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CAMPBELL BIOLOGY IN FOCUS Urry • Cain • Wasserman • Minorsky • Jackson • Reece 24 Early Life and the Diversification of Prokaryotes Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge © 2014 Pearson Education, Inc. Overview: The First Cells  Earth formed 4.6 billion years ago  The oldest fossil organisms are prokaryotes dating back to 3.5 billion years ago  Prokaryotes are single-celled organisms in the domains Bacteria and Archaea  Some of the earliest prokaryotic cells lived in dense mats that resembled stepping stones © 2014 Pearson Education, Inc. Figure 24.1 © 2014 Pearson Education, Inc.  Prokaryotes are the most abundant organisms on Earth  There are more in a handful of fertile soil than the number of people who have ever lived  Prokaryotes thrive almost everywhere, including places too acidic, salty, cold, or hot for most other organisms  Some prokaryotes colonize the bodies of other organisms © 2014 Pearson Education, Inc. Figure 24.2 © 2014 Pearson Education, Inc. Concept 24.1: 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 protocells, membranebound droplets that maintain a consistent internal chemistry 4. Origin of self-replicating molecules © 2014 Pearson Education, Inc. Synthesis of Organic Compounds on Early Earth  Earth’s early atmosphere likely contained water vapor and chemicals released by volcanic eruptions (nitrogen, nitrogen oxides, carbon dioxide, methane, ammonia, and hydrogen)  As Earth cooled, water vapor condensed into oceans, and most of the hydrogen escaped into space © 2014 Pearson Education, Inc.  In the 1920s, A. I. Oparin and J. B. S. Haldane hypothesized that the early atmosphere was a reducing environment  In 1953, Stanley Miller and Harold Urey conducted lab experiments that showed that the abiotic synthesis of organic molecules in a reducing atmosphere is possible © 2014 Pearson Education, Inc.  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 volcanoes or deep-sea vents  Miller-Urey-type experiments demonstrate that organic molecules could have formed with various possible atmospheres  Organic molecules have also been found in meteorites Video: Hydrothermal Vent © 2014 Pearson Education, Inc. Video: Tubeworms Mass of amino acids (mg) Number of amino acids Figure 24.3 20 10 0 200 100 0 1953 © 2014 Pearson Education, Inc. 2008 1953 2008 Figure 24.3a © 2014 Pearson Education, Inc. Abiotic Synthesis of Macromolecules  RNA monomers have been produced spontaneously from simple molecules  Small organic molecules polymerize when they are concentrated on hot sand, clay, or rock © 2014 Pearson Education, Inc. Protocells  Replication and metabolism are key properties of life and may have appeared together  Protocells 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 © 2014 Pearson Education, Inc.  Adding clay can increase the rate of vesicle formation  Vesicles exhibit simple reproduction and metabolism and maintain an internal chemical environment © 2014 Pearson Education, Inc. Relative turbidity, an index of vesicle number Figure 24.4 0.4 Precursor molecules plus montmorillonite clay 0.2 Precursor molecules only 0 0 40 20 Time (minutes) 60 Vesicle boundary 1 m (a) Self-assembly 20 m (b) Reproduction © 2014 Pearson Education, Inc. (c) Absorption of RNA Relative turbidity, an index of vesicle number Figure 24.4a 0.4 Precursor molecules plus montmorillonite clay 0.2 Precursor molecules only 0 0 (a) Self-assembly © 2014 Pearson Education, Inc. 40 20 Time (minutes) 60 Figure 24.4b 20 m (b) Reproduction © 2014 Pearson Education, Inc. Figure 24.4c Vesicle boundary 1 m (c) Absorption of RNA © 2014 Pearson Education, Inc. Self-Replicating RNA  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 RNA © 2014 Pearson Education, Inc.  Natural selection has produced self-replicating RNA molecules  RNA molecules that were more stable or replicated more quickly would have left the most descendant RNA molecules  The early genetic material might have formed an “RNA world” © 2014 Pearson Education, Inc.  