks striving to from each parent, () genomes double. bursts of speciation. of life forms is proteins? ensure survival, thus a few thousand mutations result in Fisher’s reach higher leaves. independently for But also The majority For example the based on similar By means of transforming into Uniformitarian- (Gregor Mendel) new species. years old. Their offspring each gene, if they horseshoe crab has new species. (, microscope developmental (James Hutton) ism inherit longer Led to theory of are not linked. remained the same Animals that survive Otto Schindewolf) As the population have undergone genes that are Extraordinary We now know the The earth as a necks. for millions of years. to breed can pass moves towards the Ecological some genome expressed earth was formed whole is subject (Jean Baptiste Lamarck) Or was he? duplication. (Niles Elredge and differently in sex ratios Functional on their successful Mendelism optimum Stephen J. Gould) 4.54 billion years to gradual change. (V. Grant, after Linnaeus) Why most species Arrows characteristics But Lamarck was “The most important phenotype, Genetics goes out distantly related Genomics ago. (Charles Lyell) wrong because have a near-1:1 sex to offspring.” mutations get organisms. What do different between theories factor of evolution of the lab and into ratio, but some (Al Jahith, Book of Animals, is the occurrence smaller and smaller, genes do? For 9thC) insects and reptiles example, the OCA2 all types, from direct Sounds a lot like of advantageous observe evolution Trade-off have many more mutations [...] microscope. in action. For gene encodes a (R. A. Fisher) theory males or females. Contrary to example, when the protein involved in intellectual Monkey Man is Because resources (W. D. Hamilton) Darwinian Principle of merely a sieve to environment producing a kinship. Tested in the wild are limited, ancestry descended Mendel’s peas: becomes more pigment that affects medicine variation save advantageous organisms must Humans developed mutations and polluted, pepper hair and eye colour. Tailor-made medicine from primates The least resurrected make compromises. The study of animal Two-fold cost from “the world of Natural eliminate moths with the based on personal Ape is “brother Transmutation well-adapted Mendel’s pea For example, they instinct and Useful for the monkeys”. selection deleterious genetic variant for of sex of man”. The fossil record individuals are experiments can’t be both behaviour: social (Ibn Khaldun, (Lord Monboddo) Any advantageous mutations.” dark colour survive, A sexual organism Muqaddimah, 1377) shows gradual “most liable to unearthed and fast growers and dominance, tool adaptation (T. H. Morgan) reproduce only passes on half Evidence? change from earlier perish”. replicated. Quantitative good competitors. manufacture, and (James Hutton) Disagreed with and become its genes to its into later species Blending (Hugo de Vries, (W. D Hamilton) transmission of Genomics woodpecker’s beak) Carl Correns) genetics more common. offspring. What part of (Robert Grant) Inheritance theory cultural traits. causing an organism Variations in (Bernard Kettlewell) (Maynard-Smith) of plasticity Daoism One generation (, our genome a) via a mysterious to leave more Offspring are a phenotype are the Criticised in How can some Nature is in constant Natural passes on its Nico Tinbergen) makes us ’internal force’ offspring than others result of blending Biometrics result of genes + individuals adapt to transformation or characteristics Gemmules (Jean Baptiste Lamarck) selection: will become more of parents’ body The statistical environment. Paved different human? ‘Dao’. Species can Change over to the next. Charles Darwin’s Germ plasm the prequel common over time. How? types. 1 tall parent study of continuous the way for modern environments change in response b) as a direct result hypothetical generations Erasmus Darwin, Gradually, the whole + 1 short parent = theory characters like evolutionary theory without being to the environment. of the environment heredity particles, Natural changes grandfather of population changes. medium height height and weight and population The spandrels Why do genetically distinct? (Laozi) (Etienne Geoffroy St-Hilaire) found in the blood. accumulate over Charles Darwin and (Charles Darwin) offspring. cut the tails off 1500 to understand the genetics. Shifting Some genetic of San Marco organisms generations, (Charles Darwin) Unit later found in rats, but after 20 link between genes (R. A. Fisher) Francis Galton: “The Eureka! But by what means? balance theory variants are bad, Not everything is an have sex? producing new strongest and most How does this work? But generations all were (genotype) and the resulting in lower Evolution can adaptation. Like the races and species. active animal still born with tails. way something produce different spandrels of a (Pierre Louis Maupertuis) should propagate Conclusion: only looks (phenotype). of bad variation in a a) Red Queen Tree of Life solutions to the cathedral dome, the species, which special ‘germ plasm’ (Francis Galton, Raphael population is the parasite Weldon, Karl Pearson) same environmental evolution can web project should thence cells are responsible genetic load, and Principle of challenges. be constrained by hypothesis All the world’s for heredity, not natural selection Ladder of life