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LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 25 The History of Life on Earth Lectures by Erin Barley Kathleen Fitzpatrick © 2011 Pearson Education, Inc. • macroevolutionary – terrestrial vertebrates – mass extinctions – origin of flight in birds • Chemical and physical processes on early Earth may have produced very simple cells through a sequence of stages: 1. Abiotic synthesis of small organic molecules 2. Joining of these small molecules into macromolecules 3. Packaging of molecules into protocells 4. Origin of self-replicating molecules © 2011 Pearson Education, Inc. Figure 25.1 Synthesis of Organic Compounds on Early Earth • Earth formed about 4.6 billion years ago, along with the rest of the solar system • Bombardment of Earth by rocks and ice likely vaporized water and prevented seas from forming before 4.2 to 3.9 billion years ago • Earth’s early atmosphere likely contained water vapor and chemicals released by volcanic eruptions (nitrogen, nitrogen oxides, carbon dioxide, methane, ammonia, hydrogen, hydrogen sulfide) © 2011 Pearson Education, Inc. • Oparin and Haldane: 1920s: early atmosphere was a reducing environment • Miller and Urey: 1953, experiments for abiotic synthesis of organic molecules in a reducing atmosphere is possible – the first organic compounds may have been synthesized near volcanoes or deep-sea vents – Amino acids have also been found in meteorites © 2011 Pearson Education, Inc. Abiotic Synthesis of Macromolecules • RNA monomers: spontaneously form from simple molecules • Small organic molecules polymerize when they are concentrated on hot sand, clay, or rock Protocells • Replication and metabolism are key properties of life • may have been fluid-filled vesicles with a membrane-like structure • In water, lipids and other organic molecules can spontaneously form vesicles with a lipid bilayer © 2011 Pearson Education, Inc. Self-Replicating RNA and the Dawn of Natural Selection • first genetic material was RNA • RNA molecules called ribozymes have been found to catalyze many different reactions • Natural selection has produced self-replicating RNA molecules • RNA molecules that were more stable or replicated more quickly would have left the most descendent RNA molecules • RNA could have provided the template for DNA, a more stable genetic material © 2011 Pearson Education, Inc. The Fossil Record • Sedimentary rocks are deposited into layers called strata and are the richest source of fossils • Few individuals have fossilized, and even fewer have been discovered • The fossil record is biased in favor of species that – Existed for a long time – Were abundant and widespread – Had hard parts Video: Grand Canyon © 2011 Pearson Education, Inc. Figure 25.4 Present Dimetrodon Rhomaleosaurus victor 100 mya 1m 0.5 m 4.5 cm Coccosteus cuspidatus 175 200 Tiktaalik 270 300 Hallucigenia 375 400 1 cm Stromatolites 2.5 cm 500 525 Dickinsonia costata 565 600 Fossilized stromatolite 1,500 3,500 Tappania How Rocks and Fossils Are Dated • Sedimentary strata reveal the relative ages of fossils • The absolute ages of fossils can be determined by radiometric dating • A “parent” isotope decays to a “daughter” isotope at a constant rate • Each isotope has a known half-life, the time required for half the parent isotope to decay © 2011 Pearson Education, Inc. Fraction of parent isotope remaining Figure 25.5 1 Accumulating “daughter” isotope 2 Remaining “parent” isotope 1 1 4 1 2 3 Time (half-lives) 8 1 4 16 Concept 25.3: Key events in life’s history include the origins of single-celled and multicelled organisms and the colonization of land • The geologic record is divided into the Archaean, the Proterozoic, and the Phanerozoic eons • The Phanerozoic encompasses multicellular eukaryotic life • The Phanerozoic is divided into three eras: the Paleozoic, Mesozoic, and Cenozoic © 2011 Pearson Education, Inc. Figure 25.7-3 Cenozoic Humans Colonization of land Origin of solar system and Earth Animals Multicellular eukaryotes 4 1 Proterozoic 2 Archaean 3 Prokaryotes Single-celled eukaryotes Atmospheric oxygen The First Single-Celled Organisms • The oldest known fossils are stromatolites, rocks formed by the accumulation of sedimentary layers on bacterial mats • Stromatolites date back 3.5 billion years ago • Prokaryotes were Earth’s sole inhabitants from 3.5 to about 2.1 billion years ago © 2011 Pearson Education, Inc. Photosynthesis and the Oxygen Revolution • Most atmospheric oxygen (O2) is