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Chapter 15 The Evolution of Microbial Life PowerPoint® Lectures for Campbell Essential Biology, Fifth Edition, and Campbell Essential Biology with Physiology, Fourth Edition – Eric J. Simon, Jean L. Dickey, and Jane B. Reece Lectures by Edward J. Zalisko © 2013 Pearson Education, Inc. Biology and Society: Has Life Been Created in the Lab? • How did life first arise on Earth? • To gain insight, scientists have – synthesized from scratch the entire genome of a small bacterium known as Mycoplasma mycoides and – transplanted the artificial genome into the cells of a closely related species called Mycoplasma capricolum. © 2013 Pearson Education, Inc. Figure 15.0 Biology and Society: Has Life Been Created in the Lab? • The newly installed genome – took over the recipient cells, – began cranking out M. mycoides proteins, and – reproduced to make more cells containing the synthetic M. mycoides genome. © 2013 Pearson Education, Inc. MAJOR EPISODES IN THE HISTORY OF LIFE • Earth was formed about 4.6 billion years ago. • Prokaryotes – evolved by about 3.5 billion years ago, – began oxygen production about 2.7 billion years ago, – lived alone for more than a billion years, and – continue in great abundance today. © 2013 Pearson Education, Inc. MAJOR EPISODES IN THE HISTORY OF LIFE • Single-celled eukaryotes first evolved about 2.1 billion years ago. • Multicellular eukaryotes first evolved at least 1.2 billion years ago. © 2013 Pearson Education, Inc. Figure 15.1a Precambrian Ancestor to all present-day life Origin of Earth 4,500 Earth’s crust solidifies 4,000 Oldest prokaryotic fossils 3,500 Millions of years ago 3,000 Atmospheric oxygen begins to appear 2,500 Figure 15.1b Precambrian Oldest eukaryotic fossils 2,000 Origin of multicellular organisms 1,500 Millions of years ago Oldest animal fossils 1,000 Figure 15.1c Paleozoic Meso- Cenozoic zoic Bacteria Archaea Fungi Animals Cambrian explosion Oldest animal fossils 1,000 Extinction of Plants colonize land dinosaurs First humans 500 Millions of years ago 0 Eukaryotes Protists Plants Prokaryotes Precambrian MAJOR EPISODES IN THE HISTORY OF LIFE • All the major phyla of animals evolved by the end of the Cambrian explosion, which – began about 540 million years ago and – lasted about 10 million years. • Plants and fungi – first colonized land about 500 million years ago and – were followed by amphibians that evolved from fish. © 2013 Pearson Education, Inc. MAJOR EPISODES IN THE HISTORY OF LIFE • What if we use a clock analogy to tick down all of the major events in the history of life on Earth? © 2013 Pearson Education, Inc. Figure 15.2 Humans Origin of solar system and Earth 0 1 4 2 3 THE ORIGIN OF LIFE • We may never know for sure how life on Earth began. © 2013 Pearson Education, Inc. Resolving the Biogenesis Paradox • All life today arises by the reproduction of preexisting life, or biogenesis. • If this is true, how could the first organisms arise? • From the time of the ancient Greeks until well into the 1800s, it was commonly believed that life regularly arises from nonliving matter, an idea called spontaneous generation. © 2013 Pearson Education, Inc. Resolving the Biogenesis Paradox • Today, most biologists think it is possible that life on early Earth evolved from simple cells produced by – chemical and – physical processes. © 2013 Pearson Education, Inc. Figure 15.3 A Four-Stage Hypothesis for the Origin of Life • According to one hypothesis, the first organisms were products of chemical evolution in four stages. © 2013 Pearson Education, Inc. Stage 1: Abiotic Synthesis of Organic Monomers • The first stage in the origin of life was the first to be extensively studied in the laboratory. © 2013 Pearson Education, Inc. The Process of Science: Can Biological Monomers Form Spontaneously? • Observation: Modern biological macromolecules are all composed of elements that were present in abundance on early Earth. • Question: Could biological molecules arise spontaneously under conditions like those on early