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History of Genetics in Evolution

Joe Felsenstein

GENOME 453, Autumn 2015

History of Genetics in Evolution – p.1/44 The Great Chain of Being Going back as far as the Ancient Greeks, a linear hierarchy of life forms was assumed, with inanimate objects at the bottom and deities at the top, such as this: Deity Angels Man Birds Reptiles Amphibians Fish Insects Worms Protists Rocks Issues: placement of birds, insects not obvious. A scale of complexity? Or what? Note: can think of it as a “chain”, hence the phrase “missing link”

History of Genetics in Evolution – p.2/44 Carl Linnæus (Carl von Linné) (1707-1778)

The great figure in biological classification. Not a nice man.

History of Genetics in Evolution – p.3/44 Monophyly Monophyletic: having a common ancestor which is not the ancestor of any of the other species being discussed.

(This definition works for cases where there are forms being included, and those where they are not, and works whether we are discussing only a fixed set of species or all species descended from some ancestor.)

History of Genetics in Evolution – p.4/44 Linnæus’s classification of

In the 10th edition of :

Class Mammalia (mammals) Class Aves (birds) Class Amphibia (amphibians and reptiles) Order Reptiles Testudo (turtles) Genus Draco (gliding lizards) Genus Lacerta (lizards, salamanders, crocodiles) Genus Rana (frogs and toads) Order Serpentes (snakes, slowworms, caecilians) Class Pisces (most fishes) Order Nantes (lampreys, rays, sharks, anglerfishes, sturgeons)

Not exactly our current scheme!

History of Genetics in Evolution – p.5/44

Example: A phylogeny of the living Craniata

Salamanders Rays Hagfishes Crocodilians

Birds Turtles Lizards Snakes, Mammals Frogs Lungfish Sharks Lampreys Tunicates Amphioxus

Ray−finned fishes Ray−finned Coelacanths

History of Genetics in Evolution – p.6/44

Vertebrates are a monophyletic group

Salamanders Rays Hagfishes Crocodilians

Birds Turtles Lizards Snakes, Mammals Frogs Lungfish Sharks Lampreys Tunicates Amphioxus

Ray−finned fishes Ray−finned Coelacanths

vertebrates ancestor

History of Genetics in Evolution – p.7/44

Reptiles and fishes are paraphyletic groups

Salamanders Rays Hagfishes Crocodilians

Birds Turtles Lizards Snakes, Mammals Frogs Lungfish Sharks Lampreys Tunicates Amphioxus

Ray−finned fishes Ray−finned Coelacanths

Reptiles

Osteichthyes

History of Genetics in Evolution – p.8/44 An intellectual world changes From the 1600s on, changes of worldview in many fields:

Commercial trade and exploration brings Europeans in contact with animals and plants of other continents

Physics: Galileo and Newton in 1600s discover the laws of motion, a single mechanism for multiple phenomena. Hugely influential.

Astronomy: Comets, galaxies, planets that have geology (Galileo, Thomas Wright, Laplace, William Herschel)

Geology: Strata, sedimentation, and (e.g. Lavoisier, Smith, Hutton, Lyell)

Paleontology: are not artifacts but remains of real organisms, different from the ones we see today (Nicholas Steno, Smith, Cuvier)

Anthropology: Diversity of people, similarity of apes to people

Chemistry: Arbitrary properties of substances replaced by the periodic table and properties of atoms and molecules (Boyle, Lavoisier, Cavendish, Priestley, Mendeleyev)

... and of course politics: the English and French revolutions and the independence of the American colonies All of this led biologists to look for unifying forces and connections between phenomena. As well as to be open to seeing ongoing changes in the natural world. History of Genetics in Evolution – p.9/44 An American in Paris (2005) Wandering east of the Panthéon on the Left Bank of Paris, you begin to notice unusual street names:

A street named for Linnaeus? This only hints at a little-known story.

History of Genetics in Evolution – p.10/44 Buffon

George-Louis Leclerc, Comte de Buffon (1707-1788)

History of Genetics in Evolution – p.11/44 Statue of Buffon at the Jardin des Plantes

History of Genetics in Evolution – p.12/44 Buffon, honored

Rue Buffon, next to the Jardin des Plantes (with plastic mastodon, Golden Arches, traffic ticket)

History of Genetics in Evolution – p.13/44 Lamarck

Jean Baptiste Pierre Antoine de Monet, Chevalier de Lamarck (1744-1829)

History of Genetics in Evolution – p.14/44 Lamarck’s tree

As published in , 1809 Can you identify some of the groups? “M.” means mammals History of Genetics in Evolution – p.15/44 Lamarck’s mechanism for evolution In Philosophie Zoologique, 1809.