become improved”. (Sewall Wright) structural factors or living diversity, now (Zoönomia, 1794) normal body cells. acts to get rid of it. Life forms are historical genes, making it chronicled online at and analogy However (Hermann Joseph Muller But how do we explain hierarchically Social and J.B.S Haldane) circumstance. easier to adapt to tolweb.org/tree/. Organisms share such high variation? organised – (Stephen J. Gould and pathogen attacks – But can we similarities and Survival of Darwinism Richard Lewontin) complex organisms Molecular until the pathogens conserve it? evolve naturally The application of are nearer the top bite back. along these lines. Principle of evolutionary ideas evolution of the ladder. Dobzhansky- Paleontology (J. Jaenike, W. D. Hamilton) (Richard Owen) divergence Galton’s to welfare. Inspired Including: (Plato, Aristotle) are those most Described in rabbits – the idea Muller model meets evolution Price’s Christians: God able to survive. b) The Hill- Darwin’s Rabbit experiments that the human race Genetic changes Did species on a) Neutral equation is at the top. (Herbert Spencer, could be genetically built up in different different continents Robertson after reading Malthus) autobiography as show that theory A character, such as ‘improved’. populations have diverge via genetic Systems “the tendency in ‘gemmules’ Random processes height, changes effect of species. (Herbert Spencer, no guarantee of drift? No: the fossil organic beings do not exist in the Francis Galton) like genetic drift depending on the biology >10,000 named descended from working well record shows Game theory blood and cannot MORALLY CONDEMNED contribute to strength of genes, allowing for The holistic by Carl Linnaeus. together. Explains mammals migrated the same stock to be transferred by genetic change. selection and how Economic models application why some hybrids across land bridges. Levels of diverge in character transfusions. Many mutations are well this character is applied to the mutations to of evolutionary like mules are (George Gaylord Simpson) natural as they become (Francis Galton) neutral – neither transmitted from evolution of co-occur, and theory to all levels sterile, or can Geographic good nor bad. selection one generation behaviour, evolution to of a system – from even die. Separation Competition for speciation (Motoo Kimura) A heated debate to the next. including human happen faster. molecules up Spontaneous (Theodosius Dobzhansky, Sociobiology resources is (Bill Hill, Alan Robertson) theory Geographic Hermann Muller) Haldane’s rule about whether (George R. Price) Used in behaviour. to ecosystems. generation Predicts (John Maynard-Smith) The same Biogeography Small colonies of a stronger between barriers like oceans Sterile or inviable b) Nearly natural Living things close relatives, so evolutionary No need for Different regions population migrate and mountains can selection acts... self-generate from species push each neutral theory principles apply have different to a different region, isolate populations hybrids are more non-living matter. God other apart. Mutations can be to social behaviour species. eventually forming Developed into from one another. likely to have two a) Sexual (Aristotle) Denis Diderot’s slightly deleterious; a) on the gene in organisms from (Comte de Buffon) new species. Over time, the different sex ’Letter on the Blind’ Why? the relative power ‘Gene’s eye view’ ants to apes to Mating Magic! Or maybe not (Moritz Wagner) populations : in Red Queen (1749) argued for of drift and (George C. Williams, It serves one sex to slime moulds. systems diverge into mammals, XY not XX, Richard Dawkins) natural evolution selection depends coevolution harm the other (E. O. Wilson) separate species. making them male. a) Male and female on population size. acting via the Each species during or after without God – () (J.B.S. Haldane) in the same (Tomoko Ohta) individual must keep ‘running’ mating. For Who’s the designer? landing its author organism (some Natural Phylogenetics (Ernst Mayr) (adapting via example, some in prison. plants, snails). theology Study of the shared c) Selectionism or family natural selection) to female spiders eat evolutionary history (W.D Hamilton) keep pace with its their mates. Irreducible Natural selection is b) One sex then the Numerical Species of species, competitors. (Robert Trivers, Geoff Parker) complexity of e.g. the dominant force other: when a Paley’s watch taxonomy represented on () human eye spells concepts in evolution. b) on groups dominant female Classification of branching ‘tree of And on a smaller scale b) Reciprocal design. Design Like an exquisitely What is a species? (Theodosius Dobzhansky, of individuals designed watch, the organisms based life’ diagrams called E. B. Ford, Ernst Mayr) Evolutionary requires a Group selection altruism a male changes sex world is too on form and phylogenies. designer – God. a) Biological (V. C. Wynne-Edwards) I’ll scratch your back psychology to replace her. complex to not structure, regardless (Willi Hennig) d) Molecular if you scratch mine, Human nature is have a creator. of shared concept clocks even if we aren’t the result of c) Separate male (William Paley) Organisms of c) on species evolutionary history. family. and female sexes (Peter Sneath, Robert Sokal) DNA and protein Species selection evolutionary different