Earth? © 2013 Pearson Education, Inc. The Process of Science: Can Biological Monomers Form Spontaneously? • Hypothesis: A closed system designed to simulate early Earth conditions could produce biologically important organic molecules from inorganic ingredients. • Prediction: Organic molecules would form and accumulate. © 2013 Pearson Education, Inc. The Process of Science: Can Biological Monomers Form Spontaneously? • Experiment: An apparatus was built to mimic the early Earth atmosphere and included – hydrogen gas (H2), methane (CH4), ammonia (NH3), and water vapor (H2O), – sparks that were discharged into the chamber to mimic the prevalent lightning of early Earth, and – a condenser that cooled the atmosphere, causing water and dissolved compounds to “rain” into the miniature “sea.” © 2013 Pearson Education, Inc. Figure 15.4 “Atmosphere” CH4 Water vapor NH3 H2 Electrode Condenser Cold water Cooled water containing organic molecules H2O “Sea” Sample for chemical analysis Miller and Urey’s experiment The Process of Science: Can Biological Monomers Form Spontaneously? • Results: After the apparatus had run for a week, an abundance of organic molecules essential for life had collected in the “sea,” including amino acids, the monomers of proteins. • These laboratory experiments – have been repeated and extended by other scientists and – support the idea that organic molecules could have arisen abiotically on early Earth. © 2013 Pearson Education, Inc. Stage 2: Abiotic Synthesis of Polymers • Researchers have brought about the polymerization of monomers to form polymers, such as proteins and nucleic acids, by dripping solutions of organic monomers onto – hot sand, – clay, or – rock. © 2013 Pearson Education, Inc. Stage 3: Formation of Pre-Cells • A key step in the origin of life was the isolation of a collection of abiotically created molecules within a membrane. • Laboratory experiments demonstrate that pre-cells could have formed spontaneously from abiotically produced organic compounds. © 2013 Pearson Education, Inc. Stage 3: Formation of Pre-Cells • Such pre-cells produced in the laboratory display some lifelike properties. They – have a selectively permeable surface, – can grow by absorbing molecules from their surroundings, and – swell or shrink when placed in solutions of different salt concentrations. © 2013 Pearson Education, Inc. Stage 4: Origin of Self-Replicating Molecules • Life is defined partly by the process of inheritance, which is based on self-replicating molecules. • One hypothesis is that the first genes were short strands of RNA that replicated themselves – without the assistance of proteins, – perhaps using RNAs that can act as enzymes, called ribozymes. © 2013 Pearson Education, Inc. Figure 15.5-4 Original “gene” RNA monomers Formation of short RNA polymers: simple “genes” Assembly of a complementary RNA chain The complementary chain serves as a template of the original “gene.” From Chemical Evolution to Darwinian Evolution • Over millions of years – natural selection favored the most efficient precells and – the first prokaryotic cells evolved. © 2013 Pearson Education, Inc. PROKARYOTES • Prokaryotes lived and evolved all alone on Earth for about 2 billion years. © 2013 Pearson Education, Inc. They’re Everywhere! • Prokaryotes – are found wherever there is life, – have a collective biomass that is at least ten times that of all eukaryotes, – thrive in habitats too cold, too hot, too salty, too acidic, or too alkaline for any eukaryote, – cause about half of all human diseases, and – are more commonly benign or beneficial. © 2013 Pearson Education, Inc. Figure 15.6 They’re Everywhere! • Compared to eukaryotes, prokaryotes are – much more abundant and – typically much smaller. © 2013 Pearson Education, Inc. Colorized SEM Figure 15.7 They’re Everywhere! • Prokaryotes living in soil and at the bottom of lakes, rivers, and oceans help to decompose dead organisms and other organic waste material, returning vital chemical elements to the environment. © 2013 Pearson Education, Inc. The Structure