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Chapter 25 The History of Life on Earth PowerPoint Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp

Overview: Lost Worlds Past organisms were very different from those now alive. The fossil record shows macroevolutionary changes over large time scales including The emergence of terrestrial vertebrates The origin of photosynthesis Long-term impacts of mass extinctions.

MacroEvolution: Large Scale Changes Over Time

Concept 25.1: Conditions on early Earth made the origin of life possible 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 protobionts. 4. Origin of self-replicating molecules.

Synthesis of Organic Compounds on Early Earth Earth formed about 4.6 billion years ago, along with the rest of the solar system. Earth s early atmosphere likely contained water vapor and chemicals released by volcanic eruptions (nitrogen, nitrogen oxides, carbon dioxide, methane, ammonia, hydrogen, hydrogen sulfide). A. I. Oparin and J. B. S. Haldane hypothesized that the early atmosphere was a reducing environment. Stanley Miller and Harold Urey conducted lab experiments that showed that the abiotic synthesis of organic molecules in a reducing atmosphere is possible.

However, the evidence is not yet convincing that the early atmosphere was in fact reducing. Instead of forming in the atmosphere, the first organic compounds may have been synthesized near submerged volcanoes and deep-sea vents. Amino acids have also been found in meteorites.

Deep Sea Vents

Abiotic Synthesis of Macromolecules Monomers --> Polymers Small organic molecules polymerize when they are concentrated on hot sand, clay, or rock. Replication and metabolism are key properties of life. Protobionts are aggregates of abiotically produced molecules surrounded by a membrane or membranelike structure. Protobionts exhibit simple reproduction and metabolism and maintain an internal chemical environment.

Experiments demonstrate that protobionts could have formed spontaneously from abiotically produced organic compounds. For example, small membrane-bounded droplets called liposomes can form when lipids or other organic molecules are added to water.

Protobionts May Have Formed Spontaneously 20 µm Glucose-phosphate Glucose-phosphate Phosphatase Phosphate Starch Maltose Amylase (a) Simple reproduction by liposomes Maltose (b) Simple metabolism

Self-Replicating RNA and the Dawn of Natural Selection The first genetic material was probably RNA, not DNA. RNA molecules called ribozymes have been found to catalyze many different reactions For example, ribozymes can make complementary copies of short stretches of their own sequence or other short pieces of RNA.

Sedimentary Rocks and Fossils 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.

Sedimentary Rock Strata -- Fossils Present Rhomaleosaurus victor, a plesiosaur Dimetrodon Casts of ammonites Coccosteus cuspidatus Hallucigenia Dickinsonia costata Stromatolites Tappania, a unicellular eukaryote Fossilized stromatolite

Radiometric Dating 1 / 2 Accumulating daughter isotope Remaining parent isotope 1 / 4 1 / 8 1 /16 1 2 3 4 Time (half-lives)

Radiocarbon dating can be used to date fossils up to 75,000 years old. For older fossils, some isotopes can be used to date sedimentary rock layers above and below the fossil. The magnetism of rocks can provide dating information. Reversals of the magnetic poles leave their record on rocks throughout the world.

The Origin of New Groups of Organisms Mammals belong to the group of animals called tetrapods. The evolution of unique mammalian features through gradual modifications can be traced from ancestral synapsids through the present.

Synapsid (300 mya) Evolution of Mammals Temporal fenestra Therapsid (280 mya) Key Articular Quadrate Dentary Squamosal Temporal fenestra Early cynodont (260 mya) EARLY TETRAPODS Dimetrodon Reptiles (including dinosaurs and birds) Temporal fenestra Later cynodont (220 mya) Therapsids Very late cynodonts Mammals Very late cynodont (195 mya)

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. Major boundaries between geological divisions correspond to extinction events in the fossil record.

Geologic Record

Geologic Time Table Humans Colonization of land Animals Origin of solar system and Earth 1 4 Proterozoic Archaean Prokaryotes Multicellular eukaryotes 2 3 Single-celled eukaryotes Atmospheric oxygen

The First Single-Celled Organisms = Prokaryotes The oldest known fossils are stromatolites, rock-like structures composed of many layers of bacteria and sediment. Stromatolites date back 3.5 billion years ago Prokaryotes were Earth s sole inhabitants from 3.5 to about 2.1 billion years ago.

