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Chapter 15 The Evolution of Microbial Life PowerPoint® Lectures for Campbell Essential Biology, Fourth Edition – Eric Simon, Jane Reece, and Jean Dickey Campbell Essential Biology with Physiology, Third Edition – Eric Simon, Jane Reece, and Jean Dickey Lectures by Chris C. Romero, updated by Edward J. Zalisko © 2010 Pearson Education, Inc. Biology and Society: Can Life Be Created in the Lab? • How did life first arise on Earth? • To gain insight, scientists have – Synthesized the entire genome of Mycoplasma genitalium, a species of bacteria found naturally in the human urinary tract – Transplanted the complete genome of one species of Mycoplasma bacteria into another © 2010 Pearson Education, Inc. Figure 15.00 • A research group led by Craig Venter hopes to – Create an artificial genome – Transplant it into a genome-free host cell • An artificial organism that could be completely controlled might – Clean up toxic wastes – Generate biofuels – Be unable to survive outside rigidly controlled conditions © 2010 Pearson Education, Inc. MAJOR EPISODES IN THE HISTORY OF LIFE • Earth was formed about 4.6 billion years ago. • Prokaryotes – Evolved by 3.5 billion years ago – Began oxygen production about 2.7 billion years ago – Lived alone for almost 2 billion years – Continue in great abundance today © 2010 Pearson Education, Inc. • Single-celled eukaryotes first evolved about 2.1 billion years ago. • Multicellular eukaryotes first evolved at least 1.2 billion years ago. © 2010 Pearson Education, Inc. Precambrian Common ancestor to all present-day life Origin of Earth 4,500 Earth cool enough for crust to solidify 4,000 Oldest prokaryotic fossils 3,500 Millions of years ago 3,000 Atmospheric oxygen begins to appear due to photosynthetic prokaryotes 2,500 Figure 15.1a Paleozoic Mesozoic Cenozoic Archaea Prokaryotes Bacteria Protists Eukaryotes Plants Fungi Animals Oldest eukaryotic fossils 2,000 Cambrian explosion Oldest Plants and animal symbiotic fungi fossils colonize land Origin of multicellular organisms 1,500 1,000 Millions of years ago 500 Extinction of dinosaurs First humans 0 Figure 15.1b • 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 – Were followed by amphibians that evolved from fish © 2010 Pearson Education, Inc. • What if we use a clock analogy to tick down all of the major events in the history of life on Earth? © 2010 Pearson Education, Inc. Humans Origin of solar system and Earth 0 4 1 2 3 Figure 15.2 THE ORIGIN OF LIFE • We may never know for sure how life on Earth began. © 2010 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 19th century, it was commonly believed that life regularly arises from nonliving matter, an idea called spontaneous generation. © 2010 Pearson Education, Inc. • Today, most biologists think it is possible that life on early Earth produced simple cells by chemical and physical processes. © 2010 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. © 2010 Pearson Education, Inc. Stage 1: Abiotic Synthesis of Organic Monomers • The first stage in the origin of life has been the most extensively studied by scientists in the laboratory. © 2010 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 the early Earth. • Question: Could biological molecules arise spontaneously under conditions like those on the early Earth? © 2010 Pearson Education, Inc. • Hypothesis: A closed system designed in the laboratory to simulate early Earth conditions could produce biologically important organic molecules from inorganic ingredients. • Prediction: Organic molecules would form and accumulate. © 2010 Pearson Education, Inc. • 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, discharged into the chamber to mimic the prevalent lightning of the early Earth – A condenser to cool the atmosphere, causing water and dissolved compounds to “rain” into the miniature “sea” © 2010 Pearson Education, Inc. “Atmosphere” CH4 Water vapor NH3 H2 Electrode Condenser Cold water Cooled water containing organic molecules H2O Stanley Miller re-creating his 1953 experiment “Sea” Sample for chemical analysis Miller and Urey’s experiment Figure 15.4 • 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. • Since Miller