9/2/14 Biology of Organisms Comparative Biology • Uses phylogenetic relationships to study the features that unite and distinguish different groups. • Allows us to understand everything in biology from genes to behavior to anatomy to ecology to distributions in a holistic manner. • Why is an introduction to life’s diversity important? Comparative Biology Escherichia coli Saccharomyces cerevisiae Caenorhabditis elegans Drosophila melanogaster Xenopus laevis Mus musculus Arabidopsis thaliana Zea mays Danio rerio • This is why we can study human diseases using mouse models. • This is why principles of genetics derived from fruit flies are generalizable. • This all goes back to the unity of life. Macaca mulatta 1 9/2/14 Comparative Biology • Some of the most exciting recent discoveries in biology include the elucidation of common genetic elements of animal development. • HOX genes. Biology of Organisms • Interactions between life’s diversity underlie essentially all of biology. • How many interactions are going on here? Biology of Organisms • Interactions between life’s diversity underlie essentially all of biology. • How might this be relevant to your future interests? • Medicine? ic zo oMes zoic Cenozoic Humans leo Pa Colonization of land Today, Module 1: Animals Origins, History, & Unity of Life Origin of solar system and Earth 4 1 Proterozoic Bil lio 2 ns of Archaean a ye rs ag 3 Prokaryotes o Multicellular eukaryotes Single-celled eukaryotes Atmospheric oxygen 2 9/2/14 What is life? • What would we look for in a life form? • What defines life? • It’s been around for 3.8-3.5 billion years What is life? • Reproduction – Oldest rocks are 4.0-3.8 billion years • Any thoughts? Asexual What is life? • Reproduction • Metabolism: – the set of chemical reactions that occur in living organisms that manage the material and energy resources of the cell. Sexual What is life? • Reproduction • Metabolism • Organization – Non-random – Hierarchies – Emergent properties 3 9/2/14 What is life? • • • • What is life? Reproduction Metabolism Organization Growth & Development • • • • Reproduction Metabolism Organization Growth & Development • Homeostasis – Heritability – Cells – Regulating internal environment What is life? • • • • • • Reproduction Metabolism Organization Growth & Development Homeostasis Responds to the environment – Temperature, moisture, sunlight, substrate What is life? • • • • • • Reproduction Metabolism Organization Growth & Development Homeostasis Responds to the environment • Evolution, Adaptation, & Extinction 4 9/2/14 III. A hierarchy of organization • From atoms to grasslands • There are increasing levels of complexity IV. Emergent Properties atom tissue organ – An upside-down pyramid of increasing structural and functional complexity. • At each increasing level, the whole is more than the sum of its parts molecule cell population organism • With increasing complexity the hierarchical level becomes more than the sum of its parts. • These are known as emergent properties. • These are novel properties that emerge from interactions at lower levels. • How is this cathedral termite mound an example of an emergent property? organ system ecosystem biome IV. Emergent Properties • As biologists, to understand the whole we need to break it down and examine its parts. • Reductionist perspective. • But we always must keep in mind that when we do this that the whole loses its emergent properties. organelle V: Correlation: structure, function, diversity • Divergence through evolution. – a.k.a. descent with modification. • Organisms have both a shared ancestry and new attributes. Mammalian Forelimb 5 9/2/14 V: Correlation: structure, function, diversity • The pentamerous (five-digit) arrangement of the mammalian forelimb indicates homology. V: Correlation: structure, function, diversity • This pentamerous arrangement has been subsequently been modified through adaptation. – Features that are similar as a result of descent. Mammalian Forelimb VI. Unity in diversity • Sometimes it is difficult to see unity in diversity. • What, for example, could a hummingbird and a mushroom have in common? VI. Unity in diversity • Fungi and Animals diverged some 965 million years ago! • Evolution will, of course, obscure these relationships. • But they are all part of the hierarchy of life. 6 9/2/14 VI. Unity in diversity • Over 1.5 million species are named. • But more than 98% of all species that have ever existed