Vesicles with RNA capable of replication would have been protocells  RNA could have provided the template for DNA, a more stable genetic material © 2014 Pearson Education, Inc. Fossil Evidence of Early Life  Many of the oldest fossils are stromatolites, layered rocks that formed from the activities of prokaryotes up to 3.5 billion years ago  Ancient fossils of individual prokaryotic cells have also been discovered  For example, fossilized prokaryotic cells have been found in 3.4-billion-year-old rocks from Australia © 2014 Pearson Education, Inc. 5 cm 30 m Figure 24.5 10 m Stromatolites Nonphotosynthetic bacteria Possible earliest appearance in fossil record 4 © 2014 Pearson Education, Inc. Cyanobacteria 3 2 Time (billions of years ago) 1 0 Figure 24.5a Stromatolites Nonphotosynthetic bacteria Possible earliest appearance in fossil record 4 © 2014 Pearson Education, Inc. Cyanobacteria 1 3 2 Time (billions of years ago) 0 30 m Figure 24.5b 3-billion-year-old fossil of a cluster of nonphotosynthetic prokaryote cells © 2014 Pearson Education, Inc. 5 cm Figure 24.5c 1.1-billion-year-old fossilized stromatolite © 2014 Pearson Education, Inc. Figure 24.5d 10 m 1.5-billion-year-old fossil of a cyanobacterium © 2014 Pearson Education, Inc.  The cyanobacteria that form stromatolites were the main photosynthetic organisms for over a billion years  Early cyanobacteria began the release of oxygen into Earth’s atmosphere  Surviving prokaryote lineages either avoided or adapted to the newly aerobic environment © 2014 Pearson Education, Inc. Concept 24.2: Diverse structural and metabolic adaptations have evolved in prokaryotes  Most prokaryotes are unicellular, although some species form colonies  Most prokaryotic cells have diameters of 0.5–5 µm, much smaller than the 10–100 µm diameter of many eukaryotic cells  Prokaryotic cells have a variety of shapes  The three most common shapes are spheres (cocci), rods (bacilli), and spirals © 2014 Pearson Education, Inc. 1 m (a) Spherical (b) Rod-shaped © 2014 Pearson Education, Inc. 3 m 1 m Figure 24.6 (c) Spiral 1 m Figure 24.6a (a) Spherical © 2014 Pearson Education, Inc. 1 m Figure 24.6b (b) Rod-shaped © 2014 Pearson Education, Inc. 3 m Figure 24.6c (c) Spiral © 2014 Pearson Education, Inc. Cell-Surface Structures  A key feature of nearly all prokaryotic cells is their cell wall, which maintains cell shape, protects the cell, and prevents it from bursting in a hypotonic environment  Eukaryote cell walls are made of cellulose or chitin  Bacterial cell walls contain peptidoglycan, a network of modified sugars cross-linked by polypeptides © 2014 Pearson Education, Inc.  Archaeal cell walls contain polysaccharides and proteins but lack peptidoglycan  Scientists use the Gram stain to classify bacteria by cell wall composition  Gram-positive bacteria have simpler walls with a large amount of peptidoglycan  Gram-negative bacteria have less peptidoglycan and an outer membrane that can be toxic © 2014 Pearson Education, Inc. Figure 24.7 (a) Gram-positive bacteria (b) Gram-negative bacteria Carbohydrate portion of lipopolysaccharide PeptidoCell glycan wall layer Plasma membrane Outer membrane Cell wall Peptidoglycan layer Plasma membrane Gram-positive bacteria Gram-negative bacteria 10 m © 2014 Pearson Education, Inc. Figure 24.7a (a) Gram-positive bacteria PeptidoCell glycan wall layer Plasma membrane © 2014 Pearson Education, Inc. Figure 24.7b (b) Gram-negative bacteria Carbohydrate portion of lipopolysaccharide Outer membrane Cell wall Peptidoglycan layer Plasma membrane © 2014 Pearson Education, Inc. Figure 24.7c Gram-negative bacteria Gram-positive bacteria 10 m © 2014 Pearson Education, Inc.  Many antibiotics target peptidoglycan and damage bacterial cell walls  Gram-negative bacteria are more likely to be antibiotic resistant  A polysaccharide or protein layer called a capsule covers many prokaryotes © 2014 Pearson Education, Inc. Figure 24.8 Bacterial cell wall Bacterial capsule Tonsil cell 200 nm © 2014 Pearson Education, Inc.  