of biological origin • O2 produced by oxygenic photosynthesis reacted with dissolved iron and precipitated out to form banded iron formations © 2011 Pearson Education, Inc. • By about 2.7 billion years ago, O2 began accumulating in the atmosphere and rusting ironrich terrestrial rocks • This “oxygen revolution” from 2.7 to 2.3 billion years ago caused the extinction of many prokaryotic groups • Some groups survived and adapted using cellular respiration to harvest energy © 2011 Pearson Education, Inc. Atmospheric O2 (percent of present-day levels; log scale) Figure 25.8 1,000 100 10 1 0.1 “Oxygen revolution” 0.01 0.001 0.0001 4 3 2 Time (billions of years ago) 1 0 • The early rise in O2 was likely caused by ancient cyanobacteria • A later increase in the rise of O2 might have been caused by the evolution of eukaryotic cells containing chloroplasts © 2011 Pearson Education, Inc. The First Eukaryotes • The oldest fossils of eukaryotic cells date back 2.1 billion years • Eukaryotic cells have a nuclear envelope, mitochondria, endoplasmic reticulum, and a cytoskeleton • The endosymbiont theory proposes that mitochondria and plastids (chloroplasts and related organelles) were formerly small prokaryotes living within larger host cells • An endosymbiont is a cell that lives within a host cell © 2011 Pearson Education, Inc. Figure 25.9-3 Plasma membrane Cytoplasm DNA Ancestral prokaryote Nucleus Endoplasmic reticulum Photosynthetic prokaryote Mitochondrion Nuclear envelope Aerobic heterotrophic prokaryote Mitochondrion Plastid Ancestral heterotrophic eukaryote Ancestral photosynthetic eukaryote • Key evidence supporting an endosymbiotic origin of mitochondria and plastids: – Inner membranes are similar to plasma membranes of prokaryotes – Division is similar in these organelles and some prokaryotes – These organelles transcribe and translate their own DNA – Their ribosomes are more similar to prokaryotic than eukaryotic ribosomes © 2011 Pearson Education, Inc. The Origin of Multicellularity • The evolution of eukaryotic cells allowed for a greater range of unicellular forms • A second wave of diversification occurred when multicellularity evolved and gave rise to algae, plants, fungi, and animals © 2011 Pearson Education, Inc. The Colonization of Land • Fungi, plants, and animals began to colonize land about 500 million years ago • Vascular tissue in plants transports materials internally and appeared by about 420 million years ago • Plants and fungi today form mutually beneficial associations and likely colonized land together • Arthropods and tetrapods are the most widespread and diverse land animals • Tetrapods evolved from lobe-finned fishes around 365 million years ago © 2011 Pearson Education, Inc. Plate Tectonics • Land masses of Earth have formed a supercontinent: 1.1 billion, 600 million, and 250 million years ago • Tectonic plates move slowly through the process of continental drift – Oceanic and continental plates can collide, separate, or slide past each other – mountains and islands, and earthquakes © 2011 Pearson Education, Inc. Figure 25.13 North American Plate Juan de Fuca Plate Eurasian Plate Caribbean Plate Philippine Plate Arabian Plate Indian Plate Cocos Plate Pacific Plate Nazca Plate South American Plate Scotia Plate African Plate Antarctic Plate Australian Plate Consequences of Continental Drift • Formation of the supercontinent Pangaea about 250 million years ago had many effects – A deepening of ocean basins – A reduction in shallow water habitat – A colder and drier climate inland • most species that have ever lived are now extinct • Mass extinction is the result of disruptive global environmental changes • In each of the five mass extinction events, more than 50% of Earth’s species became extinct © 2011 Pearson Education, Inc. Figure 25.15 1,100 1,000 25 800 20 700 600 15 500 400 10 300 200 5 100 Era Period 0 E 542 O Paleozoic D S 488 444 416 359 Mesozoic P C 299 Tr 251 J 200 Cenozoic C 145 P 65.5 0 Q N 0 Number of families: Total extinction rate (families per million years): 900 • Permian extinction: 251 million years ago – extinction of about 96% of marine animal species • Intense volcanism in what is now Siberia • Global warming resulting from the emission of large amounts of CO2 from the volcanoes • Reduced temperature gradient from equator to poles • Oceanic anoxia from reduced mixing of ocean waters © 2011 Pearson Education, Inc. • Cretaceous mass extinction: 65.5 million years – half of all marine species and many terrestrial