and Function of Prokaryotes • Prokaryotic cells – lack a membrane-enclosed nucleus, – lack other membrane-enclosed organelles, – typically have cell walls exterior to their plasma membranes, but – display an enormous range of diversity. © 2013 Pearson Education, Inc. Figure 4.4 Plasma membrane Cell wall Capsule Prokaryotic flagellum Ribosomes Nucleoid Colorized TEM Pili Figure 4.5 Ribosomes Cytoskeleton Not in most plant cells Centriole Lysosome Plasma membrane Nucleus Mitochondrion Rough endoplasmic reticulum (ER) Smooth endoplasmic reticulum (ER) Golgi apparatus Idealized animal cell Cytoskeleton Mitochondrion Central vacuole Cell wall Chloroplast Nucleus Not in animal cells Rough endoplasmic reticulum (ER) Ribosomes Plasma membrane Smooth endoplasmic reticulum (ER) Idealized plant cell Channels between cells Golgi apparatus Prokaryotic Forms • The three most common shapes of prokaryotes are 1. spherical (cocci), 2. rod-shaped (bacilli), and 3. spiral or curved. © 2013 Pearson Education, Inc. Figure 15.8 SHAPES OF PROKARYOTIC CELLS Rod-shaped (bacilli) Colorized TEM Spiral Colorized SEM Colorized SEM Spherical (cocci) Prokaryotic Forms • All prokaryotes are unicellular. • Some species – exist as groups of two or more cells, – exhibit a simple division of labor among specialized cell types, or – are very large, dwarfing most eukaryotic cells. © 2013 Pearson Education, Inc. Colorized SEM Figure 15.9 LM LM (a) Actinomycete (b) Cyanobacteria (c) Giant bacterium LM Figure 15.9c (c) Giant bacterium Prokaryotic Forms • About half of all prokaryotes are mobile, and many of these travel using one or more flagella. © 2013 Pearson Education, Inc. Prokaryotic Forms • In many natural environments, prokaryotes attach to surfaces in a highly organized colony called a biofilm, which – may consist of one or several species of prokaryotes, – may include protists and fungi, – can show a division of labor and defense against invaders, and © 2013 Pearson Education, Inc. Prokaryotic Forms – can form on almost any type of surface, including – rocks, – metal, – plastic, and – organic material including teeth. © 2013 Pearson Education, Inc. Colorized SEM Figure 15.10 Prokaryotic Reproduction • Most prokaryotes can reproduce – by dividing in half by binary fission and – at very high rates if conditions are favorable. • Some prokaryotes form endospores, which are – thick-coated, protective cells – produced when the prokaryote is exposed to unfavorable conditions. © 2013 Pearson Education, Inc. Figure 15.11 Colorized TEM Endospore Prokaryotic Nutrition • Biologists use the phrase “mode of nutrition” to describe how organisms obtain energy and carbon. – Energy – Phototrophs obtain energy from light. – Chemotrophs obtain energy from environmental chemicals. © 2013 Pearson Education, Inc. Prokaryotic Nutrition – Carbon – Autotrophs obtain carbon from carbon dioxide (CO2). – Heterotrophs obtain carbon from at least one organic nutrient—the sugar glucose, for instance. © 2013 Pearson Education, Inc. Prokaryotic Nutrition • We can group all organisms according to the four major modes of nutrition if we combine the – energy source (phototroph versus chemotroph) and – carbon source (autotroph versus heterotroph). © 2013 Pearson Education, Inc. Prokaryotic Nutrition • Dominant among multicellular organisms are – photoautotrophs and – chemoheterotrophs. • The other two modes are used only by certain prokaryotes. © 2013 Pearson Education, Inc. Figure 15.12 MODES OF NUTRITION Energy source Chemical Photoautotrophs Chemoautotrophs Elodea, an aquatic plant Photoheterotrophs Bacteria from a hot spring Chemoheterotrophs Colorized TEM Organic compounds Carbon source CO2 Colorized TEM Light Rhodopseudomonas Kingfisher with prey The Two Main Branches of Prokaryotic Evolution: Bacteria and Archaea • By comparing diverse prokaryotes at the molecular level, biologists have identified two major branches of prokaryotic evolution: 1. bacteria and 2. archaea (more closely related to eukaryotes). © 2013 Pearson Education, Inc. The Two Main Branches of