have become extinct. • This also obscures relationships. VII: Pattern & Process • Pattern: Description of the WHAT? • Process: Description of the HOW? • This course will mainly be about the description. • Because we must know what exists before we attempt to explain VI. Unity in diversity: Phylogenetics and Taxonomy • The study of diversity is known as SYSTEMATICS. • Phylogenetics is the practice of elucidating relationships. • Taxonomy is the practice of naming organisms. • Classification arranges organisms. I. Fossils & Sedimentation • Fossils are the most readily observable record of the history of life. • Key to the field of macroevolution. • Paleontology is the study of fossils. 7 9/2/14 I. Fossils & Sedimentation • Unfortunately, the fossil record is both biased and incomplete. • Why would it be biased? I. Fossils & Sedimentation • Unfortunately, the fossil record is both biased and incomplete. • Why would it be biased? • Why would it be incomplete? I. Fossils & Sedimentation • Taphonomic conditions must be appropriate. I. Fossils & Sedimentation • Taphonomic conditions depend upon: • Geological processes • Type of fossil • Age of fossils – These are the conditions that permit decaying organisms to become fossilized. Will this wombat skeleton fossilize? Shales are particularly good for preserving fossils 8 9/2/14 I. Fossils & Sedimentation • Geological Processes • Most fossils are found in sedimentary rock. • How are sediments formed? What are the implications of this for the abundance of fossils? • Also mineralized amber and ice. I. Fossils & Sedimentation I. Fossils & Sedimentation • Types of Fossils • The vast majority are of hard parts. Why? I. Fossils & Sedimentation • Types of Fossils • Types of Fossils • The vast majority are of hard parts. Why? • Trace fossils provide information on interactions, ecology, behavior, functional morphology. • How? • These are rare! – Soft tisues are broken down by bacteria and fungi – Hard parts become mineralized. • This includes animals AND plants. Dinosaur tracks Leaf-mining insects 9 9/2/14 I. Fossils & Sedimentation I. Fossils & Sedimentation • Ages of Fossils • Ages of Fossils • Older fossils are much more rare. • Why? • Older fossils are much more rare. • More opportunity for geological processes to obscure or destroy: weathering, subduction, etc. • Older rocks are rarer. Stromatolites Fossilized stromatolite I. Fossils & Sedimentation • The rarity of appropriate taphonomic conditions results in this bias and incompleteness. • Despite this, the fossil record provides remarkable insights into the history of life on earth. “eBay insect fossil is new species” Stromatolites Fossilized stromatolite II. Dating of major events • How do paleontologists estimate fossil/strata ages? – Relative – Absolute One second before the end of the dinosaurs… 10 9/2/14 II. Dating of major events • Relative dating: • Usually older fossils at bottom of strata, younger towards top. II. Dating of major events • Absolute dating. • Radioactive elements: isotopes that decay at a constant rate. • The ratio of these versus the stable isotopes that they decay into gives us a metric of the age that the sediment was formed – or the fossil itself if any organic Carbon is lucky enough to be preserved. II. Dating of major events Common isotope ratios used in radiometric dating II. Dating of major events • Generalizations: • Index fossils help correlate ages of strata over wide areas. • Based on welldocumented fossils of short-lived (but abundant) species. Viviparus glacialis is an index fossil for 2.3-1.8 mya 11 9/2/14 III. The Geological Time Scale IV. Major Episodes The geologic record is divided into the Archaean, the Proterozoic, and the Phanerozoic eons. • A combination of: – Relative dating – Absolute dating – Major events in the history of life • Give us the Geological Time Scale – You should become familiar with the names, dates, and major events in this time scale. The Archaean: 4.6-2.5 bya • Probably absent of life until 3.5 bya (first rocks 3.8 bya) • Prokaryotes appear (3.5 bya) • Massive increase in Oxygen (of biological origin) and first significant extinction at end of Archaean 2.5 bya I highly recommend that you study Table 25.1 from your book! Stromatolites Fossilized stromatolite The Archaean & Proterozoic together are commonly known as the Precambrian Era The Proterozoic: 2500-542 mya • First Eukaryotes and multicellular organisms appear. • Familiarize