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Microbial Ecology Chapter 30 Principles of Microbial Ecology, Definitions  Ecology   Ecosystem   The study of relationships among organisms and their environment. Includes all of the biotic (living) components and the abiotic (physical and chemical) components of an environment. Biosphere  That region of the earth that is inhibited by living organisms. Principles of Microbial Ecology  Definitions  Biodiversity   Biomass   Evenness of distribution of the # of species present Weight of all organisms present Ecological Community  Comprised of a variety of different species in a given environment; more stable than an environment with fewer organisms. Principles of Microbial Ecology  Ecological Niche   The role that an organism plays in its particular ecosystem as well as the physical space it occupies. Microenvironment   Environment immediately surrounding an individual cell Biofilm (Fig. 30.1, pg. 766) Principles of Microbial Ecology  Indigenous   Native organisms Nonindigenous  Temporary inhabitants Principles of Microbial Ecology  Nutrient Acquisition  Primary Producers  Autotrophs      Convert CO2 organic material Photoautotrophs – plants, algae, cyanobacteria Anoxygenic phototrophs  Use sunlight for energy Chemolithoautotrophs  Oxidize inorganic compounds for energy Food source for consumers and decomposers Principles of Microbial Ecology  Consumers  Heterotrophs    Utilize organic material Food chain  Herbivores – primary consumers  Carnivores – secondary consumers  Carnivores – tertiary consumers Food web  Interacting food chains Principles of Microbial Ecology  Decomposers  Heterotrophs     Primarily bacteria and fungi Digest remains or primary producers and consumers  Detritus - Fresh or partially decomposed organic matter Specialize in digesting complex materials Mineralization  Complete breakdown of organic matter into inorganic molecules such as ammonia, sulfates, phosphates & CO2 Principles of Microbial Ecology  Low Nutrient Environments  Common in nature  Dilute aqueous solutions    Lakes, rivers, streams Distilled water reservoirs Respiratory equipment Principles of Microbial Ecology  Microbial Competition  Ability of microbes to compete successfully for a habitat generally related to   Rate at which organism multiples Ability to withstand adverse environmental conditions Principles of Microbial Ecology  Antagonism  Promotes biodiversity through competition  Bactericins  Proteins produced by some soil microbes that kill closely related strains of bacteria Principles of Microbial Ecology  Microbes and Environmental Change  Examples  Enzyme induction     Inactivates mercury Only formed when mercury is present Antibiotic resistant bacteria Growth and metabolism of organism can change environment.  Figure 30..4, pg. 768 Principles of Microbial Ecology  Microbial Communities  Biofilms (discussed in ch. 4)  Microbial Mat  A thick, dense, highly organized structure composed of distinctive layers (fig. 30.5, pg. 769) Principles of Microbial Ecology  Microbial Ecology Studies  Traditional    Cultures Microscopy Molecular Techniques  Microscopy   Dyes that made are fluorescent by metabolic activities Fluorescence in situ hybridization (FISH)  Nucleic acid probes to observe only cells with specific nucleotide sequences Principles of Microbial Ecology    Confocal scanning laser microscopes  To observe sectional views of a 3-dimensional specimen (biofilm) Polymerase chain reaction (PCR)  To detect only certain organisms  Denaturing gradient gel electrophoresis (DGGE)  PCR & DGGE studies conform that standard cultures techniques can be poor indicators of natural microbial population composition Genomics  Sequence information can apply to more than one group of microbes Aquatic Habitats  Water   Extremely efficient solvent Can absorb various wavelengths of light  Important aspect relating to photosynthesis Aquatic Habitats  Marine Environment  