plants and animals, including most dinosaurs • iridium in sedimentary rocks suggests a meteorite • Dust clouds blocked sunlight and disturbed global climate • The Chicxulub crater off the coast of Mexico is evidence of a meteorite that dates to the same time © 2011 Pearson Education, Inc. Adaptive Radiations • Adaptive radiation is the evolution of diversely adapted species from a common ancestor • Adaptive radiations may follow – Mass extinctions – The evolution of novel characteristics – The colonization of new regions • Mammals underwent an adaptive radiation after the extinction of terrestrial dinosaurs © 2011 Pearson Education, Inc. Changes in Rate and Timing • Heterochrony evolutionary change in the rate or timing of developmental events – It can have a significant impact on body shape • Paedomorphosis, the rate of reproductive development accelerates compared with somatic development – The sexually mature species may retain body features that were previous juvenile structures Animation: Allometric Growth © 2011 Pearson Education, Inc. Figure 25.21 Chimpanzee infant Chimpanzee adult Chimpanzee fetus Chimpanzee adult Human fetus Human adult Figure 25.22 Gills Changes in Spatial Pattern • Homeotic genes: control the placement and organization of body parts • Hox genes: class of homeotic genes that provide positional information during development – If Hox genes are expressed in the wrong location, body parts can be produced in the wrong location • Specific changes in the Ubx gene have been identified that can “turn off” leg development © 2011 Pearson Education, Inc. Figure 25.24 Hox gene 6 Hox gene 7 Hox gene 8 Ubx About 400 mya Drosophila Artemia Evolutionary Novelties • Most novel biological structures evolve in many stages from previously existing structures • Complex eyes have evolved from simple photosensitive cells independently many times • Exaptations are structures that evolve in one context but become co-opted for a different function • Natural selection can only improve a structure in the context of its current utility © 2011 Pearson Education, Inc. Figure 25.26 (a) Patch of pigmented cells (b) Eyecup Pigmented cells (photoreceptors) Pigmented cells Epithelium Nerve fibers Nerve fibers (c) Pinhole camera-type eye (d) Eye with primitive lens Epithelium Cellular mass (lens) Fluid-filled cavity (e) Complex camera lens-type eye Cornea Cornea Lens Retina Optic nerve Pigmented layer (retina) Optic nerve Optic nerve LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 26 Phylogeny and the Tree of Life Lectures by Erin Barley Kathleen Fitzpatrick © 2011 Pearson Education, Inc. • Phylogeny: evolutionary history of a species or related species • Systematics: classifies organisms and determines their evolutionary relationships – fossil, molecular, and genetic data • Taxonomy is the ordered division and naming of organisms © 2011 Pearson Education, Inc. Binomial Nomenclature • Linnaeus (18th Century) taxonomy based on resemblances – two-part names for species and hierarchical classification • Genus: first part of the name – The first letter is capitalized • specific epithet : second part is unique for each species within the genus • Entire species name is italicized • Taxon: a taxonomic unit – domain, kingdom, phylum, class, order, family, genus, and species © 2011 Pearson Education, Inc. Figure 26.3 Species: Panthera pardus Genus: Panthera Family: Felidae Order: Carnivora Class: Mammalia Phylum: Chordata Domain: Bacteria Kingdom: Animalia Domain: Eukarya Domain: Archaea Linking Classification and Phylogeny • Phylogenetic Tree: evolutionary relationships in branching trees – hypothesis about evolutionary relationships – Each branch represents the divergence of two species • Systematists have proposed the PhyloCode, – Only recognizes groups with a common ancestor and its descendents • Sister taxa: groups with immediate common ancestor • rooted tree: a branch to represent the last common ancestor • basal taxon: diverges early in the group history and originates near the common ancestor • Polytomy: branch from which more than two groups emerge © 2011 Pearson Education, Inc. Figure 26.4 Order Family Genus Species Panthera Felidae Panthera pardus (leopard) Taxidea Lutra Mustelidae Carnivora Taxidea taxus (American badger) Lutra lutra (European otter) Canis Canidae Canis latrans (coyote) Canis lupus (gray wolf) Figure 26.5 Branch point: where lineages diverge Taxon A Taxon B Taxon C Sister taxa Taxon D ANCESTRAL LINEAGE Taxon E Taxon F Taxon