Photosynthesis and the Oxygen Revolution Most atmospheric oxygen (O 2 ) is of biological origin. O 2 produced by oxygenic photosynthesis reacted with dissolved iron and precipitated out to form banded iron formations. The source of O 2 was likely bacteria similar to modern cyanobacteria.

By about 2.7 billion years ago, O 2 began accumulating in the atmosphere and rusting iron-rich terrestrial rocks. This oxygen revolution from 2.7 to 2.2 billion years ago Posed a challenge for life Provided opportunity to gain energy from light Allowed organisms to exploit new ecosystems.

About 2.7 billion years ago, O 2 began accumulating in the atmosphere and rusting iron-rich terrestrial rocks.

The First Eukaryotes The oldest fossils of eukaryotic cells date back 2.1 billion years. The hypothesis of endosymbiosis 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.

The prokaryotic ancestors of mitochondria and plastids probably gained entry to the host cell as undigested prey or internal parasites. In the process of becoming more interdependent, the host and endosymbionts would have become a single organism. Serial endosymbiosis supposes that mitochondria evolved before plastids through a sequence of endosymbiotic events.

Invagination of Plasma Membrane Plasma membrane Ancestral prokaryote Cytoplasm DNA Endoplasmic reticulum Nucleus Nuclear envelope

Serial Endosymbiosis Aerobic heterotrophic prokaryote Mitochondrion Ancestral heterotrophic eukaryote

Serial Endosymbiosis Photosynthetic prokaryote Mitochondrion Plastid Ancestral photosynthetic eukaryote

Endosymbiotic Sequence: Ancestral Prokaryote Invagination of Plasma Membrane Cytoplasm DNA Endoplasmic reticulum Serial Endosymbiosis: Aerobic heterotrophic prokaryote Nuclear envelope Mitochondrion Ancestral heterotrophic eukaryote Nucleus Photosynthetic prokaryote Mitochondrion Plastid Ancestral photosynthetic eukaryote

Key evidence supporting an endosymbiotic origin of mitochondria and plastids: Similarities in inner membrane structures and functions. These organelles transcribe and translate their own DNA. Their ribosomes are more similar to prokaryotic than eukaryotic ribosomes.

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. Comparisons of DNA sequences date the common ancestor of multicellular eukaryotes to 1.5 billion years ago. The oldest known fossils of multicellular eukaryotes are of small algae that lived about 1.2 billion years ago.

The snowball Earth hypothesis suggests that periods of extreme glaciation confined life to the equatorial region or deep-sea vents from 750 to 580 million years ago. The Cambrian explosion refers to the sudden appearance of fossils resembling modern phyla in the Cambrian period (535 to 525 million years ago). The Cambrian explosion provides the first evidence of predator-prey interactions. Fossils in China provide evidence of modern animal phyla tens of millions of years before the Cambrian explosion.

Millions of years ago Sponges Cnidarians Echinoderms Chordates Brachiopods Annelids Molluscs Arthropods Cambrian Explosion 500 Early Paleozoic era (Cambrian period) 542 Late Proterozoic eon

Proterozoic Fossils that may be animal embryos (SEM) (a) Two-cell stage 150 µm (b) Later stage 200 µm

The Colonization of Land Fungi, plants, and animals began to colonize land about 500 million years ago. Plants and fungi likely colonized land together by 420 million years ago. Arthropods and tetrapods are the most widespread and diverse land animals. Tetrapods evolved from lobe-finned fishes around 365 million years ago.

Concept 25.4: The rise and fall of dominant groups reflect continental drift, mass extinctions, and adaptive radiations At three points in time, the land masses of Earth have formed a supercontinent: 1.1 billion, 600 million, and 250 million years ago. Earth s continents move slowly over the underlying hot mantle through the process of continental drift. Oceanic and continental plates can collide, separate, or slide past each other. Interactions between plates cause the formation of mountains and islands, and earthquakes.