Some bacteria develop resistant cells called endospores when they lack an essential nutrient  Other bacteria have fimbriae, which allow them to stick to their substrate or other individuals in a colony  Pili (or sex pili) are longer than fimbriae and allow prokaryotes to exchange DNA © 2014 Pearson Education, Inc. Figure 24.9 Fimbriae 1 m © 2014 Pearson Education, Inc. Motility  In a heterogeneous environment, many bacteria exhibit taxis, the ability to move toward or away from a stimulus  Chemotaxis is the movement toward or away from a chemical stimulus © 2014 Pearson Education, Inc.  Most motile bacteria propel themselves by flagella scattered about the surface or concentrated at one or both ends  Flagella of bacteria, archaea, and eukaryotes are composed of different proteins and likely evolved independently © 2014 Pearson Education, Inc. Figure 24.10 Flagellum Filament Hook Motor Cell wall Plasma membrane © 2014 Pearson Education, Inc. Rod Peptidoglycan layer 20 nm Figure 24.10a 20 nm Hook Motor © 2014 Pearson Education, Inc. Evolutionary Origins of Bacterial Flagella  Bacterial flagella are composed of a motor, hook, and filament  Many of the flagella’s proteins are modified versions of proteins that perform other tasks in bacteria  Flagella likely evolved as existing proteins were added to an ancestral secretory system  This is an example of exaptation, where existing structures take on new functions through descent with modification © 2014 Pearson Education, Inc. Internal Organization and DNA  Prokaryotic cells usually lack complex compartmentalization  Some prokaryotes do have specialized membranes that perform metabolic functions  These are usually infoldings of the plasma membrane © 2014 Pearson Education, Inc. Figure 24.11 1 m 0.2 m Respiratory membrane Thylakoid membranes (a) Aerobic prokaryote © 2014 Pearson Education, Inc. (b) Photosynthetic prokaryote Figure 24.11a 0.2 m Respiratory membrane (a) Aerobic prokaryote © 2014 Pearson Education, Inc. Figure 24.11b 1 m Thylakoid membranes (b) Photosynthetic prokaryote © 2014 Pearson Education, Inc.  The prokaryotic genome has less DNA than the eukaryotic genome  Most of the genome consists of a circular chromosome  The chromosome is not surrounded by a membrane; it is located in the nucleoid region  Some species of bacteria also have smaller rings of DNA called plasmids © 2014 Pearson Education, Inc. Figure 24.12 Chromosome Plasmids 1 m © 2014 Pearson Education, Inc.  There are some differences between prokaryotes and eukaryotes in DNA replication, transcription, and translation  These allow people to use some antibiotics to inhibit bacterial growth without harming themselves © 2014 Pearson Education, Inc. Nutritional and Metabolic Adaptations  Prokaryotes can be categorized by how they obtain energy and carbon  Phototrophs obtain energy from light  Chemotrophs obtain energy from chemicals  Autotrophs require CO2 as a carbon source  Heterotrophs require an organic nutrient to make organic compounds © 2014 Pearson Education, Inc.  Energy and carbon sources are combined to give four major modes of nutrition  Photoautotrophy  Chemoautotrophy  Photoheterotrophy  Chemoheterotrophy © 2014 Pearson Education, Inc. Table 24.1 © 2014 Pearson Education, Inc. The Role of Oxygen in Metabolism  Prokaryotic metabolism varies with respect to O2  Obligate aerobes require O2 for cellular respiration  Obligate anaerobes are poisoned by O2 and use fermentation or anaerobic respiration, in which substances other than O2 act as electron acceptors  Facultative anaerobes can survive with or without O2 © 2014 Pearson Education, Inc. Nitrogen Metabolism  Nitrogen is essential for the production of amino acids and nucleic acids  Prokaryotes can metabolize nitrogen in a variety of ways  In nitrogen fixation, some prokaryotes convert atmospheric nitrogen (N2) to ammonia (NH3) © 2014 Pearson Education, Inc. Metabolic Cooperation  Cooperation between prokaryotes allows them to use environmental resources they could not use as individual cells  In the cyanobacterium Anabaena, photosynthetic cells and nitrogen-fixing cells called heterocysts (or heterocytes) exchange metabolic products © 2014 Pearson Education, Inc. Figure 24.13 Photosynthetic cells Heterocyst 20 m © 2014 Pearson Education, Inc.  