Oceans     Cover more than 70% of earth’s surface Most abundant aquatic habitat Represent 95% of global water Fresh Water Environment  Lakes, Rivers   Fraction of global water source Important source of fresh water Aquatic Habitats  Oceans and lakes  Characteristic zones influence distribution of microbial populations  Upper layers   Sufficient light penetration - photosynthetic microorganisms Oligotrophic waters   Nutrient poor Growth of photosynthetic organisms & autotrophs limited by lack of phosphate, nitrate and iron Aquatic Habitats  Eutrophic waters  Nutrient rich (fig. 30.6, pg. 770)    Photosynthetic activities in upper layers produce organic compounds Organic compounds permit growth of heterotrophs in lower layers Heterotrophs consume dissolved O2 during metabolism  O2 consumption can outpace slow rate of atmospheric O2 diffusion into water  Can create a hypoxic environment Aquatic Habitats  Definitions  Eutrophic  A body of water rich in nutrients   Oligotrophic   A body of water low in nutrients Eutrophication   swamps, bog lakes, etc. Natural nutrient enrichment of waters Accelerated Eutrophication  Rapid loading of nutrients Aquatic Habitats  Potable Water   Safe for drinking Rainwater  Distillate   Ground Water   contaminated by air pollutants aquifers, underground lakes & rivers Surface Waters  Creeks, rivers, ponds, lakes Aquatic Habitats  Factors Affecting Presence of Organisms  Nutrients  Oceans typically oligotrophic  Inshore areas not as stable as deep ocean  Dramatically affected by run-off  Dead zone in Gulf of Mexico every spring Aquatic Habitats  Oxygen (limiting factor)      low solubility in water, quantities limited well mixed cold water ~8-9mg/l warm water ~ 5mg/l Deep marine water is O2 saturated due to mixing associated with tides, currents and wind Temperature - Worldwide 0oC to ~100oC Aquatic Habitats  Freshwater environments  Oligotrophic lakes may have anaerobic layers due to thermal stratification  Epilimmion     Hypolimmion     Colder deeper layers (~5o-4oC) May be anaerobic due consumption of O2 by heterotrophs Water most dense at 4oC (39oF) Thermocline (~20o-10oC)   Warm upper layer (25o-22oC) Generally oxygen rich due to photosynthetic organisms Generally aerobic Zone (layer) of rapid temperature change As weather cools, water mixes oxygenating deep water Aquatic Habitats  Freshwater Environments  Rivers and Streams  Usually shallow and turbulent     Facilitates O2 circulation Generally aerobic Generally good sunlight penetration for photosynthesis Sheathed bacteria adhere to stable structures to allow utilization of nutrients flowing pass  Examples: Sphaerotilus & Leptothrix Aquatic Habitats  Factors Affecting Presence of Organisms  Sunlight Penetration (Photic Zone)     depth of sunlight penetration algae & cyanobacteria photosynthesis provides nutrients & oxygen for other organisms pH Range 2 - 9  fish hypersensitive to bacterial parasites at pH 5.5, usually die if pH drops below 4.5 Aquatic Habitats  Specialized Aquatic Environments  Salt lakes – no outlets    Iron springs   Water evaporates, concentrates salt Halophilic organisms Contain large quantities of ferrous ions Sulfur Springs  Support both photosynthetic and non-photosynthetic sulfur bacteria Aquatic Habitats  Lake Zones  Littoral Zone    Limnetic Zone   Extending from shore to the limit of occupancy of rooted plants Part of the photic zone Region of open water bounded by zone of emergent (rooted) vegetation Benthic Zone  Sediment (regardless of depth) Aquatic Habitats  Freshwater  Composition of the water reflects its source   Stagnant ponds to free flowing rivers and lakes Ground water   Surface water    Normally relatively free of nutrients and toxins Affected by surface runoff of materials Organics, fertilizers, herbicides, pesticides, etc. Inshore Marine  Affected by freshwater runoff and pollutants Aquatic Habitats  Marine Environment  Factors affecting presence of miroorganisms   