Earth - Plate Tectonics: Continental Drift Crust Mantle Inner core Outer core (a) Cutaway view of Earth Juan de Fuca Plate Cocos Plate Pacific Plate North American Plate Caribbean Plate Nazca Plate South American Plate Scotia Plate (b) Major continental plates Arabian Plate African Plate Eurasian Plate Antarctic Plate Indian Plate Philippine Plate Australian Plate

Consequences of Continental Drift Formation of the supercontinent Pangaea about 250 million years ago had many effects: A reduction in shallow water habitat A colder and drier climate inland Changes in climate as continents moved toward and away from the poles Changes in ocean circulation patterns leading to global cooling.

History of Continental Drift Millions of years ago Paleozoic Mesozoic Cenozoic Present Eurasia 65.5 South America Africa India Madagascar Antarctica 135 251

The break-up of Pangaea lead to allopatric speciation. The current distribution of fossils reflects the movement of continental drift. Similarity of fossils in parts of South America and Africa supports the idea that these continents were formerly attached. The fossil record shows that most species that have ever lived are now extinct. At times, the rate of extinction has increased dramatically and caused a mass extinction. In each of the five mass extinction events, more than 50% of Earth s species became extinct.

Five Big Mass Extinctions Total extinction rate (families per million years): Number of families: 20 15 800 700 600 500 10 400 300 5 200 100 0 Era Period Paleozoic Mesozoic E O S D C P Tr J Cenozoic C P N 0 542 488 444 416 359 299 251 200 145 65.5 0 Time (millions of years ago)

The Permian extinction defines the boundary between the Paleozoic and Mesozoic eras. This mass extinction caused the extinction of about 96% of marine animal species and might have been caused by volcanism, which lead to global warming, and a decrease in oceanic oxygen. The Cretaceous mass extinction 65.5 million years ago separates the Mesozoic from the Cenozoic. Organisms that went extinct include about half of all marine species and many terrestrial plants and animals, including most dinosaurs.

Massive Meterorite Impact Evidence The presence of iridium in sedimentary rocks suggests a meteorite impact about 65 million years ago. The Chicxulub crater off the coast of Mexico is evidence of a meteorite that dates to the same time.

Evidence of Meteroite Impact Yucatán Peninsula NORTH AMERICA Chicxulub crater

Is a Sixth Mass Extinction Under Way? Consequences Scientists estimate that the current rate of extinction is 100 to 1,000 times the typical background rate. Data suggest that a sixth human-caused mass extinction is likely to occur unless dramatic action is taken. Mass extinction can alter ecological communities and the niches available to organisms. It can take from 5 to 100 million years for diversity to recover following a mass extinction. Mass extinction can pave the way for adaptive radiations.

Adaptive Radiations - New Environmental Opportunities Adaptive radiation is the evolution of diversely adapted species from a common ancestor upon introduction to new environmental opportunities. Mammals underwent an adaptive radiation after the extinction of terrestrial dinosaurs. The disappearance of dinosaurs (except birds) allowed for the expansion of mammals in diversity and size. Other notable radiations include photosynthetic prokaryotes, large predators in the Cambrian, land plants, insects, and tetrapods.

World-Wide Adaptive Radiations Ancestral mammal ANCESTRAL CYNODONT Monotremes (5 species) Marsupials (324 species) Eutherians (placental mammals; 5,010 species) 250 200 150 100 50 Millions of years ago 0

Regional Adaptive Radiations Adaptive radiations can occur when organisms colonize new environments with little competition. The Hawaiian Islands are one of the world s great showcases of adaptive radiation.

Hawaiian Islands -- Regional Adaptive Radiations Close North American relative, the tarweed Carlquistia muirii Dubautia laxa KAUAI 5.1 million years MOLOKAI 1.3 MAUI million years OAHU 3.7 LANAI million years HAWAII 0.4 million years Argyroxiphium sandwicense Dubautia waialealae Dubautia scabra Dubautia linearis

Major changes in body form result from changes in the sequences and regulation of developmental genes Studying genetic mechanisms of change can provide insight into large-scale evolutionary change. Genes that program development control the rate, timing, and spatial pattern of changes in an organism s form as it develops into an adult. Heterochrony is an evolutionary change in the rate or timing of developmental events. It can have a significant impact on body shape. The contrasting shapes of human and chimpanzee skulls are the result of small changes in relative growth rates.