In some prokaryotic species, metabolic cooperation occurs in surface-coating colonies called biofilms © 2014 Pearson Education, Inc. Reproduction  Prokaryotes reproduce quickly by binary fission and can divide every 1–3 hours  Key features of prokaryotic biology allow them to divide quickly  They are small  They reproduce by binary fission  They have short generation times © 2014 Pearson Education, Inc. Adaptations of Prokaryotes: A Summary  The ongoing success of prokaryotes is an extraordinary example of physiological and metabolic diversification  Prokaryotic diversification can be viewed as a first great wave of adaptive radiation in the evolutionary history of life © 2014 Pearson Education, Inc. Concept 24.3: Rapid reproduction, mutation, and genetic recombination promote genetic diversity in prokaryotes  Prokaryotes have considerable genetic variation  Three factors contribute to this genetic diversity  Rapid reproduction  Mutation  Genetic recombination © 2014 Pearson Education, Inc. Rapid Reproduction and Mutation  Prokaryotes reproduce by binary fission, and offspring cells are generally identical  Mutation rates during binary fission are low, but because of rapid reproduction, mutations can accumulate rapidly in a population  High diversity from mutations allows for rapid evolution  Prokaryotes are not “primitive” but are highly evolved © 2014 Pearson Education, Inc. Figure 24.14 Experiment Daily serial transfer 0.1 mL (population sample) Old tube (discarded after transfer) New tube (9.9 mL growth medium) Population growth rate (relative to ancestral population) Results 1.8 1.6 1.4 1.2 1.0 0 © 2014 Pearson Education, Inc. 5,000 10,000 15,000 Generation 20,000 Figure 24.14a Population growth rate (relative to ancestral population) Results 1.8 1.6 1.4 1.2 1.0 0 © 2014 Pearson Education, Inc. 5,000 10,000 15,000 Generation 20,000 Genetic Recombination  Genetic recombination, the combining of DNA from two sources, contributes to diversity  Prokaryotic DNA from different individuals can be brought together by transformation, transduction, and conjugation  Movement of genes among individuals from different species is called horizontal gene transfer © 2014 Pearson Education, Inc. Transformation and Transduction  A prokaryotic cell can take up and incorporate foreign DNA from the surrounding environment in a process called transformation  Transduction is the movement of genes between bacteria by bacteriophages (viruses that infect bacteria) © 2014 Pearson Education, Inc. Figure 24.15-1 1 Phage infects bacterial donor cell with A and B alleles. Phage DNA A B Donor cell © 2014 Pearson Education, Inc. Figure 24.15-2 1 Phage infects bacterial donor cell with A and B alleles. Phage DNA A B Donor cell 2 Phage DNA is replicated and proteins synthesized. © 2014 Pearson Education, Inc. A B Figure 24.15-3 1 Phage infects bacterial donor cell with A and B alleles. Phage DNA A B Donor cell 2 Phage DNA is replicated and proteins synthesized. A B 3 Fragment of DNA with A allele is packaged within a phage capsid. © 2014 Pearson Education, Inc. A Figure 24.15-4 Phage DNA 1 Phage infects bacterial donor cell with A and B alleles. A B Donor cell 2 Phage DNA is A replicated and proteins synthesized. B 3 Fragment of DNA with A allele is packaged within a phage capsid. 4 Phage with A allele infects bacterial recipient cell. A Crossing over A A− B− Recipient cell © 2014 Pearson Education, Inc. Figure 24.15-5 Phage DNA 1 Phage infects bacterial donor cell with A and B alleles. A B Donor cell 2 Phage DNA is A replicated and proteins synthesized. B 3 Fragment of DNA with A allele is packaged within a phage capsid. 4 Phage with A A Crossing over allele infects bacterial recipient cell. 5 Incorporation of phage DNA creates recombinant cell with genotype AB. © 2014 Pearson Education, Inc. A A− B− Recipient cell Recombinant cell A B− Conjugation and Plasmids  Conjugation is the process where genetic material is transferred between prokaryotic cells  In bacteria, the DNA transfer is one way  In E. coli, the donor cell attaches to a recipient by a pilus, pulls it closer, and transfers DNA © 2014 Pearson Education, Inc. Figure 24.16 1 m Sex pilus © 2014 Pearson Education, Inc.  