| Microbiology, 5/e Lansing M Prescott,
Augustana College Donald A Klein,
Colorado State University John P Harley,
Eastern Kentucky University
Microorganisms in Terrestrial Environments
Study Outline- Soil as an Environment for Microorganisms
- Soil consists primarily of inorganic geological materials, which are modified by the biotic community
- Soils are dynamic and develop over time
- Changes in the activity of the microbial community caused by changes in plant growth, temperature, and other factors can significantly alter soil
- In most soils, the major producers of organic material are vascular plants; algae, cyanobacteria and photosynthetic bacteria also contribute
- Bacteria, fungi, protozoa, insects, nematodes, worms and many other animals live in soil
- Soil varies in terms of the amount of oxygen available to microorganisms
- Highly oxygenated areas are found on particle surfaces; microorganisms are often found in thin films of water on these surfaces
- The interior of a soil particle can be anaerobic
- Under certain conditions, soil may contain isolated pockets of water, which serve as mini-aquatic environments; oxygen concentrations are generally lower in these pockets of water
- Water logged soil is very similar to anaerobic lake sediments
- Changes in water content, gas fluxes, and the growth of plant roots can affect the concentration of carbon dioxide and other gases in the soil
- Microorganisms in the Soil Environment
- Bacteria are found primarily on the surfaces of soil particles, most frequently on surfaces of pores in these particles; this probably protects them from predation by protozoa and gives them access to soluble nutrients
- Filamentous fungi form bridges between separated particles or aggregates called peds; this exposes the fungi to high levels of oxygen; filamentous fungi move nutrients and water over great distances in the soil
- Protozoa, soil insects, and other animals are located in soil and often feed on bacteria and fungi
- Microbial populations can be very high; yet only a small portion of them have been cultured; actinobacteria are important, but less studied, members of terrestrial soil ecosystems
- Microorganisms are constantly being added to soil, but most do not survive
- The microbial community makes important contributions to biogeochemical cycling
- Soil microorganisms can be categorized on the basis of their preference for easily available or more resistant substrates
- Some prefer higher nutrient levels and they respond rapidly to the addition of easily utilizable substrates such as sugars and amino acids
- Others are indigenous forms that tend to utilize native organic matter
- Some grow in oligotrophic (low nutrient) environments
- Soil insects and other animals serve as decomposer-reducers; they not only decompose but also physically reduce the size of organic aggregates; this increases the surface area and makes nutrients more available for utilization by bacteria and fungi
- Protozoa influence nutrient cycling by microbivory (feeding on microorganisms), causing an increase in the rates of nitrogen and phosphorus mineralization
- Free hydrolytic enzymes released from animals, plants, insects, and microorganisms contribute to soil biochemistry
- Microbial loop-rapid turn over of organic matter making the nutrients available to plants rather than to higher trophic levels; bacteria and predatory protozoa are involved in this process
- Microorganisms and the Formation of Different Soils
- Soil formation
- Colonization takes place after newly exposed geological material begins to weather
- Nutrients such as phosphorus and iron may be present, but carbon and nitrogen must be imported; cyanobacteria are active in the pioneer stage of nutrient accumulation
- Once formed, most soils are rich sources of nutrients because of the plant and animal life that lives and dies there
- Tropical and temperate region soils
- In tropical soils, organic matter is decomposed rapidly and mobile inorganic nutrients can be leached out causing rapid loss of fertility; fertility can be further decreased by slash-and-burn agriculture
- In temperate region soils, decomposition rates are less than the rate of primary production
- In temperate grasslands, deep root penetration leads to formation of fertile soils
- In cooler coniferous environments, there can be excessive accumulations of organic matter; fire is a major means to control nutrient cycles
- In bog soils, decomposition is slowed by waterlogged, anoxic conditions; this leads to peat accumulation
- Cold moist area soils-colder temperatures decrease both the rate of decomposition and the rate of plant growth; below the plant growth zone is permafrost; these soils are very sensitive to physical disturbances and pollution
- Desert soils-microbial communities called desert crusts, which consist of cyanobacteria and associated commensals, can retain rainfall
- Geologically heated hyperthermal soils-populated by bacteria and archaea, many of which are chemolithoautotrophs; chemoorganotrophs are also present
- Soil Microorganism Associations with Vascular Plants
- Microorganisms on the outside of plants
- Phyllosphere microorganisms-a wide variety of microorganisms are found on and in the aerial surfaces of plants (phyllosphere), where they can utilize organic compounds released by the leaves and stems; phyllosphere microorganisms can either protect or harm the plant host
- Rhizosphere and rhizoplane microorganisms
- The rhizosphere is the volume of soil around plant roots influenced by materials released by the plants; the rhizoplane is the root surface; both provide unique environments for microorganisms; microorganisms in the rhizosphere serve as a labile source of nutrients and play a critical role in organic matter synthesis and degradation
- Numerous rhizosphere bacteria influence plant growth through the release of auxins, gibberellins, cytokinins, and other molecules
- Associative nitrogen fixation-nitrogen fixation carried out by bacteria on the rhizoplanes and in the rhizosphere
- Microorganism growth within plants
- Rhizobium-a prominent member of the rhizosphere community; it can also establish an endosymbiotic association with legumes and fix nitrogen for use by the plant
- The infection process is under the control of a bacterial gene and a plant regulatory protein
- Rhizobium is stimulated to produce Nod factors, which activate the host responses necessary for root hair infection and module development