Allometric Growth Newborn 2 5 15 Adult Age (years) (a) Differential growth rates in a human Chimpanzee fetus Chimpanzee adult Human fetus Human adult (b) Comparison of chimpanzee and human skull growth

Heterochrony can alter the timing of reproductive development relative to the development of nonreproductive organs In paedomorphosis, the rate of reproductive development accelerates compared with somatic development. The sexually mature species may retain body features that were juvenile structures in an ancestral species.

Paedomorphosis - Juvenile Gills Retained by Adult Salamander Gills

Changes in Spatial Pattern - Hox genes Substantial evolutionary change can also result from alterations in genes that control the placement and organization of body parts. Homeotic genes determine such basic features as where wings and legs will develop on a bird or how a flower s parts are arranged. Hox genes are a 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. For example, in crustaceans, a swimming appendage can be produced instead of a feeding appendage.

Evolution of vertebrates from invertebrate animals was associated with alterations in Hox genes. Two duplications of Hox genes have occurred in the vertebrate lineage. These duplications may have been important in the evolution of new vertebrate characteristics. The tremendous increase in diversity during the Cambrian explosion is a puzzle. Changes in developmental genes can also result in new morphological forms.

Hox Genes Alterations Hypothetical vertebrate ancestor (invertebrate) with a single Hox cluster First Hox duplication Hypothetical early vertebrates (jawless) with two Hox clusters Second Hox duplication Vertebrates (with jaws) with four Hox clusters

Changes in developmental genes can result in new morphological forms Hox gene 6 Hox gene 7 Hox gene 8 Ubx About 400 mya Drosophila Artemia

Concept 25.6: Evolution is not goal oriented Evolution is like tinkering it is a process in which new forms arise by the slight modification of existing forms. 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.

Evolution: new forms arise by the slight modification of existing forms Pigmented cells (photoreceptors) (a) Patch of pigmented cells Fluid-filled cavity Epithelium Nerve fibers Epithelium (b) Eyecup Cellular mass (lens) Pigmented cells Nerve fibers Cornea Optic nerve Pigmented layer (retina) (c) Pinhole camera-type eye Optic nerve (d) Eye with primitive lens Cornea Lens Optic nerve Retina (e) Complex camera-type eye

Horse Evolution Recent (11,500 ya) Pleistocene (1.8 mya) Pliocene (5.3 mya) Equus Hippidion and other genera Nannippus Pliohippus Hipparion Neohipparion Sinohippus Megahippus Callippus Archaeohippus Miocene (23 mya) Anchitherium Hypohippus Merychippus Parahippus Oligocene (33.9 mya) Miohippus Mesohippus Paleotherium Epihippus Propalaeotherium Eocene (55.8 mya) Pachynolophus Orohippus Key Hyracotherium Grazers Browsers

The appearance of an evolutionary trend does not imply that there is some intrinsic drive toward a particular phenotype 1.2 bya: First multicellular eukaryotes 2.1 bya: First eukaryotes (single-celled) 3.5 billion years ago (bya): First prokaryotes (single-celled) 535 525 mya: Cambrian explosion (great increase in diversity of animal forms) 500 mya: Colonization of land by fungi, plants and animals Millions of years ago (mya)

You should now be able to: 1. Define radiometric dating, serial endosymbiosis, Pangaea, snowball Earth, exaptation, heterochrony, and paedomorphosis. 2. Describe the contributions made by Oparin, Haldane, Miller, and Urey toward understanding the origin of organic molecules. 3. Explain why RNA, not DNA, was likely the first genetic material.

4. Describe and suggest evidence for the major events in the history of life on Earth from Earth s origin to 2 billion years ago. 5. Briefly describe the Cambrian explosion. 6. Explain how continental drift led to Australia s unique flora and fauna. 7. Describe the mass extinctions that ended the Permian and Cretaceous periods. 8. Explain the function of Hox genes.