The F factor is a piece of DNA required for the production of pili  Cells containing the F plasmid (F+) function as DNA donors during conjugation  Cells without the F factor (F–) function as DNA recipients during conjugation  The F factor is transferable during conjugation © 2014 Pearson Education, Inc. Figure 24.17-1 F plasmid F cell (donor) Bacterial chromosome Mating bridge F− cell (recipient) Bacterial chromosome 1 One strand of F cell plasmid DNA breaks at arrowhead. © 2014 Pearson Education, Inc. Figure 24.17-2 F plasmid F cell (donor) Bacterial chromosome Mating bridge F− cell (recipient) Bacterial chromosome 1 One strand of F cell plasmid DNA breaks at arrowhead. © 2014 Pearson Education, Inc. 2 Broken strand peels off and enters F− cell. Figure 24.17-3 F plasmid F cell (donor) Bacterial chromosome Mating bridge F− cell (recipient) Bacterial chromosome 1 One strand of F cell plasmid DNA breaks at arrowhead. © 2014 Pearson Education, Inc. 2 Broken strand peels off and enters F− cell. 3 Donor and recipient cells synthesize complementary DNA strands. Figure 24.17-4 F plasmid F cell (donor) Bacterial chromosome F cell Mating bridge F− cell (recipient) F cell Bacterial chromosome 1 One strand of F cell plasmid DNA breaks at arrowhead. © 2014 Pearson Education, Inc. 2 Broken strand peels off and enters F− cell. 3 Donor and recipient cells synthesize complementary DNA strands. 4 Recipient cell is now a recombinant F cell.  The F factor can also be integrated into the chromosome  A cell with the F factor built into its chromosomes functions as a donor during conjugation  The recipient becomes a recombinant bacterium, with DNA from two different cells © 2014 Pearson Education, Inc. R Plasmids and Antibiotic Resistance  Genes for antibiotic resistance are carried in R plasmids  Antibiotics kill sensitive bacteria, but not bacteria with specific R plasmids  Through natural selection, the fraction of bacteria with genes for resistance increases in a population exposed to antibiotics  Antibiotic-resistant strains of bacteria are becoming more common © 2014 Pearson Education, Inc. Concept 24.4: Prokaryotes have radiated into a diverse set of lineages  Prokaryotes have radiated extensively due to diverse structural and metabolic adaptations  Prokaryotes inhabit every environment known to support life © 2014 Pearson Education, Inc. An Overview of Prokaryotic Diversity  Applying molecular systematics to the investigation of prokaryotic phylogeny has produced dramatic results  Molecular systematics led to the splitting of prokaryotes into bacteria and archaea  Molecular systematists continue to work on the phylogeny of prokaryotes © 2014 Pearson Education, Inc. Figure 24.18 Euryarchaeotes Crenarchaeotes UNIVERSAL ANCESTOR Nanoarchaeotes Domain Archaea Korarchaeotes Domain Eukarya Eukaryotes Proteobacteria Spirochetes Cyanobacteria Gram-positive bacteria © 2014 Pearson Education, Inc. Domain Bacteria Chlamydias  The use of polymerase chain reaction (PCR) has allowed for more rapid sequencing of prokaryote genomes  A handful of soil may contain 10,000 prokaryotic species  Horizontal gene transfer between prokaryotes obscures the root of the tree of life © 2014 Pearson Education, Inc. Bacteria  Bacteria include the vast majority of prokaryotes familiar to most people  Diverse nutritional types are scattered among the major groups of bacteria Video: Tubeworms © 2014 Pearson Education, Inc. Figure 24.UN01 Eukarya Archaea Bacteria © 2014 Pearson Education, Inc. Figure 24.19a Beta subgroup Alpha subgroup © 2014 Pearson Education, Inc. 1 m Nitrosomonas (TEM) Delta subgroup Epsilon subgroup Chondromyces crocatus (SEM) 2 m Rhizobium (arrows) (TEM) 300 m Thiomargarita namibiensis (LM) 200 m Gamma subgroup 2.5 m Alpha Beta Gamma Proteobacteria Delta Epsilon Helicobacter pylori (TEM)  Proteobacteria are gram-negative bacteria including photoautotrophs, chemoautotrophs, and heterotrophs  Some are anaerobic and others aerobic © 2014 Pearson Education, Inc. Figure 24.19aa Alpha Beta Gamma Proteobacteria Delta Epsilon © 2014 Pearson Education, Inc.  