Same Factors as Fresh Water plus Barometric pressure (hydrostatic pressure)     1 atm / 33 feet of seawater ocean 35,750 feet (11,000 meters) deep, hydrostatic pressure 1,083 atm Organisms are barophilic (barophiles) Salinity   Marine averages 3.5% (fresh averages ~0.5%) Organisms are halophilic (halophiles) or halotolerant Aquatic Habitats  Microbial Flora  Dictated by Available Nutrients  Bulk of Microbial Mass   Aerobic Chemoheterotrophic Bacteria    algae, cyanobacteria & protozoa degrade organic materials Cytophaga, Caulobacter Chemoautotrophic Bacteria  obtain energy from aerobic oxidation of reduced inorganic compounds Aquatic Habitats  Sulfur Oxidizers - Thiobacillus   Nitrifiers    oxidize H2S dissolved in water to inorganic sulfur or sulfate more important in marine environments oxidize ammonia - nitrite - nitrate Sediment  Methanogenic Bacteria  Foraminiferans & Radiolarians (oil and gas markers) Aquatic Habitats  Marine Waters  Microbial Flora  Most bacteria are found    Deep ocean vents    In association with organic particles (often less than 0.1mm in size) near the surface In association with skin or gut of fish Chemoautotrophic bacteria Some Vibrios are of major importance as fish pathogens Some microbes cause human-like diseases in fish   Pasteuralla piscicida (like tularemia) Mycobacterium marinum (TB like disease) Aquatic Habitats  Major Functions of Freshwater and Marine Bacteria  Decompose Organic Matter   Transform Essential Minerals   liberate mineral nutrients cycling them through forms other organisms can use Release Dissolved Organic Compounds  into the food web to support growth of other organisms Aquatic Habitats  Determining Microbial Flora  Epifluorescence Counting   Stain with acridine orange (stains DNA) view slide under UV light   tedious and can be inaccurate, counts DNA from living and dead organisms Luciferin-luciferase Enzyme System   Gives estimate of the number of viable organisms in a given volume of water Based on carbon:ATP ratio (~250 for most microbes) Terrestrial Habitats  Characteristics of Soil  Composed of   Pulverized rocks, decaying organic material, air & water Life   Bacteria, fungi, algae, protozoa, worms, insects, and plants roots May contain    More than 4,000 different species per gram of soil More than 2 tons of bacteria and fungi per acre Can be a rapidly and dramatically changing environment Terrestrial Habitats  Soil Layers (Horizons)  Topsoil (A Horizon)     Subsoil (B Horizon)   Accumulation of clays, salts & various nutrients C Horizon   Dark, nutrient-rich Supports plant growth Depth – few inches to several feet Partially weathered bedrock R Horizon  Unweathered bedrock Terrestrial Habitats  Microorganisms in Soil  Composition affected by environmental conditions  Moisture    Finely textured soils (clay) tend to be waterlogged and anaerobic Sandy soils (dry quickly) tend to be aerobic Acidity   Suppresses bacterial growth Fungi thrive with less competition for nutrients Terrestrial Habitats  Temperature    Mesophiles comprise the bulk of the soil bacteria, they grow best between 20oC and 50oC Thermophiles occur in compost piles where they generate heat Available Nutrients  The size of the microbial population in soil is limited by on the amount of organic matter available Terrestrial Habitats  Soil Organisms  Prokaryotes   Most numerous soil inhabitants Most common genera    Nocardia, Arthrobacter, Streptomyces Streptomyces  Produce conidia (dessication resistant spore)  Produce geosmins (give soil musty odor)  Produce many medically useful antibiotics Gram (+) bacteria more abundant than Gram (-) bacteria Terrestrial Habitats  Not all Soil Organisms are Beneficial  Human Bacterial Pathogens   Clostridium and Norcardia Human Fungal Pathogens  Coccidioides, Histoplasma, and Blastomyces Terrestrial Habitats  Fungi   Make up bulk of soil biomass Most are aerobic     Some are free-living Some occur in symbiotic relationship with