and Urey’s experiments, laboratory analogues of the primeval Earth have produced – All 20 amino acids – Several sugars © 2010 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 – Rock © 2010 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. © 2010 Pearson Education, Inc. • 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 – Swell or shrink when placed in solutions of different salt concentrations © 2010 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. © 2010 Pearson Education, Inc. Original “gene” Complementary RNA chain Figure 15.5 From Chemical Evolution to Darwinian Evolution • Over millions of years – Natural selection favored the most efficient pre-cells – The first prokaryotic cells evolved © 2010 Pearson Education, Inc. PROKARYOTES • Prokaryotes lived and evolved all alone on Earth for 2 billion years before eukaryotes evolved. © 2010 Pearson Education, Inc. They’re Everywhere! • Prokaryotes – Are found wherever there is life – Far outnumber eukaryotes – Can cause disease – Can be beneficial © 2010 Pearson Education, Inc. • Prokaryotes live deep within the Earth and in habitats too cold, too hot, too salty, too acidic, or too alkaline for any eukaryote to survive. © 2010 Pearson Education, Inc. • Compared to eukaryotes, prokaryotes are – Much more abundant – Typically much smaller © 2010 Pearson Education, Inc. Figure 15.7 Colorized SEM • Prokaryotes – Are ecologically significant, recycling carbon and other vital chemical elements back and forth between organic matter, the soil, and atmosphere – Cause about half of all human diseases – Are more typically benign or beneficial © 2010 Pearson Education, Inc. The Structure and Function of Prokaryotes • Prokaryotic cells – Lack true nuclei – Lack other membrane-enclosed organelles – Have cell walls exterior to their plasma membranes © 2010 Pearson Education, Inc. Plasma membrane (encloses cytoplasm) Cell wall (provides Rigidity) Capsule (sticky coating) Colorized TEM Prokaryotic flagellum (for propulsion) Ribosomes (synthesize proteins) Nucleoid (contains DNA) Pili (attachment structures) Figure 4.4 Ribosomes Cytoskeleton Centriole Lysosome Flagellum Not in most plant cells Plasma membrane Nucleus Mitochondrion Rough endoplasmic reticulum (ER) Golgi apparatus Cytoskeleton Mitochondrion Nucleus Rough endoplasmic reticulum (ER) Idealized animal cell Smooth endoplasmic reticulum (ER) Central vacuole Cell wall Not in animal cells Chloroplast Ribosomes Plasma membrane Smooth endoplasmic reticulum (ER) Channels between cells Idealized plant cell Golgi apparatus Figure 4.5 Procaryotic Forms • Prokaryotes come in several shapes: – Spherical (cocci) – Rod-shaped (bacilli) – Spiral © 2010 Pearson Education, Inc. SHAPES OF PROKARYOTIC CELLS Colorized TEM Spiral Colorized SEM Rod-shaped (bacilli) Colorized SEM Spherical (cocci) Figure 15.8 • Most prokaryotes are – Unicellular – Very small • Some prokaryotes – Form true colonies – Show specialization of cells – Are very large © 2010 Pearson Education, Inc. (b) Cyanobacteria LM LM Colorized SEM (a) Actinomycete (c) Giant bacterium Figure 15.9 • About half of all prokaryotes are mobile, using flagella. • Many have one or more flagella that propel the cells away from unfavorable places or toward more favorable places, such as nutrient-rich locales. © 2010 Pearson Education, Inc. Colorized TEM Flagellum Plasma membrane Cell wall Rotary movement of each flagellum Figure 15.10 Procaryotic Reproduction • Most prokaryotes can reproduce by binary fission and at very high rates if conditions are favorable. • Some prokaryotes – Form endospores, thick-coated, protective cells that are produced within the cells when they are exposed to unfavorable conditions – Can survive very harsh conditions for extended periods, even centuries © 2010 Pearson Education, Inc. Colorized SEM Endospore Figure 15.11 Procaryotic Nutrition • Prokaryotes exhibit four major modes of nutrition. – Phototrophs obtain energy from light. – Chemotrophs obtain energy from environmental chemicals. – Species that obtain carbon from carbon dioxide (CO2) are autotrophs. – Species that obtain carbon from at least one organic nutrient—the sugar glucose, for instance—are called heterotrophs. © 2010 Pearson Education, Inc. • 