Members of the subgroup alpha proteobacteria are closely associated with eukaryotic hosts in many cases  Scientists hypothesize that mitochondria evolved from aerobic alpha proteobacteria through endosymbiosis  Example: Rhizobium, which forms root nodules in legumes and fixes atmospheric N2  Example: Agrobacterium, which produces tumors in plants and is used in genetic engineering © 2014 Pearson Education, Inc. Figure 24.19ab Alpha subgroup 2.5 m Rhizobium (arrows) inside a root cell of a legume (TEM) © 2014 Pearson Education, Inc.  Members of the subgroup beta proteobacteria are nutritionally diverse  Example: the soil bacterium Nitrosomonas, which converts NH4+ to NO2– © 2014 Pearson Education, Inc. Figure 24.19ac Beta subgroup 1 m Nitrosomonas (colorized TEM) © 2014 Pearson Education, Inc.  The subgroup gamma proteobacteria includes sulfur bacteria such as Thiomargarita namibiensis and pathogens such as Legionella, Salmonella, and Vibrio cholerae  Escherichia coli resides in the intestines of many mammals and is not normally pathogenic © 2014 Pearson Education, Inc. Figure 24.19ad Gamma subgroup 200 m Thiomargarita namibiensis containing sulfur wastes (LM) © 2014 Pearson Education, Inc.  The subgroup delta proteobacteria includes the slime-secreting myxobacteria and bdellovibrios, a bacteria that attacks other bacteria © 2014 Pearson Education, Inc. Figure 24.19ae Delta subgroup 300 m Fruiting bodies of Chondromyces crocatus, a myxobacterium (SEM) © 2014 Pearson Education, Inc.  The subgroup epsilon proteobacteria contains many pathogens including Campylobacter, which causes blood poisoning, and Helicobacter pylori, which causes stomach ulcers © 2014 Pearson Education, Inc. Figure 24.19af Epsilon subgroup 2 m Helicobacter pylori (colorized TEM) © 2014 Pearson Education, Inc. Figure 24.19b Cyanobacteria Chlamydia (arrows) (TEM) 5 m 2.5 m Spirochetes Oscillatoria Leptospira (TEM) 5 m 2 m Gram-positive bacteria Streptomyces (SEM) © 2014 Pearson Education, Inc. 40 m Chlamydias Mycoplasmas (SEM)  Chlamydias are parasites that live within animal cells  Chlamydia trachomatis causes blindness and nongonococcal urethritis by sexual transmission © 2014 Pearson Education, Inc. Figure 24.19ba Chlamydias 2.5 m Chlamydia (arrows) inside an animal cell (colorized TEM) © 2014 Pearson Education, Inc.  Spirochetes are helical heterotrophs  Some are parasites, including Treponema pallidum, which causes syphilis, and Borrelia burgdorferi, which causes Lyme disease © 2014 Pearson Education, Inc. Figure 24.19bb Spirochetes 5 m Leptospira, a spirochete (colorized TEM) © 2014 Pearson Education, Inc.  Cyanobacteria are photoautotrophs that generate O2  Plant chloroplasts likely evolved from cyanobacteria by the process of endosymbiosis © 2014 Pearson Education, Inc. Figure 24.19bc Cyanobacteria 40 m Oscillatoria, a filamentous cyanobacterium © 2014 Pearson Education, Inc.  Gram-positive bacteria include  Actinomycetes, which decompose soil  Streptomyces, which are a source of antibiotics  Bacillus anthracis, the cause of anthrax  Clostridium botulinum, the cause of botulism  Some Staphylococcus and Streptococcus, which can be pathogenic  Mycoplasms, the smallest known cells © 2014 Pearson Education, Inc. Figure 24.19bd Gram-positive bacteria 5 m Streptomyces, the source of many antibiotics (SEM) © 2014 Pearson Education, Inc. Figure 24.19be Gram-positive bacteria 2 m Hundreds of mycoplasmas covering a human fibroblast cell (colorized SEM) © 2014 Pearson Education, Inc. Archaea  Archaea share certain traits with bacteria and other traits with eukaryotes © 2014 Pearson Education, Inc. Figure 24.UN02 Eukarya Archaea Bacteria © 2014 Pearson Education, Inc. Table 24.2 © 2014 Pearson Education, Inc. Table 24.2a © 2014 Pearson Education, Inc. Table 24.2b © 2014 Pearson Education, Inc.  Some archaea live in extreme environments and are called extremophiles  Extreme halophiles live in highly saline environments  Extreme thermophiles thrive in very hot environments Video: Cyanobacteria (Oscillatoria) © 2014 Pearson Education, Inc. Figure 24.20 © 2014 Pearson Education, Inc.  Methanogens produce methane as a waste product  Methanogens are strict anaerobes and are poisoned by O2  Methanogens live in swamps and marshes, in the guts of cattle, and near deep-sea hydrothermal vents © 2014 Pearson Education, Inc. Figure 24.21 2 m © 2014 Pearson Education, Inc. Figure 24.21a © 2014 Pearson Education, Inc. Figure 24.21b 2 m © 2014 Pearson Education, Inc.  