plant roots   Usually found in top 10 cm of soil Crucial in decomposing plant matter Mycorrhizae Algae  Live mostly on or near surface Terrestrial Habitats  Algae    Dependent on sunlight and photosynthesis to provide energy needs. Sensitive to environmental conditions of drought and low temperature Major nutrient source for   Earthworms and nematodes Protozoa    Aerobic - generally found near the surface Found in moist soils at a density of ~104 to 105 organisms per gram of soil Predators of soil bacteria and algae Terrestrial Habitats  Rhizosphere   Zone of soil that adheres to plant roots Roots cells extract organic molecules   Sugars, amino acids and vitamins Fosters growth of microorganisms   Gram (-) more prevalent than surrounding soil Certain grasses – enriched with Azospirillum species Biochemical Cycling & Energy Flow  Biochemical Cycles   Cyclical paths elements take as they flow through living (biotic) and non-living (abiotic) components of ecosystem Fixed and limited amount of elements available  Carbon and nitrogen particularly important    Stable gaseous forms CO2 and N gas enter atmosphere Global impacts Elements continually cycle in ecosystem  Energy does not, must be continually added to fuel life Biochemical Cycling  Elements - three general purposes  Biomass production  Incorporated into cell material   All organisms require nitrogen to produce amino acids Energy source   Reduced form of element is used to generate energy – ATP Energy yielding reactions oxidize the energy source   Chemoorganotrophs use reduced carbon compounds – sugar, lipids and amino acids Chemolithotrophs use reduced inorganic molecules – H2S, ammonia (NH3) and hydrogen gas (H4) Biochemical Cycling  Terminal electron acceptor  Electrons from energy source transferred to an oxidized form of element during respiration    Aerobic conditions  O2 is terminal electron acceptor Anaerobic conditions some prokaryotes use  Nitrate (NO3), nitrite (NO2), sulfate (SO4)and CO2 The following pages will review cycling processes for oxygen, carbon, nitrogen, phosphorus and sulfur Oxygen Cycle  During photosynthesis cyanobacteria, algae and green plants produce oxygen from water. The oxygen is utilized via respiration.  The level of oxygen in the atmosphere is maintained by chemical reactions in the upper atmosphere, aerobic respiration and photosynthesis Carbon Cycle  Carbon    Carbon enters producers during photosynthesis or chemosynthesis In turn enters consumers via consumption of the producers. Carbon returned to the atmosphere in the form of CO2 by respiration and the actions of decomposers consuming dead or decaying waste.  Oxygen has profound influence on cycle      Allows degradation of certain compounds Helps determine the types of carbon containing gases produced Aerobic decomposition Great deal of OC2 formed through aerobic respiration (CH2O)n + (O2)n CO2 + H2O Carbon Cycle  Low oxygen (wet soils, marshes, swamps, etc.)  Degradation is incomplete  Generate CO2 and other gases  Some CO2 used by methanogens (ex: Archaea) as terminal electron acceptor generating methane (CH4)  4H2 + CO2 CH4 + H2O  Methane entering atmosphere is oxidized by UV light and chemical ions to CO and CO2 Nitrogen Cycle  Nitrogen (Fig. 30.11; pg. 775)   Most important constituent of proteins and nucleic acids Consumers obtain required nitrogen from ingested plants and animals and use it to build biomass  Prokaryotes – diverse in use of nitrogen compounds    Some use oxidized compounds like nitrate and nitrite Some use reduced nitrogen compounds like ammonium All of these metabolic activities represent steps in the N cycle Nitrogen Cycle  Nitrogen Fixation  Nitrogen gas reduced to form ammonium    Ammonium can be incorporated into cellular material Atmosphere 79% N2  Relatively few organisms use atmospheric (gaseous) nitrogen – rely on prokaryotes to convert atmospheric nitrogen into a useable form Nitrogenase  Enzyme complex that