yourself with pages 516-517 and figure 25.9 in the textbook for this. 12 9/2/14 The Proterozoic: 2500-542 mya • First Eukaryotes and multicellular organisms appear. • Low diversity early (Snowball Earth) • Later characterized by the “Ediacaran” or “Vendian” biota. • Mass extinction of these forms at the end of this boundary. • Why? The Paleozoic Era: 542-251 mya • Began with the Cambrian Explosion. • Sudden appearance of modern animal phyla in the fossil record. • Localized fossils and DNA evidence suggest earlier origins (Conway Morris’ long fuse). • BUT the explosion refers to their widespread emergence and dominance. Phanerozoic: 542 mya-present • The Phanerozoic encompasses multicellular eukaryotic life • The Phanerozoic is divided into three eras: the Paleozoic, Mesozoic, and Cenozoic • Major boundaries between geological divisions correspond to extinction events in the fossil record The Paleozoic Era: 542-251 mya • Major features: – Colonization of land – Appearance of vascular plants – Origins of seed plants – Diversification of insect orders – Radiation of vertebrates 13 9/2/14 The Paleozoic Era: 542-251 mya • Ended with the PERMIAN extinction. • Correlated with formation of PANGEA. The Paleozoic Era: 542-251 mya • Ended with the PERMIAN extinction. • Correlated with formation of PANGEA. • Correlated with high levels of volcanism. • Loss of approximately 90% of all animal species. • Transition into the Mesozoic Era Continental Drift • The movement of earth’s continents relative to each other. • Based on the theory of plate tectonics. • Tectonic plates move in relation to each other causing continental drift, earthquakes, volcanoes, mountain-building, and oceanic trench formation The Mesozoic Era: 251-65mya • Divided into three Periods: – Triassic (251-200 mya) – Jurassic (200-145 mya) – Cretaceous (145-65 mya) • End of each characterized by extinctions 14 9/2/14 The Mesozoic Era: 251-65mya • Characterized by rise and dominance of Gymnosperms (Triassic and Jurassic). • Rise, diversification, and extinction of most dinosaur groups. • Origins and early diversification of mammals and Angiosperms. The Mesozoic Era: 251-65mya • Ended with Cretaceous Extinction The Mesozoic Era: 251-65mya • Ended with Cretaceous Extinction. • Significant evidence for massive impact event at Chicxulub crater in the Yucatán Peninsula. A split second before the extinction of the dinosaurs The Mesozoic Era: 251-65mya • Ended with Cretaceous Extinction. • Significant evidence for massive impact event at Chicxulub crater in the Yucatán Peninsula. • Worldwide Iridium layer at Cretaceous-Tertiary boundary (a.k.a. K-T boundary). • Extinction of 50% of all marine and terrestrial species, including all but one lineage of dinosaurs. Cenotes in the Yucatán indicate the rim of the ancient crater 15 9/2/14 The Cenozoic Era: 65mya-present • Divided into two periods: What is the objective of systematics? – Paleogene (65-23 mya) – Neogene (23myapresent) • Rise of angiosperms, mammals, and extreme diversification of insects. A. Objectives of Taxonomy A. Objectives of Taxonomy Small canine teeth; language 1. Sort and identify organisms into species. • This includes the unique derived characteristics that distinguish them. Cranial features; brain size; brain morphology 1. Sort and identify organisms into species. • This includes the unique derived characteristics that distinguish them. Presence of a chin Advanced toolmaking Dimensions of pelvis; upright gait; s-shaped spine 16 9/2/14 A. Objectives of Taxonomy Skull balanced upright on vertebral column 1. Sort and identify organisms into species. • • This includes the unique derived characteristics that distinguish them. This includes giving a name to undescribed species. Reduced hair cover Elongated thumb and shortened finger; Limb length Species: Panthera pardus A. Objectives of Taxonomy Genus: Panthera 1. Sort and identify organisms into species. 2. Arrange (classify) species into broader hierarchical taxonomic categories • • Family: Felidae Order: Carnivora Class: Mammalia From genera to domains What is our classification? Phylum: Chordata Kingdom: Animalia Bacteria 1. • • Provides the most information. Becomes predictive. Archaea B. How to classify--the rationale B. How to classify--the rationale 1. Classification should as much as possible reflect evolutionary history. Domain: Eukarya 2. Classification should as much as possible reflect evolutionary history. Single taxon should be composed of all species derived from a common ancestor. 