Recent metagenomic studies have revealed many new groups of archaea  Some of these may offer clues to the early evolution of life on Earth © 2014 Pearson Education, Inc. Concept 24.5: Prokaryotes play crucial roles in the biosphere  Prokaryotes are so important that if they were to disappear, the prospects for any other life surviving would be dim © 2014 Pearson Education, Inc. Chemical Recycling  Prokaryotes play a major role in the recycling of chemical elements between the living and nonliving components of ecosystems  Chemoheterotrophic prokaryotes function as decomposers, breaking down dead organisms and waste products © 2014 Pearson Education, Inc.  Prokaryotes can sometimes increase the availability of nitrogen, phosphorus, and potassium for plant growth  Prokaryotes can also “immobilize” or decrease the availability of nutrients © 2014 Pearson Education, Inc. Uptake of K by plants (mg) Figure 24.22 1.0 0.8 0.6 0.4 0.2 Seedlings growing in the lab 0 No Strain 1 Strain 2 Strain 3 bacteria Soil treatment © 2014 Pearson Education, Inc. Figure 24.22a Seedlings growing in the lab © 2014 Pearson Education, Inc. Ecological Interactions  Symbiosis is an ecological relationship in which two species live in close contact: a larger host and smaller symbiont  Prokaryotes often form symbiotic relationships with larger organisms © 2014 Pearson Education, Inc.  In mutualism, both symbiotic organisms benefit  In commensalism, one organism benefits while neither harming nor helping the other in any significant way  In parasitism, an organism called a parasite harms but does not kill its host  Parasites that cause disease are called pathogens © 2014 Pearson Education, Inc. Figure 24.23 © 2014 Pearson Education, Inc.  The ecological communities of hydrothermal vents depend on chemoautotrophic bacteria for energy © 2014 Pearson Education, Inc. Impact on Humans  The best-known prokaryotes are pathogens, but many others have positive interactions with humans © 2014 Pearson Education, Inc. Mutualistic Bacteria  Human intestines are home to about 500–1,000 species of bacteria  Many of these are mutualists and break down food that is undigested by our intestines © 2014 Pearson Education, Inc. Pathogenic Bacteria  Prokaryotes cause about half of all human diseases  For example, Lyme disease is caused by a bacterium and carried by ticks © 2014 Pearson Education, Inc. Figure 24.24 5 m © 2014 Pearson Education, Inc. Figure 24.24a © 2014 Pearson Education, Inc. Figure 24.24b © 2014 Pearson Education, Inc. Figure 24.24c 5 m © 2014 Pearson Education, Inc.  Pathogenic prokaryotes typically cause disease by releasing exotoxins or endotoxins  Exotoxins are secreted and cause disease even if the prokaryotes that produce them are not present  Endotoxins are released only when bacteria die and their cell walls break down © 2014 Pearson Education, Inc.  Horizontal gene transfer can spread genes associated with virulence  For example, pathogenic strains of the normally harmless E. coli bacteria have emerged through horizontal gene transfer © 2014 Pearson Education, Inc. Prokaryotes in Research and Technology  Experiments using prokaryotes have led to important advances in DNA technology  For example, E. coli is used in gene cloning  For example, Agrobacterium tumefaciens is used to produce transgenic plants © 2014 Pearson Education, Inc.  Bacteria can now be used to make natural plastics  Prokaryotes are the principal agents in bioremediation, the use of organisms to remove pollutants from the environment  Bacteria can be engineered to produce vitamins, antibiotics, and hormones  Bacteria are also being engineered to produce ethanol from waste biomass © 2014 Pearson Education, Inc. Figure 24.25 (b) (a) © 2014 Pearson Education, Inc. Figure 24.25a (a) © 2014 Pearson Education, Inc. Figure 24.25b (b) © 2014 Pearson Education, Inc. Figure 24.26 © 2014 Pearson Education, Inc. Figure 24.UN03 © 2014 Pearson Education, Inc. Figure 24.UN04 Fimbriae Cell wall Circular chromosome Capsule Sex pilus Internal organization Flagella © 2014 Pearson Education, Inc. Figure 24.UN05 © 2014 Pearson Education, Inc.