Prokaryotic Evolution: Bacteria and Archaea • Thus, life is organized into three domains: 1. Bacteria, 2. Archaea, and 3. Eukarya. © 2013 Pearson Education, Inc. The Two Main Branches of Prokaryotic Evolution: Bacteria and Archaea • Some archaea are “extremophiles.” – Halophiles thrive in salty environments. – Thermophiles inhabit very hot water. – Methanogens – inhabit the bottoms of lakes and swamps and – aid digestion in cattle and deer. © 2013 Pearson Education, Inc. Figure 15.13 (a) Salt-loving archaea (b) Heat-loving archaea Bacteria and Disease Bacteria That Cause Disease • Bacteria and other organisms that cause disease are called pathogens. • Most pathogenic bacteria produce poisons. – Exotoxins are proteins bacterial cells secrete into their environment. – Endotoxins are – not cell secretions but instead – chemical components of the outer membrane of certain bacteria. © 2013 Pearson Education, Inc. Colorized SEM Figure 15.14 Haemophilus influenzae Cells of nasal lining Bacteria That Cause Disease • The best defenses against bacterial disease are – sanitation, – antibiotics, and – education. © 2013 Pearson Education, Inc. Bacteria That Cause Disease • Lyme disease is – caused by bacteria carried by ticks and – treated with antibiotics, if detected early. © 2013 Pearson Education, Inc. SEM Figure 15.15 Tick that carries the Lyme disease bacterium “Bull’s-eye” rash Spirochete that causes Lyme disease Figure 15.15a “Bull’s-eye” rash Figure 15.15b Tick that carries the Lyme disease bacterium Figure 15.15c Tick that carries the Lyme disease bacterium SEM Figure 15.15d Spirochete that causes Lyme disease Biological Weapons • In October 2001, endospores of the bacterium that causes anthrax were mailed to members of the news media and the U.S. Senate. • Five people died from this attack. © 2013 Pearson Education, Inc. Biological Weapons • Another bacterium considered to have dangerous potential as a weapon is Clostridium botulinum, producer of the exotoxin botulinum, which – blocks transmission of nerve signals that cause muscle contraction and – is the deadliest poison on Earth. © 2013 Pearson Education, Inc. Biological Weapons • The bacterium that causes plague – is also a potential biological weapon, – is carried by rodents, and transmitted by fleas, – produces egg-size swellings called buboes under the skin, and – can be treated with antibiotics if diagnosed early. © 2013 Pearson Education, Inc. Figure 15.16 The Ecological Impact of Prokaryotes • Pathogenic bacteria are in the minority among prokaryotes. • Far more common are species that are essential to our well-being, either directly or indirectly. © 2013 Pearson Education, Inc. Prokaryotes and Chemical Recycling • Prokaryotes play essential roles in – chemical cycles in the environment and – the breakdown of organic wastes and dead organisms. © 2013 Pearson Education, Inc. Prokaryotes and Bioremediation • Bioremediation is the use of organisms to remove pollutants from – water, – air, and – soil. • A familiar example is the use of prokaryotic decomposers in sewage treatment. © 2013 Pearson Education, Inc. Figure 15.17 Rotating arm spraying liquid wastes Rock bed coated with aerobic prokaryotes and fungi Liquid wastes Outflow Prokaryotes and Bioremediation • Certain bacteria – can decompose petroleum and – are useful in cleaning up oil spills. © 2013 Pearson Education, Inc. Figure 15.18 PROTISTS • Protists are – eukaryotes that are not fungi, animals, or plants, – mostly unicellular, and – ancestral to all other eukaryotes. © 2013 Pearson Education, Inc. The Origin of Eukaryotic Cells • Eukaryotic cells evolved by – the infolding of the plasma membrane of a prokaryotic cell to form the endomembrane system and – a process known as endosymbiosis. © 2013 Pearson Education, Inc. The Origin of Eukaryotic Cells • Symbiosis is a more general association between organisms of two or more species. • Endosymbiosis – refers to one species living inside another host species and – is the process by which eukaryotes gained mitochondria and chloroplasts. © 2013 