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VCUG3001/VCPG6001 Unravelling Complexity April 2013 Does evolution drive toward ever-increasing complexity? Lindell Bromham Centre for Macroevolution and Macroecology Evolution, Ecology & Genetics, Research School of Biology, Australian National University www.tempoandmode.com www.macroevoeco.com Great Chain of Being God Angels Heaven Humans Beasts Plants Flame Rocks You are here Sacculina Sacculina Sacculina barnacles crustaceans arthropods language sociality Major Transitions in Evolution multicellularity cells chromosomes genes replication Does this trend indicate a drive toward complexity? 100 300 400 500 600 700 0 50 100 150 Cell type number 200 250 Millions of Years before Present 200 100 300 400 500 600 700 0 50 100 150 Cell type number 200 250 Millions of Years before Present 200 Genome size Diversity and complexity increase over time because uniformity and simplicity must be actively maintained in the face of inevitable change. Predicts non-adaptive trend toward complexity Increase more likely than decrease More DNA is costly More DNA is risky So if more DNA is bad, why does it accumulate? R. A. Fisher In a large population, selection can promote tiny advantages and build adaptation in small steps Tomoko Ohta The power of selection depends on population size In small populations, chance effects can lead to nonadaptive change ‘‘Where, then, is the direct supporting evidence for the assumption that complexity is entirely rooted in adaptive processes? No existing observations support such a claim, and, given the massive global dominance of unicellular species over multicellular eukaryotes, in terms of both species richness and numbers of individuals, if there is an advantage of organismal complexity, one can only marvel at the inability of natural selection to promote it.’’ Michael Lynch Origins of Genome Architecture Does gain in complexity = failure to stay simple? Complexity is not a measure of success and does not necessarily convey advantage Stepwise evolution of complexity is not evidence for a driven trend Neutral models explain complex patterns using simple processes Diversity Availability Effort Sheehan PM (1977) Species diversity in the phanerozoic: a reflection of labor by systematicists? Paleobiology 3:325-328 Arthropoda Mollusca Chordata Platyhelminthes Nematoda Annelida Cnidaria Echinodermata Porifera Bryozoa Rotifera Myxozoa Nemertea Acanthocephala Tardigrada Gastrotricha Acoelomorpha Nematomorpha Brachiopoda Onychophora Kinorhyncha Entoprocta Sipuncula Echiura Hemichordata Gnathostomulida Ctenophora Chaetognatha Loricifera Mesozoa Phoronida Priapulida Cycliophora Xenoturbellida Placozoa Micrognathozoa 1 Ecdysozoa Lophotrochozoa Deuterostomia 10 100 1000 10000 100000 1000000 Approximate number of described species