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) © 2010 Pearson Education, Inc. MODES OF NUTRITION Energy source Chemical Photoautotrophs Colorized TEM Light Elodea, an aquatic plant Bacteria from a hot spring Colorized TEM Photoheterotrophs Organic compounds Carbon source CO2 Chemoautotrophs Rhodopseudomonas Chemoheterotrophs Little Owl (Athene noctua) Figure 15.12 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: – Bacteria – Archaea (more closely related to eukaryotes) © 2010 Pearson Education, Inc. • 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. © 2010 Pearson Education, Inc. (a) Salt-loving archaea (b) Heat-loving archaea Figure 15.13 Bacteria and Humans • Bacteria interact with humans in many ways. © 2010 Pearson Education, Inc. Bacteria That Cause Disease • Bacteria and other organisms that cause disease are called pathogens. • Most pathogenic bacteria produce poisons. – Exotoxins are poisonous proteins secreted by bacterial cells. – Endotoxins are not cell secretions but instead chemical components of the outer membrane of certain bacteria. © 2010 Pearson Education, Inc. Colorized SEM Haemophilus influenzae Cells of nasal lining Figure 15.14 • The best defenses against bacterial disease are – Sanitation – Antibiotics – Education © 2010 Pearson Education, Inc. • Lyme disease is – Caused by bacteria carried by ticks – Treated with antibiotics, if detected early © 2010 Pearson Education, Inc. SEM Tick that carries the Lyme disease bacterium “Bull’s-eye” rash Spirochete that causes Lyme disease Figure 15.15 Bioterrorism • Humans have a long and ugly history of using organisms as weapons. – During the Middle Ages, armies hurled the bodies of plague victims into enemy ranks. – Early conquerors, settlers, and warring armies in South and North America gave native peoples items purposely contaminated with infectious bacteria. – In 1984, members of a cult in Oregon contaminated restaurant salad bars with Salmonella bacteria. – In the fall of 2001, five Americans died from the disease anthrax in a presumed terrorist attack. © 2010 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. © 2010 Pearson Education, Inc. Prokaryotes and Chemical Recycling • Prokaryotes play essential roles in – Chemical cycles in the environment – The breakdown of organic wastes and dead organisms © 2010 Pearson Education, Inc. Prokaryotes and Bioremediation • Bioremediation is the use of organisms to remove pollutants from – Water – Air – Soil • A familiar example is the use of prokaryotic decomposers in sewage treatment. © 2010 Pearson Education, Inc. Rotating spray arm Rock bed coated with aerobic prokaryotes and fungi Liquid wastes Outflow Figure 15.17 • Certain bacteria – Can decompose petroleum – Are useful in cleaning up oil spills © 2010 Pearson Education, Inc. Figure 15.18 PROTISTS • Protists – Are eukaryotic – Evolved from prokaryotic ancestors – Are ancestral to all other eukaryotes, which are – Plants – Fungi – Animals © 2010 Pearson Education, Inc. Figure 15.19 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 – Endosymbiosis, one species living inside another host species, in which free-living bacteria came to reside inside a host cell, producing mitochondria and chloroplasts © 2010 Pearson Education, Inc. Plasma membrane Photosynthetic prokaryote DNA Cytoplasm Membrane infolding (Some cells) Endosymbiosis Endoplasmic reticulum Nucleus Ancestral prokaryote Aerobic heterotrophic prokaryote Chloroplast Mitochondrion Nuclear envelope Photosynthetic eukaryotic cell Cell with nucleus and endomembrane system (a) Origin of the endomembrane system (b) Origin of mitochondria and chloroplasts Figure 15.20 The Diversity of Protists • Protists can be – Unicellular – Multicellular • More than any other group, protists vary in – Structure – Function © 2010 Pearson Education, Inc. • Protists are not one distinct group but instead represent all the eukaryotes that are not plants, animals, or fungi. © 2010 Pearson Education, Inc. • The classification of protists remains a work in progress. • The four major categories of protists, grouped by lifestyle, are – Protozoans – Slime molds – Unicellular algae – Seaweeds © 2010 Pearson Education, Inc. Protozoans • Protists that live