species (Robert Trivers) sequences can be (Stephen Jay Gould) adaptations (mammals). Ignores Sympatric cannot interbreed. (Ernst Mayr) used to calculate affecting our But Physiological speciation when species split d) on more c) Parent- psychological Malthusianism selection Individuals of the b) Ecological off from others. offspring makeup. Nature limits same population (Marcel Florkin, Emil Smith) than one of Variation accum- Convergent CONTROVERSIAL TODAY population size via develop concept the above ulates in different evolution Sibling competition disease, war, preferences Different species Multi-level How many populations. At one The same traits starvation, and for certain habitats occupy different selection theory partners? point, individuals evolve repeatedly offspring can take other tragedies. or foods. These niches, or can only breed with and independently. it out of parents. – Most birds pair ( Malthus) preferences are specialised Wings evolved (Robert Trivers) bond for life members of their inherited, and the environments. three times in birds, because raising group. The pop- different types no (Leigh van Valen) ulation splits into bats and bees. young is hard work. longer interbreed. Memetics But cheating is different species. Could explain the c) Phylogenetic Catastrophism (George Romanes) still common. huge diversity of Cultural ideas or Evolution is not concept – In most mammals ’memes’ (concepts, gradual. Fixed All members of a the male mates with Runaway African lakes. stories, tunes) are a species arise and species come from many females sexual (John Maynard Smith) lot like genes: they are perfectly Sexual same branching replicate, mutate because selection point on reproduction is adapted to their selection and are subject environment, but Females prefer evolutionary ‘tree’. Handicap to selective easy for him, but Differences (Joel Cracraft) can also go extinct. showy males, which principle pressure. hard for her. (Georges Cuvier) between males’ leads them to select (Luigi Cavalli-Sforza (S. T. Emlen, L. W. Oring) ability to father exaggerated sexual The peacock and Marcus Feldman, offspring result in ornaments like the evolved an Richard Dawkins) physically different peacock’s long tail. impractically long sexes – the peacock (R. A. Fisher) tail – to show the Intelligent and the peahen. peahens his ‘good design (Charles Darwin) genes’ mean he Natural theology But why the long tail? can afford one. (Amotz Zahavi) But pretending to be science. Creationism dies hard. (Michael Behe)

4thC BCE onwards 9th-13thC 18th-19thC 1859 1860s 1880-1920 1900-1960 1960s-present 2000-present

Just four genetic Step by step, from The four elements How life is Ecosystem Sources/ processes govern the simple to wildly Community Species A dynamic complex of proliferation of species complex Population Populations of different biological of evolution organised Organism different species that communities – plants, credits Cell Groups of organisms One is ‘a group of live in the same area animals and microbes – Collections of cells that living in the same area organisms that can and interact via in interaction with the Darwin, C. (1859). by Genome The smallest unit of collaborate to form a that can interbreed interbreed and create competition, predation surrounding environment means of natural selection: or the preservation of favoured races in the struggle for life. : 1. Gene A threadlike structure life. Most cells have larger entity such as viable offspring’ and parasitism John Murray DNA All an organism’s DNA containing the DNA one or two copies of an animal or plant The billions of bacteria 2. living in your gut form Unit of heredity that molecule and There are estimated to be each chromosome their own ecosystem. Ridley, M. (2004). Evolution. 3rd edition. Oxford: Natural nearly 9 million species on 3. Four nucleotides The human genome is supporting proteins Recent research suggests codes for proteins or earth, of which only just Blackwell Science Ltd. made up of 6 billion DNA human gut ecosystems Drift that form the building RNA within an organism bases split into 24 Some ants have only one over 1 million (about 15%) Selection 4. of each chromosome in form into three main chromosomes. It contains Human chromosome have been named. blocks of most their cells, while some types dominated by Ruse, M. and Travis J.. eds. (2009). Evolution: about 25,000 genes. number 1 contains 220 Not all parts of a gene plants have 12. particular species. The change in a genetic variant living organisms million nucleotides. . Harvard: Harvard Genetic variants linked to have to be on the same from one generation to the next Migration stretch of DNA or even on University Press. advantageous traits (such as height, Genes undergo accidental resistance to disease) cause an due to chance – and not to Stretched out, the DNA the same chromosome. changes – or mutations – Did you molecules in a single Research: Tania Jenkins, Miriam Quick natural selection The process by which genes human cell would be organisms to leave more offspring caused by , viruses, know? Design: Stefanie Posavec or gene variants transfer from about 2 metres long. or simple copying errors one population to another Kindly funded by a grant from the European Society for http://www.eseb.org/

Thanks to Sarah Otto, Tadeusz Kawecki and Christine Clavien for comments on drafts