G This branch point represents the common ancestor of taxa A–G. This branch point forms a polytomy: an unresolved pattern of divergence. Basal taxon What We Can and Cannot Learn from Phylogenetic Trees • Phylogenetic trees show patterns of descent, not phenotypic similarity • Phylogenetic trees do not indicate when species evolved or how much change occurred in a lineage • It should not be assumed that a taxon evolved from the taxon next to it © 2011 Pearson Education, Inc. Phylogenies are inferred from morphological and molecular data • Homologies: phenotypic and genetic similarities due to shared ancestry – Organisms with similar morphologies or DNA sequences are likely to be related • In phylogeny, must distinguish whether a similarity is the result of: – Homology is similarity due to shared ancestry – Analogy is similarity due to convergent evolution • similar environmental pressures/ natural selection produce similar adaptations in different evolutionary lineages © 2011 Pearson Education, Inc. Figure 26.7 • Bat and bird wings are homologous as forelimbs, but analogous as functional wings • Analogous structures or molecular sequences that evolved independently are also called homoplasies • Homology can be distinguished from analogy by comparing fossil evidence and the degree of complexity • The more complex two similar structures are, the more likely it is that they are homologous • Molecular systematics uses DNA and other molecular data to determine evolutionary relationships © 2011 Pearson Education, Inc. Concept 26.3: Shared characters are used to construct phylogenetic trees • Cladistics: group organisms by common descent – Clade: group of species that includes an ancestral species and all its descendants – A valid clade is monophyletic, signifying that it consists of the ancestor species and all its descendants – A paraphyletic grouping consists of an ancestral species and some, but not all, of the descendants – A polyphyletic grouping consists of various species with different ancestors © 2011 Pearson Education, Inc. Figure 26.10 (a) Monophyletic group (clade) (b) Paraphyletic group (c) Polyphyletic group A A B B C C C D D D E E F F F G G G A B Group Group E Group Shared Ancestral and Shared Derived Characters • A shared ancestral character: character that originated in a taxon ancestor • A shared derived character: evolutionary novelty unique to a particular clade • In some trees, the length of a branch can reflect the number of genetic changes that have taken place in a particular DNA sequence • In other trees, branch length can represent chronological time, and branching points can be determined from the fossil record © 2011 Pearson Education, Inc. Figure 26.11 Lancelet (outgroup) CHARACTERS Lancelet (outgroup) Lamprey Bass Frog Turtle Leopard TAXA Lamprey 0 1 1 1 1 1 Bass Vertebral column (backbone) Hinged jaws 0 0 1 1 1 1 Four walking legs 0 0 0 1 1 1 Amnion 0 0 0 0 1 1 Hair 0 0 0 0 0 1 Vertebral column Frog Hinged jaws Turtle Four walking legs Amnion Leopard Hair (a) Character table (b) Phylogenetic tree Figure 26.13 Drosophila Lancelet Zebrafish Frog Chicken Human Mouse PALEOZOIC 542 MESOZOIC 251 Millions of years ago CENOZOIC 65.5 Present • Maximum parsimony assumes that the tree that requires the fewest evolutionary events is the most likely • maximum likelihood: given certain rules about how DNA changes over time, a tree can be found that reflects the most likely sequence of evolutionary events • Computer programs are used to search for trees that are parsimonious and likely © 2011 Pearson Education, Inc. Figure 26.14 Human Mushroom Tulip 0 30% 40% 0 40% Human Mushroom Tulip 0 (a) Percentage differences between sequences 15% 5% 5% 15% 15% 10% 25% 20% Tree 1: More likely Tree 2: Less likely (b) Comparison of possible trees Phylogenetic Trees as Hypotheses • The best phylogenetic trees fits the most data: morphological, molecular, and fossil • Phylogenetic bracketing allows us to predict features of an ancestor from features of its descendants – Birds and crocodiles share several features: fourchambered hearts, song, nest building, and brooding – The fossil record supports nest building and brooding in dinosaurs © 2011 Pearson Education, Inc. Figure 26.16 Lizards and snakes Crocodilians Common ancestor of crocodilians, dinosaurs, and birds Ornithischian dinosaurs Saurischian dinosaurs Birds Concept 26.4: An organism’s evolutionary history is documented in its genome • DNA that codes for rRNA changes relatively slowly and is useful