mediates nitrogen fixation and is readily inactivated by oxygen  Nitrogen fixing aerobes must have a mechanism for protection Nitrogen Cycle  Nitrogen fixing prokaryotes (diazotrophs)  Free living     Symbiotic - significant in benefiting plant growth    Azotobacter - chief suppliers of fixed nitrogen in grasslands & similar ecosystems Cyanobacteria - most significant nitrogen fixer in aquatic environments Clostridium spp. - dominant free-living anaerobes in soils Found in association with all leguminous plants including alfalfa, clover, peas, beans, peanuts and vetch Rhizobium Synthetic nitrogen compounds Nitrogen Cycle  Ammonification  The decomposition of organic nitrogen into ammonia    Occurs when extracellular proteolytic enzymes convert proteins into amino acids. Other enzymes then decompose amino acids into ammonium (NH4+) and sulfate ions. Ammonium ions in turn can be oxidized to nitrite (NO2-) and nitrate (NO3-) through Nitrification Nitrogen Cycle  Nitrification   Oxidation of ammonium to nitrite Nitrifiers - encompass two groups of chemolithotrophic bacteria     Ammonia oxidizers  Nitrosomonas - (NH4+ to NO2-) (ammonium to nitrite) Nitrite oxidizers  Nitrobacter & Nitrospira (NO2- to NO3-) (nitrite to nitrate) Obligate aerobes – use molecular O2 as final electron acceptor  Nitrification does not occur in waterlogged soils or anaerobic aquatic environments Important because it supplies plants with nitrates which is the most useable form of nitrogen for plant metabolism Nitrogen Cycle  Denitification   Process to convert nitrate to gaseous nitrogen Nitrate represents fully oxidized nitrogen      Pseudomonas spp. can use nitrate as terminal electron acceptor  Anaerobic respiration Nitrate reduced to gaseous nitrogen compounds – nitrous oxide and molecular nitrogen Release to atmosphere represents a loss of nitrogen and contributes to global warming Benefit – waste water treatment processes to remove nitrate Anammox   Brocadia anamoxidans oxidizes ammonium anaerobically Potential benefit in waste water treatment Sulfur Cycle  Sulfur    Occurs in all living things Chiefly a compound of amino acids methioine & cysteine Key steps of cycle rely on prokaryotes     Some use reduced form of H2S, some elemental S Others use sulfate Most plants and microbes assimulate sulfur as sulfate (SO42-) Is present in the soil (like nitrogen) chiefly as a part of proteins Sulfur Cycle  Hydrogen sulfide is toxic to lining things    Under aerobic conditions, H2S oxidizes spontaneously to sulfur and is then converted to sulfate (SO42-) (its most readily utilized form) by sulfur bacteria Under anaerobic conditions sulfate-reducing bacteria reduce sulfate to hydrogen sulfide Oxidation of hydrogen sulfide to sulfate carried out principally by nonphotosynthetic autotrophs, Thiobacillus, Thiothrix and Beggiatoa and less commonly by photosynthetic autotrophs (green and purple sulfur bacteria) Sulfur Cycle  Sulfur Reduction  Reduction of sulfate to sulfide   Carried out by anaerobic bacteria that are capable of utilizing sulfate as the final electron acceptor in their anaerobic respiration Include Desulfovibrio and Desulfomonas Phosphorus Cycle  Involves movement of phosphorus between inorganic and organic forms  Microorganisms play three major roles in phosphorus transformations    Mineralize organic phosphorus Convert insoluble forms of inorganic phosphorus to soluble forms Immobilize inorganic phosphorus Phosphorus Cycle  Overall Transformations of Phosphorus  Soil Organisms   Break down organic phosphate into to inorganic phosphates Then convert inorganic phosphates to orthophosphate (PO43-)    Orthophosphate is water soluble and readily used by most plants and microorganisms When plants & animals die decomposers convert organic phosphate back into inorganic phosphate Phosphorus is often the limiting nutrient in many environments
 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                            