primarily by ingesting food are called protozoans. • Protozoans with flagella are called flagellates and are typically free-living, but sometimes are nasty parasites. © 2010 Pearson Education, Inc. Colorized SEM A flagellate: Giardia Figure 15.21a Colorized SEM Another flagellate: trypanosomes Figure 15.21b • Amoebas are characterized by – Great flexibility in their body shape – The absence of permanent organelles for locomotion • Most species move and feed by means of pseudopodia (singular, pseudopodium), temporary extensions of the cell. © 2010 Pearson Education, Inc. • Amoebas may have a shell, as seen in forams, or no shell at all. © 2010 Pearson Education, Inc. LM An amoeba Figure 15.21c LM A foram Figure 15.21d • Apicomplexans are – Named for a structure at their apex (tip) that is specialized for penetrating host cells and tissues – All parasitic, such as Plasmodium, which causes malaria © 2010 Pearson Education, Inc. TEM An apicomplexan Figure 15.21e • Ciliates – Are mostly free-living (nonparasitic), such as the freshwater ciliate Paramecium – Use structures called cilia to move and feed © 2010 Pearson Education, Inc. LM A ciliate Figure 15.21f LM Colorized SEM A flagellate: Giardia An amoeba Another flagellate: trypanosomes Cilia An apicomplexan LM Red blood cell Oral groove TEM LM Apical complex A foram Pseudopodium of amoeba Colorized SEM Food being ingested A ciliate Figure 15.21 Slime Molds • Slime molds resemble fungi in appearance and lifestyle, but the similarities are due to convergence, and slime molds are not at all closely related to fungi. • The two main groups of these protists are – Plasmodial slime molds – Cellular slime molds © 2010 Pearson Education, Inc. • Plasmodial slime molds – Can be large – Are decomposers on forest floors – Are named for the feeding stage in their life cycle, an amoeboid mass called a plasmodium © 2010 Pearson Education, Inc. Figure 15.22 • Cellular slime molds have an interesting and complex life cycle that changes between a – Feeding stage of solitary amoeboid cells – Sluglike colony that moves and functions as a single unit – Stalklike reproductive structure © 2010 Pearson Education, Inc. LM Slug-like colony Amoeboid cells Reproductive structure Figure 15.23 Unicellular and Colonial Algae • Algae are – Photosynthetic protists – Found in plankton, the communities of mostly microscopic organisms that drift or swim weakly in aquatic environments © 2010 Pearson Education, Inc. • Unicellular algae include – Diatoms, which have glassy cell walls containing silica – Dinoflagellates, with two beating flagella and external plates made of cellulose © 2010 Pearson Education, Inc. SEM (a) A dinoflagellate, with its wall of protective plates Figure 15.24a LM (b) A sample of diverse diatoms, which have glossy walls Figure 15.24b • Green algae are – Unicellular – Sometimes flagellated, such as Chlamydomonas – Colonial, sometimes forming a hollow ball of flagellated cells, as seen in Volvox © 2010 Pearson Education, Inc. Colorized SEM (c) Chlamydomonas, a unicellular green alga with a pair of flagella Figure 15.24c LM (d) Volvox, a colonial green alga Figure 15.24d Seaweeds • Seaweeds – Are only similar to plants because of convergent evolution – Are large, multicellular marine algae – Grow on or near rocky shores – Are often edible © 2010 Pearson Education, Inc. • Seaweeds are classified into three different groups, based partly on the types of pigments present in their chloroplasts: – Green algae – Red algae – Brown algae (including kelp) © 2010 Pearson Education, Inc. Green algae Red algae Brown algae Figure 15.25 Evolution Connection: The Origin of Multicellular Life • Multicellular organisms have interdependent, specialized cells that perform different functions, such as feeding, waste disposal, gas exchange, and protection—and are dependent on each other. © 2010 Pearson Education, Inc. • Colonial protists likely formed the evolutionary links between unicellular and multicellular organisms. © 2010 Pearson Education, Inc. Unicellular protist Gamete Locomotor cells Food-synthesizing cells Somatic cells Colony Early multicellular organism with specialized, interdependent cells Later organism with gametes and somatic cells Figure 15.26-3