Pearson Education, Inc. Figure 15.19 Plasma membrane DNA Cytoplasm Membrane infolding Endoplasmic reticulum Nucleus Ancestral prokaryote Nuclear envelope Cell with nucleus and endomembrane system (a) Origin of the endomembrane system Photosynthetic prokaryote Endosymbiosis Aerobic heterotrophic prokaryote Chloroplast Mitochondrion Photosynthetic eukaryotic cell (b) Origin of mitochondria and chloroplasts The Diversity of Protists • The group called protists – consists of multiple clades but – remains a convenient term to refer to eukaryotes that are not plants, animals, or fungi. © 2013 Pearson Education, Inc. The Diversity of Protists • Protists obtain their nutrition in a variety of ways. – Algae are autotrophs, producing their food by photosynthesis. © 2013 Pearson Education, Inc. The Diversity of Protists – Other protists are heterotrophs. – Some protists eat bacteria or other protists. – Other protists are fungus-like and obtain organic molecules by absorption. – Parasites derive their nutrition from a living host, which is harmed by the interaction. Parasitic trypanosomes infect blood and cause sleeping sickness. © 2013 Pearson Education, Inc. (a) An autotroph: Caulerpa, a multicellular alga (b) A heterotroph: parasitic trypanosome LM Colorized SEM Figure 15.20 (c) A mixotroph: Euglena The Diversity of Protists • Protist habitats are diverse and include – oceans, lakes, and ponds, – damp soil and leaf litter, and – the bodies of host organisms with which they share mutually beneficial relationships, such as – unicellular algae and reef-building coral animals, and – cellulose-digesting protists and termites. © 2013 Pearson Education, Inc. Figure 15.21 LM Protozoans • Protists that live primarily by ingesting food are called protozoans. © 2013 Pearson Education, Inc. Another flagellate: Trichomonas A foram Red blood cell An apicomplexan Colorized TEM Cilia Cell “mouth” TEM LM Apical complex An amoeba A ciliate LM A flagellate: Giardia Colorized SEM Colorized SEM Figure 15.22 Protozoans • Protozoans with flagella are called flagellates and – are typically free-living, but – some are nasty parasites. © 2013 Pearson Education, Inc. Colorized SEM Figure 15.22a A flagellate: Giardia Colorized SEM Figure 15.22b A flagellate: Trichomonas Protozoans • Amoebas are characterized by – great flexibility in their body shape and – the absence of permanent organelles for locomotion. • Most species move and feed by means of pseudopodia (singular, pseudopodium), temporary extensions of the cell. © 2013 Pearson Education, Inc. Colorized TEM Figure 15.22c An amoeba Protozoans • Other protozoans with pseudopodia include the forams, which have shells. © 2013 Pearson Education, Inc. LM Figure 15.22d A foram Protozoans • Apicomplexans are – named for a structure at their apex (tip) that is specialized for penetrating host cells and tissues, – all parasitic, and – able to cause serious human diseases, such as malaria caused by Plasmodium. © 2013 Pearson Education, Inc. Figure 15.22e Red blood cell An apicomplexan TEM Apical complex Protozoans • Another apicomplexan is Toxoplasma, – occurring in the digestive tracts of millions of people in the United States but – held in check by the immune system. • A woman newly infected with Toxoplasma during pregnancy can pass the parasite to her unborn child, who may suffer nervous system damage. © 2013 Pearson Education, Inc. Protozoans • Ciliates are protozoans that – are named for their use of hair-like structures called cilia to move and sweep food into their mouths, – are mostly free-living (nonparasitic), such as the freshwater ciliate Paramecium, and – include heterotrophs and mixotrophs. © 2013 Pearson Education, Inc. Figure 15.22f Cell “mouth” A ciliate LM Cilia Slime Molds • Slime molds – resemble fungi in appearance and lifestyle due to convergence, but – are more closely related to amoebas. • The two main groups of these protists are – plasmodial slime molds and – cellular slime molds. © 2013 Pearson Education, Inc. Slime Molds • Plasmodial slime molds – are named for the