1. 2. 3. Older: Family Pongidae Subfamily Ponginae Family Hominidae This is known as monophyletic Contrast older notion of primate families Pongidae + Hominidae vs current notion of Hominidae Illustrates concepts of paraphyly and polyphyly Tribe Pongini Current: Hominidae 17 9/2/14 B. How to classify--the rationale B. How to classify--the rationale • Paraphyletic: A taxon that does not include all of the descendants of the most recent common ancestor. C. The Importance of Homology • Structures that share a common ancestor. – forelimb of tetrapods. – opposable thumb of primates – flower of angiosperms • These provide information into relationships via monophyly. • Polyphyletic: A taxon that includes members derived from two or more ancestors. Note that this is part of a continuum with paraphyletic. C. The Importance of Homology • Similar structures that do not share a common ancestry are analogous or are homoplasies. • These result from convergent evolution. • Are not informative to relationships. Informative to adaptation. 18 9/2/14 PHYLOGENETICS • The study of evolutionary relatedness between organisms. • (Contrast with taxonomy) • Without phylogenetics, comparative biology would not exist and diversity would make no sense. PHYLOGENY • Graphical representation of the evolutionary history of a group as expressed in terms of relatedness. • Classifications based on phylogenies are known as natural classifications. • Phylogenies are based on shared, derived (or unique) homologous features. These are known as synapomorphies. Phylogenies Phylogenies Change through time Change through time Change through time Change through time Change through time Relative position is the only thing that matters! 19 9/2/14 Sister groups Nodes, Tips, Internodes The two taxa on either side of a split Lancelet (outgroup) Lamprey Fish Vertebral column Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear Outgroups Polytomies When resolu<on of the branching diagram is difficult Not part of the group in ques<on, but is closely related to the group Lancelet (outgroup) Lamprey Fish Vertebral column Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear 20 9/2/14 Terminology • Phylogenies are based on shared, derived (or unique) homologous features. These are known as apomorphies. • Synapomorphies are traits that are unique, derived, and indicate rela<onships. They denote clades. • Autapomorphies are traits that are unique and derived, but do not indicate rela<onships. They denote <ps. • Plesiomorphies are traits shared by a number of groups, and are inherited from ancestors older than the last common ancestor. They do not denote clades. Synapomorphies, Autapomorphies, & Plesiomorphies Lancelet (outgroup) Lamprey Fish Vertebral column Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear Lancelet (outgroup) Lancelet (outgroup) Lamprey Lamprey Fish Vertebral column Fish Vertebral column Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear Phylogenetic tree: Endothermic vertebrates with hair, mammary glands, three bones in the middle ear (etc.) are all related and thus called MAMMALS. Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear Is the vertebral column a defining feature of mammals? How or how not? 21 9/2/14 Lancelet (outgroup) Lancelet (outgroup) Lamprey Lamprey Fish Vertebral column Fish Vertebral column Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear NO! This is a shared ancestral trait or plesiomorphy. At what level does this become an synapomorphy? Understanding the branching diagram • Recall that species arise from a splitting of an ancestral (parent) population (speciation); • Thus, all life could be represented by millions of such branches going back to the first organisms. Hinged jaws Four walking legs Amphibians Reptiles Amniotic egg Mammals Hair, mammary glands, 3 bones in the middle ear Tree-thinking questions: Which is most closely related to a fish: amphibians, reptiles, or mammals? Which is most evolved: amphibians, reptiles, or mammals? Understanding the branching diagram • Recall that species arise from a splitting of an ancestral (parent) population (speciation); • Thus, all life could be represented by millions of such branches going back to the first organisms. 