Evidence for Evolution Graduate Seminar Introduction and overview The three main claims of Darwinian evolution • Living species are related by common ancestry • Change through time occurs at the population not the organism level • The main cause of adaptive evolution is natural selection (and related mechanisms) The three main claims of Darwinian evolution • Living species are related by common ancestry Common ancestor The importance of common ancestry • If two different species descended from a single ancestor then change (=evolution) is implied All differences evolved along the lineages What did people believe before Darwin? • Separate ancestry (many versions) A special case: Jean-Baptiste Lamarck • French Naturalist (17441829) • The first scientific theory of evolution • Struggled to reconcile evolution and the Scala naturae (great scale of being) A special case: Lamarck’s theory • Life progresses upward due to an internal drive towards perfection A special case: Lamarck’s theory • Life progresses upward due to an internal drive towards perfection • Why are “primitive” organisms still around? A special case: Lamarck’s theory • Life progresses upward due to an internal drive towards perfection • Why are “primitive” organisms still around? – Spontaneous generation of new life constantly Lamarck’s view Because all species follow the same trajectory of origin: they will form a ladder of advancement Lamarck’s view is basically separate ancestry First articulation of common ancestry Lyell, C. Principles of Geology, Vol. II, Chap. 1 Species 1 Species 2 Species 3 Common Ancestor Common Ancestor Darwin envisaged evolution as a tree The affinities of all the beings of the same class have sometimes be represented by a great tree. I believe this simile largely speaks the truth…… …The green and budding twigs may represent existing species; and those produced during former years may represent the long succession of extinct species….. ….the great Tree of Life….covers the earth with ever-branching and beautiful ramifications Charles Darwin, On the Origin of Species; pages 131-132 The only figure in The Origin of Species Two claims • That major groups of organisms share descent from common ancestry – vs. separate ancestry • That all living organisms share descent from common ancestry – vs. several origins Evidence for common ancestry (against separate ancestry) • • • • • • • Fossil record Homology Vestigial Structures Classification Hierarchical distribution of traits Agreement between gene trees Evolution during domestication Evidence for common ancestry • Fossil record – Transitional fossils (sometimes in temporal sequence): consistent with the existence of real common ancestors Living sister group Transitional fossils Major clade Traits Deep Homology • Distantly related organisms share structural similarities • Function varies • Explicable by common ancestry grasping leaping flying swimming running human “fish” whale Amphibia Reptilia Pentadactyl limb bat Vestigial structures • Structures that are non-functional (but functional in related species). For example: – – – – Human appendix, tail bones, Gill slits in mammal embryos Hip bones of whales and snakes Eyes in cave fish Trees explain patterns in trait distribution Fur; milk Amnion Four legs; lungs Vertebral column Trees explain patterns in trait distribution Fur; milk Amnion Four legs; lungs Vertebral column Applies a forteriori to molecular data Biogeography: closely related species live near each other Molecular phylogeny of Hawaiian and other Campanulaceae (Givnish et al.) Hawaii Correlation among gene trees (Penny et al. 1982. Testing the theory of evolution by comparing phylogenetic trees constructed from five different protein sequences. Nature 297: 197-200.) • When we estimate the phylogeny from different genes, we get trees that are much more similar than could happen by chance during separate ancestry • Amenable to statistical analysis We see diverse forms that are descended from single ancestor Brassica oleracea What about the claim of a single ancestor of all living organisms? • Shared biochemistry (e.g., same 4 nitrogenous bases, same 20 L-amino acids, ATP) – There are many possible nitrogenous bases and amino acids; Many sugars could have form the NA backbone – No chemical reason for L- vs. D-amino acids • Shared structures (ribosomes, lipid bilayer membranes) • Shared metabolic pathways (e.g., glycolysis) • Share information processing (genetic code) – The code is a “frozen accident” The three main claims of Darwinian evolution • Living species are related by common ancestry • Change through time occurs at the population not the organism level – No organism level mechanism is currently plausible • The main cause of adaptive evolution is natural selection (and related mechanisms) The main cause of adaptive evolution is natural selection • Claim 1: Natural selection happens – vs. Natural selection does not/cannot happen • Claim 2: Natural selection is sufficient to explain even the most complex traits of living organisms – vs. natural selection is not sufficient The main cause of adaptive evolution is natural selection • Claim 1: Natural selection happens • Evidence: – Artificial selection and rapid natural selection – Theoretically must occur if only minimal assumptions are met • Genetic variation • Limited resources The main cause of adaptive evolution is natural selection • Claim 2: Natural selection is sufficient to explain even the most complex traits of living organisms • “Evidence” – On short time scales it is very effective – Time is long – No other natural mechanisms are known