feeding stage in their life cycle, an amoeboid mass called a plasmodium, – are decomposers on forest floors, and – can be large. © 2013 Pearson Education, Inc. Figure 15.23 A plasmodial slime mold Slime Molds • Cellular slime molds have an interesting and complex life cycle of successive stages: – a feeding stage of solitary amoeboid cells, – a swarming stage as a slug-like colony that can move and function as a single unit, and – a stage during which they generate a stalk-like multicellular reproductive structure. © 2013 Pearson Education, Inc. Figure 15.24 LM Life stages of a cellular slime mold 2 Slug-like colony 1 Amoeboid cells 3 Reproductive structure Unicellular and Colonial Algae • Algae are – photosynthetic protists whose chloroplasts support food chains in – freshwater and – marine ecosystems. • Many unicellular algae are components of plankton, the communities of mostly microscopic organisms that drift or swim weakly in aquatic environments. © 2013 Pearson Education, Inc. Unicellular and Colonial Algae • Unicellular algae include – dinoflagellates, with – two beating flagella and – external plates made of cellulose, – diatoms, with glassy cell walls containing silica, and – green algae. © 2013 Pearson Education, Inc. Unicellular and Colonial Algae • Green algae are – unicellular in most freshwater lakes and ponds, – sometimes flagellated, such as Chlamydomonas, and – sometimes colonial, forming a hollow ball of flagellated cells as seen in Volvox. © 2013 Pearson Education, Inc. (b) A sample of diverse diatoms, which have glassy walls (c) Chlamydomonas, a unicellular green alga with a pair of flagella LM Colorized SEM (a) A dinoflagellate, with its wall of protective plates LM SEM Figure 15.25 (d) Volvox, a colonial green alga Seaweeds • Seaweeds – are large, multicellular marine algae, – grow on or near rocky shores, – are only similar to plants because of convergent evolution, – are most closely related to unicellular algae, and – are often edible. © 2013 Pearson Education, Inc. Seaweeds • Seaweeds are classified into three different groups, based partly on the types of pigments present in their chloroplasts: 1. green algae, 2. red algae, and 3. brown algae (including kelp). © 2013 Pearson Education, Inc. Figure 15.26 Green algae Red algae Brown algae Evolution Connection: The Origin of Multicellular Life • Multicellular organisms have – specialized cells that are dependent on each other and perform different functions, such as – feeding, – waste disposal, – gas exchange, and – protection. © 2013 Pearson Education, Inc. Evolution Connection: The Origin of Multicellular Life • Colonial protists likely formed the evolutionary links between – unicellular and – multicellular organisms. • The colonial green alga Volvox demonstrates one level of specialization and cooperation. © 2013 Pearson Education, Inc. Figure 15.27-3 Unicellular protist An ancestral colony may have formed when a cell divided and remained attached to its offspring. Cells in the colony may have become specialized and interdependent. Additional specialization may have led to sex cells (gametes) and nonreproductive cells (somatic cells). Foodsynthesizing cells Locomotor cells Gamete Somatic cells Figure 15.UN01 Bacteria Prokaryotes Archaea Protists Eukarya Plants Fungi Animals Figure 15.UN03 Major episode Millions of years ago Plants and fungi colonize land All major animal phyla established First multicellular organisms Oldest eukaryotic fossils Accumulation of O2 in atmosphere Oldest prokaryotic fossils Origin of Earth 500 530 1,200 1,800 2,400 3,500 4,600 Figure 15.UN04 Inorganic compounds 1 Abiotic synthesis of organic monomers Organic monomers 2 Abiotic synthesis of polymers 3 Formation of pre-cells Polymer Membrane-enclosed compartment 4 Complementary chain Self-replicating molecules Figure 15.UN05 Spherical Rod-shaped Spiral Figure 15.UN06 Nutritional Mode Energy Source Photoautotroph Sunlight Carbon Source CO2 Chemoautotroph Inorganic chemicals Photoheterotroph Sunlight Organic compounds Chemoheterotroph Organic compounds