22 9/2/14 Understanding the branching diagram • Recall that species arise from a splitting of an ancestral (parent) population (speciation); • Thus, all life could be represented by millions of such branches going back to the first organisms. Understanding the branching diagram All the way to the relationships linking all species of cats (Johnson et al 2006) D. CHARACTERS • • A set of alternative conditions (character state) that are considered able to evolve one to another. Must search for and evaluate homologous structures. – Must follow Recognition Criteria of Homology: 1. Similarity in position 2. Detailed resemblance 3. Continuance through intermediate forms D. CHARACTERS • • A set of alternative conditions (character state) that are considered able to evolve one to another. Must search for and evaluate homologous structures. – Must follow Recognition Criteria of Homology: 1. Similarity in position ✔ 2. Detailed resemblance 3. Continuance through intermediate forms 23 9/2/14 D. CHARACTERS D. CHARACTERS • • • A set of alternative conditions (character state) that are considered able to evolve one to another. Must search for and evaluate homologous structures. • – Must follow Recognition Criteria of Homology: – Must follow Recognition Criteria of Homology: 1. Similarity in position ✔ 2. Detailed resemblance ✖ 3. Continuance through ✖ intermediate forms 1. Similarity in position ✔ 2. Detailed resemblance ✖ 3. Continuance through intermediate forms D. CHARACTERS • Types of Characters: – Must be products of evolutionary process – Must be heritable – What kinds of things fall under this? A set of alternative conditions (character state) that are considered able to evolve one to another. Must search for and evaluate homologous structures. • Morphological Characters • Physiological characters • Molecular characters • Behavioral characters • Ecological characters • Geographic characters D. CHARACTERS A simple rule for hypothesis testing • The more data, the better! – This applies to testing phylogenetic relationships as well: the more characters, the better – Also, the more character systems, the better. 24 9/2/14 Phylogenetic Analysis • How do we reconstruct phylogenetic trees? • Based on using characters to test hypotheses of phylogenetic relationships. • Remember that the branching diagram is the hypothesis. Phylogenetic Analysis 1. A set of data (character X taxon matrix) 2. A set of possible evolutionary trees 3. A means of evaluating the alternative trees given the data. • Identify homologous characters and delineate alternative character states. Phylogenetic Analysis 1. A set of data (character X taxon matrix) 2. A set of possible evolutionary trees 3. A means of evaluating the alternative trees given the data. Phylogenetic Analysis 1. A set of data (character X taxon matrix) 2. A set of possible evolutionary trees 3. A means of evaluating the alternative trees given the data. • These are the alternative hypotheses. 25 9/2/14 Phylogenetic Analysis 1. A set of data (character X taxon matrix) 2. A set of possible evolutionary trees 3. A means of evaluating the alternative trees given the data. • Based on distribution of using shared derived characters (apomorphies) to identify clades. • Evaluated based on maximum parsimony or maximum likelihood as the optimality criterion. Which tree is preferred? • Parsimony • Maximum-likelihood • We will spend time in lab on these AND your book is quite thorough with this (pp 542-547) • We should first investigate the simplest explanation for observed character state distributions. • Minimizes the number of evolutionary events on a tree. • Maximizes apomorphic characters while mimizing homoplasious characters. Which tree is preferred? • Parsimony • Maximum-likelihood • We will spend time in lab on these AND your book is quite thorough with this (pp 542-547) How would we draw a phylogeny of these lizards with only the information we have right here? C A B 26 9/2/14 Which tree is preferred? • Parsimony Which tree is preferred? A – Data matrix & alternative hypotheses. • Parsimony A – Minimize character changes on the trees. B B C C B Tail black Legs blue Back orange Front red A C A C B B Tail black Legs blue Back orange Front red C B A C A C B B Which tree is preferred? • Parsimony Which tree is preferred? • Parsimony A – Minimize character changes on the trees. – Minimize character changes on the trees. – Do so for every character. B C B Tail black A B A C A Legs blue Back orange Front red C Tail black A B A C B C B A C A Legs blue Back orange Front red A C C A C B B 27 9/2/14 Which tree