for investigating branching points hundreds of millions of years ago • mtDNA evolves rapidly and can be used to explore recent evolutionary events © 2011 Pearson Education, Inc. • Gene duplication increases the number of genes in the genome, providing more opportunities for evolutionary changes – Repeated gene duplications result in gene families – traced to a common ancestor • Orthologous genes are found in a single copy in the genome and are homologous between species – diverge only after speciation • Paralogous genes result from gene duplication, so are found in more than one copy in the genome – diverge within the clade that carries them and often evolve new functions © 2011 Pearson Education, Inc. Figure 26.18 Formation of orthologous genes: a product of speciation Species A Formation of paralogous genes: within a species Ancestral gene Ancestral gene Ancestral species Species C Speciation with divergence of gene Gene duplication and divergence Orthologous genes Paralogous genes Species C after many generations Species B Genome Evolution • Orthologous genes are widespread and extend across many widely varied species – For example, humans and mice diverged about 65 million years ago, and 99% of our genes are orthologous • Gene number and the complexity of an organism are not strongly linked – For example, humans have only four times as many genes as yeast, a single-celled eukaryote • Genes in complex organisms appear to be very versatile, and each gene can perform many functions © 2011 Pearson Education, Inc. Concept 26.5: Molecular clocks help track evolutionary time • A molecular clock uses constant rates of evolution in some genes to estimate the absolute time of evolutionary change – orthologous genes: nucleotide substitutions are proportional to the time of the last common ancestor – paralogous genes: nucleotide substitutions are proportional to the time gene duplication • Molecular clocks are calibrated against branches whose dates are known from the fossil record • Individual genes vary in how clocklike they are © 2011 Pearson Education, Inc. Neutral Theory • much evolutionary change in genes and proteins has no effect on fitness and is not influenced by natural selection • the rate of molecular change in these genes and proteins should be regular like a clock – The molecular clock does not run as smoothly as neutral theory predicts – Irregularities result from natural selection in which some DNA changes are favored over others – Estimates of evolutionary divergences older than the fossil record have a high degree of uncertainty – The use of multiple genes may improve estimates © 2011 Pearson Education, Inc. Applying a Molecular Clock: The Origin of HIV • Phylogenetic analysis shows that HIV is descended from viruses that infect chimpanzees and other primates • HIV spread to humans more than once • Comparison of HIV samples shows that the virus evolved in a very clocklike way • Application of a molecular clock to one strain of HIV suggests that that strain spread to humans during the 1930s © 2011 Pearson Education, Inc. Figure 26.20 Index of base changes between HIV gene sequences 0.20 0.15 HIV 0.10 Range Adjusted best-fit line (accounts for uncertain dates of HIV sequences) 0.05 0 1900 1920 1940 1960 Year 1980 2000 From Two Kingdoms to Three Domains • Early taxonomists: plants or animals • Five kingdoms: Monera (prokaryotes), Protista, Plantae, Fungi, and Animalia • Three-domains: Bacteria, Archaea, and Eukarya – tree of life: eukaryotes and archaea are more closely related to each other than to bacteria – tree of life is based largely on rRNA genes, as these have evolved slowly Animation: Classification Schemes © 2011 Pearson Education, Inc. Figure 26.21 Eukarya Land plants Green algae Cellular slime molds Dinoflagellates Forams Ciliates Red algae Diatoms Amoebas Euglena Trypanosomes Leishmania Animals Fungi Green nonsulfur bacteria Sulfolobus Thermophiles (Mitochondrion) Spirochetes Halophiles COMMON ANCESTOR OF ALL LIFE Methanobacterium Archaea Chlamydia Green sulfur bacteria Bacteria Cyanobacteria (Plastids, including chloroplasts) • Horizontal gene transfer is the movement of genes from one genome to another – exchange of transposable elements and plasmids, viral infection, and fusion of organisms • Some researchers suggest that eukaryotes arose as an fusion between a bacterium and archaean • If so, early evolutionary relationships might be better depicted by a ring of life instead of a tree of life © 2011 Pearson Education, Inc.