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Prokaryotes Chapter 27 • They’re (almost) Everywhere! • Most prokaryotes are microscopic Prokaryotes: Domains Bacteria & Archaea – But what they lack in size they more than make up for in numbers • The number of prokaryotes in a single handful of fertile soil – Is greater than the number of people who have ever lived PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Prokaryotes thrive almost everywhere • Biologists are discovering – Including places too acidic, too salty, too cold, or too hot for most other organisms – These organisms have an astonishing genetic diversity – Not too surprising, given their 3.5+ billion years of evolution! • Structural, functional, and genetic adaptations contribute to prokaryotic success • Most prokaryotes are unicellular – Although some species form colonies Figure 27.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Domain Bacteria: a prokaryote • Prokaryotic cells have a variety of shapes Fimbriae Nucleoid – The three most common of which are spheres (cocci), rods (bacilli), and spirals (spirilla) Ribosomes Plasma membrane Bacterial chromosome (a) A typical rod-shaped bacterium Cell wall Capsule Flagella 0.5 µm (b) A thin section through the bacterium Bacillus coagulans (TEM) 1 µm Figure 27.2a–c (a) Spherical (cocci) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 µm (b) Rod-shaped (bacilli) (c) Spiral 5 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 1 Cell-Surface Structures The Gram Stain • One of the most important features of nearly all prokaryotic cells • Allows scientists to classify many bacterial species into two groups based on cell wall composition: Gram-positive (clade?) and Gram-negative – Is their cell wall, which maintains cell shape, provides physical protection, and prevents the cell from bursting in a hypotonic environment Lipopolysaccharide Cell wall Peptidoglycan layer Cell wall Outer membrane Peptidoglycan layer Plasma membrane Plasma membrane Protein Protein Grampositive bacteria Gramnegative bacteria 20 µm (a) Gram-positive. Gram-positive bacteria have a cell wall with a large amount of peptidoglycan that traps the violet dye in the cytoplasm. The alcohol rinse does not remove the violet dye, which masks the added red dye. Figure 27.3a, b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) Gram-negative. Gram-negative bacteria have less peptidoglycan, and it is located in a layer between the plasma membrane and an outer membrane. The violet dye is easily rinsed from the cytoplasm, and the cell appears pink or red after the red dye is added. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Streptococcus on tonsil cell • The cell wall of many prokaryotes • Some prokaryotes have fimbriae and pili – Is covered by a capsule, a sticky layer of polysaccharide or protein – Which allow them to stick to their substrate or other individuals in a colony 200 nm Fimbriae Capsule 200 nm Figure 27.5 Figure 27.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Motility • Most motile bacteria propel themselves by flagella – Which are structurally and functionally different from eukaryotic flagella • In a heterogeneous environment, many bacteria exhibit taxis – The ability to move toward or away from certain stimuli Flagellum Filament 50 nm Cell wall Hook Basal apparatus Figure 27.6 Plasma membrane Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 Internal and Genomic Organization Escherichia coli cell and its chromosome • Most Prokaryotic cells lack complex compartmentalization (membrane-bound organelles) • The typical prokaryotic genome • But some have specialized membranes that perform metabolic functions (recall endosymbiotic theory) 0.2 µm – Is a ring of DNA that is not surrounded by a membrane and that is located in a nucleoid region 1 µm Chromosome Respiratory membrane Thylakoid membranes Figure 27.7a, b (a) Aerobic prokaryote (b) Photosynthetic prokaryote Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 27.8 1 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fig. 27-8 Chromosome Plasmids • Some species of bacteria – Also have smaller rings of DNA called plasmids 1 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Reproduction and Adaptation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fig. 27-12 • Prokaryotes reproduce quickly by binary fission – Every 1–3 hours (or faster: 20 min) – Asexual reproduction • Prokaryotes can exchange DNA: Horizontal transfer - often via plasmids Fig. 27-12 – Conjugation: passing DNA between cells – Transformation: DNA can be acquired from the environment – Transduction: DNA transfer via viruses Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Conjugation via Sex pilus 1 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 3 Fig. 27-13 Conjugation and the F factor (fertility) Fertility (F) and Resistance (R) Plasmids • Rapid reproduction and horizontal gene transfer F plasmid Bacterial chromosome F+ cell – Facilitate the evolution of prokaryotes to changing environments F+ cell Mating bridge F– cell – Adaptations can readily be propagated within a ‘species’ and to