is preferred? • Parsimony – Minimize character changes on the trees. – Do so for every character. – Count the number of changes. – Which is most parsimonious? A B C B A Which tree is preferred? • Parsimony • Maximum-likelihood • We will spend time in lab on these AND your book is quite thorough with this (pp 542-547) C C A • Similar, but now optimality no longer based on principle of parsimony. • Optimality based on specified model of evolution. • Generally applied to molecular data. • Uses external information. B Testing evolutionary hypotheses Using Phylogenies Mapping evolutionary transitions Some horned lizards squirt blood from their eyes when attacked by canids. How many times has blood-squirting evolved? • Now that you have a tree, what can you do with it? – Testing hypotheses about evolution – Learning about the characteristics of extinct species and ancestral lineages – Classifying organisms (later) Blood squirting? No Yes 28 9/2/14 Testing evolutionary hypotheses Testing evolutionary hypotheses Mapping evolutionary transitions Mapping evolutionary transitions Some horned lizards squirt blood from their eyes when attacked by canids. How many times has blood-squirting evolved? This phylogeny suggests a single evolutioary gain and a single loss of blood squirting Blood squirting? No Yes But a new phylogeny using multiple characters suggests that blood squirting has been lost many times in the evolution of this group Our interpretation of these evolutionary scenarios depends on phylogeny Leaché and McGuire. Molecular Phylogenetics and Evolution 39: 628-644 Testing evolutionary hypotheses Reconstructing ancestral characters This phylogeny also shows how we can use data from living species to infer character states in ancestral taxa ? ? Testing evolutionary hypotheses Mapping evolutionary transitions How many times has venom evolved in squamate reptiles? Once in the large “venom clade” Groups within this clade then evolved different venom types e.g., different proteins found in Snakes versus Gila monsters Ancestral state could be blue, purple, or intermediate…outgroup comparison indicates blue is most parsimonious Leaché and McGuire. Molecular Phylogenetics and Evolution 39: 628-644 Even non-venomous lizards in this clade (Iguania) share ancestral toxins Fry et al. (2006) Nature 439: 584-588 29 9/2/14 Testing evolutionary hypotheses Testing evolutionary hypotheses Convergence and modes of speciation Coevolution What can this phylogeny tell us about homology/analogy and speciation? Lake Tanganyika Aphids and bacteria are symbiotic Lake Malawi Given this close relationship, we might expect that speciation in an aphid would cause parallel speciation in the bacteria 1. Similarities between each pair are the result of convergence 2. Sympatric speciation more likely than allopatric speciation When comparing phylogenies for each group we see evidence for reciprocal cladogenesis (but also contradictions) Clark et al. (2000) Testing evolutionary hypotheses Testing evolutionary hypotheses Geographic origins Geographic origins Where did domestic corn (Zea mays maize) originate? Where did humans originate? A B Each tip is one of 135 different mitochondrial DNA types found among 189 individual humans Populations from Highland Mexico are at the base of each maize clade African mtDNA types are clearly basal on the tree, with the non-African types derived Vigilant et al. (1991) Science Matsuoka et al. (2002) Suggests that humans originated in Africa 30 9/2/14 The power of Phylogenetic Biology Phylogenetics and Dinosaur behavior • From understanding physiological processes to identifying endangered species. • To understanding dinosaur behavior… (?!) Belinda Chang and colleagues began by reconstructing a phylogeny of vertebrates using numerous genes. Phylogenetics and Dinosaur behavior Phylogenetics and Dinosaur behavior They then sequenced a rhodopsin gene from all of these species and tested their photoactive properties. Using this information, they reconstructed what the ancestral protein looked like. 31 9/2/14 Phylogenetics and Dinosaur behavior Phylogenetics and Dinosaur behavior They then synthesized this protein, expressed it in a mammalian cell line tissue culture, and tested its photoactive properties. The photoactive properties were significantly red-shifted from modern birds, suggesting that early archosaurs hunted at dawn and dusk! 32
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