Organisms’ characters are altered by the effects of use and disuse. These changes are passed on to descendants by inheritance of acquired characters.

Note that Lamarck did not originate “Lamarckian inheritance": it was something everyone believed in at that time.

History of Genetics in Evolution – p.16/44 Old displays in the Museum of , Paris

History of Genetics in Evolution – p.17/44 Statue of Lamarck in the Jardin des Plantes, Paris

History of Genetics in Evolution – p.18/44 Lamarck’s works listed

History of Genetics in Evolution – p.19/44 “My father, you will be vindicated”

History of Genetics in Evolution – p.20/44 Buffon’s (and Lamarck’s) house next to the Museum

History of Genetics in Evolution – p.21/44 Plaque on house commemorating Buffon

“George Louis Leclerc, Count of Buffon, born in Montbard, 7 September 1707, director of the Royal Botanical Garden from 1739 on, died in this house, 16 April 1788”

History of Genetics in Evolution – p.22/44 Plaque on house commemorating Lamarck

“Jean Baptiste Lamarck, born in Bazentin Le Petit, 1 August 1744, Professor at the Museum, Author of the first theory of evolution, lived in this house from 1795 on, died on the 18th of December 1829”

History of Genetics in Evolution – p.23/44 Geoffroy versus Cuvier

Etienne Geoffroy St. Georges Lèopole Hilaire Chrètien (1772-1844) Frèdèric Dagobert, Baron Cuvier (1769-1832)

History of Genetics in Evolution – p.24/44 Memorials in Paris

Fountainoncornerof RueCuvier, RueLinnéandRueCuvier alongsideoffountain (“Á ”) and side of Jardin des Plantes

History of Genetics in Evolution – p.25/44 Allee Cuvier, within the Jardin

History of Genetics in Evolution – p.26/44 Rue Geoffroy St. Hilaire

History of Genetics in Evolution – p.27/44 Paris: Rue Lamarck and Rue Darwin

History of Genetics in Evolution – p.28/44 Johann Wolfgang von Goethe (1749-1832)

History of Genetics in Evolution – p.29/44 Goethe (1790) on the origin of parts of flowers

History of Genetics in Evolution – p.30/44 The Naturphilosophen

The Naturphilosophen and Evolutionary views

Ape Ape

Monkey Monkey Mouse Mouse

Reptile Reptile Amphibian Amphibian Fish Fish

common developmental pathway evolutionary tree

Note − The picture here is very much a Great Chain of Being

History of Genetics in Evolution – p.31/44 Fossil forms becoming known in 1700s, 1800s

Charles Willson Peale Mary Anning’s Plesiosaur, 1821 The Exhumation of the Mastodon (in 1801, painted 1808)

History of Genetics in Evolution – p.32/44 Robert Chambers discusses evolution in 1844

History of Genetics in Evolution – p.33/44 (1809-1882)

History of Genetics in Evolution – p.34/44 Wallace

Alfred Russel Wallace (1823-1913) in 1869

History of Genetics in Evolution – p.35/44 Lamarck’s theory versus Darwin’s Lamarck Darwin/Wallace Geneticvariationimportant? No Yes Differential survival or reproduction? No Yes Mutationsare inwhat direction? adaptive random Phenotypicchangesinherited? Yes maybe

History of Genetics in Evolution – p.36/44 Fleeming Jenkin

Fleeming Jenkin (1833-1885) Fleeming Jenkin Building University of Edinburgh

History of Genetics in Evolution – p.37/44 Blending inheritance and selection

10

8

4

2

0 0 10 20 30 40 Value of character

History of Genetics in Evolution – p.38/44 The Biometricians

Francis Galton (1822-1911) Karl Pearson (1857-1936)

History of Genetics in Evolution – p.39/44 Gregor Mendel (1822-1884)

History of Genetics in Evolution – p.40/44 Mendel in his school

The faculty of Mendel’s monastery school (Mendel is top center-right with flower)

History of Genetics in Evolution – p.41/44 Rediscoverers of Mendel

Carl Correns Erich von Tschermak-Seysenegg Hugo De Vries

History of Genetics in Evolution – p.42/44 Founders of theoretical population genetics

R. A. Fisher J. B. S. Haldane Sewall Wright

History of Genetics in Evolution – p.43/44 Developers and popularizers of the Neodarwinian Synthesis

Ernst Mayr George Gaylord Simpson Sir Julian Huxley

G.LedyardStebbins TheodosiusDobzhansky

History of Genetics in Evolution – p.44/44