other prokaryotes F+ cell Bacterial chromosome (a) Conjugation and transfer of an F plasmid Hfr cell A+ F factor F– cell A+ A+ A+ A– Recombinant F– bacterium A– A– A+ • E.g. R Plasmids spread anti-biotic resistance A+ A– (b) Conjugation and transfer of part of an Hfr bacterial chromosome Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Prokaryotes Exhibit Diverse Nutritional Modes • Many prokaryotes form endospores – Which can remain viable in harsh conditions for centuries Bacillus anthracis • A great diversity of nutritional and metabolic adaptations have evolved in prokaryotes – All major biochemical pathways • Examples of all four modes of nutrition are found among prokaryotes Endospore – Photoautotrophy – Chemoautotrophy – Photoheterotrophy Figure 27.9 0.3 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings – Chemoheterotrophy Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Prokaryotes Exhibit Diverse Nutritional Modes Prokaryotes Exhibit Diverse Nutritional Modes • Phototrophs: energy from sun • All four possible modes of nutrition are found among prokaryotes • Chemotrophs: energy from molecules • Autotrophs: able to build their own organic cds • Heterotrophs: require organic nutrient(s) – Photoautotrophy – Chemoautotrophy – Photoheterotrophy – Chemoheterotrophy Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4 Metabolic Relationships to Oxygen • Major nutritional modes in prokaryotes • Prokaryotic metabolism – Also varies with respect to oxygen • Obligate aerobes: Require oxygen • Facultative anaerobes: Can survive with or without oxygen • Obligate anaerobes: Are poisoned by oxygen Table 27.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Nitrogen Metabolism Some Prokaryotes Exhibit Multicellularity • Prokaryotes can metabolize nitrogen • In the cyanobacterium Anabaena (and Nostoc) – In a variety of ways • In a process called nitrogen fixation – Photosynthetic cells and nitrogen-fixing cells exchange metabolic products – Some prokaryotes convert atmospheric nitrogen to ammonia Photosynthetic cells Heterocyst Figure 27.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 20 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Metabolic Cooperation Metabolic Cooperation in Prokaryotes • Cooperation between prokaryotes • Surface-coating colonies called biofilms – Dental plaque 1 µm – Allows them to use environmental resources they could not use as individual cells Figure 27.11 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 5 Lessons from Molecular Systematics Nanoarchaeotes Domain Eukarya Eukaryotes Korarchaeotes Euryarchaeotes Gram-positive bacteria Cyanobacteria Spirochetes Chlamydias Epsilon Delta Beta • Molecular systematics Crenarchaeotes Domain Archaea Domain Bacteria Proteobacteria Alpha – Systematists based prokaryotic taxonomy on phenotypic criteria: e.g. shape, gram stain, nutritional mode • A tentative phylogeny of some of the major taxa of prokaryotes based on molecular systematics Gamma • Until the late 20th century – Is leading to a phylogenetic classification of prokaryotes, and – Identification of major new clades Universal ancestor Figure 27.12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Bacteria • Proteobacteria α proteobacteria closest living relatives of mitochondria 2.5 µm • Diverse nutritional types – Are scattered among the major groups of bacteria 1 µm Rhizobium (arrows) inside a root cell of a legume (TEM) • The two largest groups are Nitrosomonas (colorized TEM) 0.5 µm – The proteobacteria and the Gram-positive bacteria Fruiting bodies of Chondromyces crocatus, a myxobacterium (SEM) 5 µm 10 µm Chromatium; the small globules are sulfur wastes (LM) 2 µm Bdellovibrio bacteriophorus Attacking a larger bacterium (colorized TEM) Figure 27.13 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Helicobacter pylori (colorized TEM). Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Archaea 2.5 µm • Chlamydias, spirochetes, Gram-positive bacteria, and cyanobacteria • Archaea share certain traits with bacteria – And other traits with eukaryotes 5 µm Chlamydia (arrows) inside an animal cell (colorized TEM) 1 µm 5 µm Leptospira, a spirochete (colorized TEM) 50 µm Hundreds of mycoplasmas covering a human fibroblast cell Streptomyces, the source of many antibiotics (colorized SEM) (colorized SEM) Figure 27.13 Two species of Oscillatoria, filamentous cyanobacteria (LM) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Table 27.2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 6 Some Archaea are ‘Extremophiles’ Archaea • Extremophiles ‘love’ extreme environments • Extreme halophiles – Live in extreme environments – Live in high saline environments • Extreme thermophiles – Thrive in very hot environments • e.g. Hydro-thermal vents Figure 27.14 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Archaea • Methanogens Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Text Art 5.02 Prokaryotes are Superabundant; New Genomics: Up to 50,000 species from Sargasso Sea – Live in swamps and marshes – Produce methane as a waste product – Obligate anaerobes • Archaeans are also found in ‘normal’ environments Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ecology of Prokaryotes Chemical Recycling • Prokaryotes play crucial roles in the biosphere • Prokaryotes play a major role • Prokaryotes are so important to the biosphere that if they were to disappear – The prospects for any other life surviving would be dim • e.g. Photosynthesis – The loss of Eukaryotes would not have such an effect on Prokaryotes – In the continual recycling of chemical elements between the living and nonliving components of the environment in ecosystems • Many chemoheterotrophic prokaryotes function as decomposers – Breaking down corpses, dead vegetation, and waste products • Nitrogen-fixing prokaryotes – Add usable nitrogen to the environment Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 7 Symbiotic Relationships: Living together Symbiotic Relationships • Some prokaryotes • Mutualism: light-producing bacteria live in special organs of the flashlight fish – Live inside hosts as parasites or disease agents • Some prokaryotes – Live with mutualistic relationships with their hosts (both benefit): • Legumes (peas & beans) host nitrogenfixing bacteria in their roots – Others may be commensals (commensalism) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Symbiotic Relationships Figure 27.15 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Escherichia coli - some forms parasitic/pathogens • Prokaryotes have both harmful and beneficial impacts on humans – Some prokaryotes are human pathogens – But many others have positive interactions with humans Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Pathogenic Prokaryotes • Prokaryotes cause about half of all human diseases – Lyme disease is an example • Pathogenic prokaryotes typically cause disease – By releasing exotoxins or endotoxins • Many pathogenic bacteria – Are potential weapons of bioterrorism Figure 27.16 5 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 8 Prokaryotes in Research and Technology • Experiments using prokaryotes – Have led to important advances in DNA technology – Escherichia coli makes insulin for diabetics! • Prokaryotes are the principal agents in bioremediation – The use of organisms to remove pollutants from the environment Figure 27.17 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fig. 28-03a Excavata Diplomonads Parabasalids Euglenozoans • Prokaryotes are also major tools in Dinoflagellates Alveolates Apicomplexans Chromalveolata – Mining Ciliates Diatoms Stramenopiles – The synthesis of vitamins Golden algae Brown algae Oomycetes Chlorarachniophytes Forams Radiolarians Green algae Archaeplastida Red algae Chlorophytes Charophyceans Land plants Amoebozoans Slime molds Gymnamoebas Nucleariids Sperm Haploid (n) Diploid (2n) Key “Bud” Protonemata Animals 1 Sporangia release spores. Most fern species produce a single type of spore that gives rise to a bisexual gametophyte. Key Male gametophyte 2 The haploid protonemata produce “buds” that grow into gametophytes. Choanoflagellates • The life cycle of a fern Raindrop Spores develop into threadlike protonemata. Fungi Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The life cycle of a moss 1 Unikonta Entamoebas Opisthokonts Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Rhizaria – Production of antibiotics, hormones, and other products 4 A sperm swims through a film of moisture to an archegonium and fertilizes the egg. Antheridia 3 Most mosses have separate male and female gametophytes, with antheridia and archegonia, respectively. 2 The fern spore develops into a small, photosynthetic gametophyte. 3 Although this illustration shows an egg and sperm from the same gametophyte, a variety of mechanisms promote cross-fertilization between gametophytes. Haploid (n) Diploid (2n) Antheridium Spore MEIOSIS Young gametophyte “Bud” Sporangium Egg Spores Gametophore Female Archegonia Meiosis occurs and haploid spores develop in the sporangium gametophyte of the sporophyte. When the Rhizoid sporangium lid pops off, the peristome “teeth” regulate 6 The sporophyte grows a gradual release of the spores. long stalk, or seta, that emerges Seta from the archegonium. Archegonium 8 Peristome Sporangium MEIOSIS Mature Mature sporophytes sporophytes Capsule (sporangium) Calyptra Zygote Mature sporophyte 6 On the underside of the sporophyte‘s reproductive leaves are spots called sori. Each sorus is a cluster of sporangia. Embryo Female gametophytes Young 5 The diploid zygote sporophyte develops into a sporophyte embryo within 7 Attached by its foot, the the archegonium. sporophyte remains nutritionally dependent on the gametophyte. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings FERTILIZATION Sorus FERTILIZATION Archegonium Capsule with peristome (LM) Sperm Egg Zygote (within archegonium) Foot Figure 29.8 New sporophyte Sporangium Gametophyte Fiddlehead Figure 29.12 4 Fern sperm use flagella to swim from the antheridia to eggs in the archegonia. 5 A zygote develops into a new sporophyte, and the young plant grows out from an archegonium of its parent, the gametophyte. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 9