- The bacterium induces formation of an infection thread by the plant; the infection thread grows down the root hair
- Rhizobium spreads within the infection thread into the underlying root cells
- Bacteria multiply and develop into swollen, branched bacteroids enclosed by a plant-derived membrane called the peribacteroid membrane
- Further growth and differentiation lead to the formation of nitrogen-fixing forms called symbiosomes
- Nitrogen fixation occurs within symbiosomes within the root nodules, and the nitrogen is then assimilated into various organic compounds and distributed throughout the plant
- Once differentiated, the bacterial cells cannot reproduce; this sacrifice has been called altruism in the rhizosphere: it may promote root exudation and the maintenance of high levels of Rhizobium in the rhizosphere
- A major goal of biotechnology is to introduce nitrogen-fixing genes into plants that do not normally form such associations
- Fungal and bacterial endophytes of plants-some fungi and bacteria infect and live within plants as endophytes; can be mutualistic or parasitic (pathogenic)
- Mycorrhizae
- Fungus-root associations
- Five associations have been described; they fall into two broad categories
- ) Ectomycorrhizae-grow as an external sheath around the root with limited penetration of the fungus into the cortical regions of the root; found primarily in temperate regions; mycelia extend far into the soil forming a mycorrhizosphere and mediate nutrient transfer to the plant; process is aided by mycorrhizal helper bacteria (MRB)
- ) Endomycorrhizae-fungi that penetrate the outer cortical cells of the plant root, where they form characteristic structures known as arbuscules
- Mycorrhizae increase the competitiveness of the plant and increase water uptake by the plant in arid environments; they also make it possible to share resources (e.g., carbon, minerals and water)
- Actinorrhizae-actinomycete-root associations between members of the genus Frankia and woody plants; Frankia can fix nitrogen and are important in the life of woody, shrublike plants
- Stem-nodulating rhizobia-bacteria that form nodules at the base of adventitious roots branching out of the stem part above the soil surface; observed primarily in tropical legumes
- Agrobacterium-members of this genus containing the Ti (tumor-inducing) plasmid cause the formation of galls (tumors) on the plant; is a complex process that involves transfer of Ti DNA into the plant host; recent interest in these bacteria centers on the use of the Ti plasmid as a vector to transfer new genetic characteristics (eg., herbicide resistance, bioluminescence) to plants
- Fungi and bacteria as plant pathogens-many fungi and bacteria are plant pathogens, causing rusts, blights, rots and other plant diseases
- Viruses-many viruses infect plants and cause disease and economic losses (e.g., TMV)
- Tripartite and tetrapartite associations-involve some combination of plant, rhizobia, mycorrhizae, and actinorrhizae; enable plant to better cope with nutrient-deficient environments
- Soils, Plants, and Nutrients
- Organic matter in soil helps retain nutrients, maintain soil structure, and hold water for plant use; plowing, irrigation, and other soil disturbances can increase microbial degradation of this organic matter, thereby decreasing soil fertility
- Several agricultural methods have been developed to promote and maintain soil fertility
- No-till and minimum-till practices reduce aeration and use herbicides to control weed infestation
- Composting allows for controlled decomposition and produces physiologically stabilized compost material; this material, when added to the soil, increases the organic content without stimulating soil microorganisms to increase decomposition
- Soils are increasingly being impacted by mineral nitrogen releases resulting from human activities
- This nitrogen is derived from two major sources: agricultural fertilizers and fossil fuel combustion
- The nitrogen releases have had a number of effects
- When the nitrogen added is greater than can be used by plants, it remains in mobile form and can enter surface water and groundwater; higher nitrate levels in water are related to a number of animal health problems
- Nitrogen fertilizers alter microbial community structure and function
- Excessive use of fertilizers can have global-level impacts when nitrogen enters coastal and marine waters
- Phosphorus fertilizers cause eutrophication of surface waters when excess phosphorus moves into those waters
- Soils, Plants, and the Atmosphere
- Ice-nucleating bacteria serve as nucleation centers for snow and precipitation formation; genetically engineered ice-minus bacteria can be sprayed on frost-sensitive plants to protect the plants from frost-damage; however, the release of such genetically engineered microorganisms into the environment is of continuing concern
- Soil microorganisms can influence the atmosphere by degrading airborne pollutants such as methane, hydrogen, and carbons dioxide; they can also improve air in closed buildings
- Soil microorganisms can influence the global fluxes of both relatively stable and reactive gases
- Relatively stable gases include carbon dioxide, nitrous oxide, nitric oxide, and methane
- Reactive gases include ammonia, hydrogen sulfide, and dimethylsulfide
- Use of nitrogen-containing fertilizers, use of automobiles, conversion of soils to agricultural use, and use of landfills can stimulate production of NO, N20, methane and other greenhouse gases
- Methane is produced by methanogens found in ruminants, rice paddies, termite guts, and landfills
- Methane is degraded by methanotrophs
- The balance between methane production and methane utilization is important for preventing excessive accumulation of this greenhouse gas
- Microorganisms and Plant Decomposition
- Released soluble materials from plants create a residuesphere, an area between decaying plant material and the soil; residuesphere has much microbial activity
- Soluble plant materials are degraded first
- Starch, cellulose, and proteins are then degraded, both aerobically and anaerobically
- Lignin is degraded primarily by aerobic basidiomycetes, but only after physical rot leads to cavitation of wood
- Unusual gases are produced in the process of woody plant decomposition (e.g., chloromethanes, and cyanide)
- The Subsurface Biosphere
- Studied by examining outcrops, surface excavations, petroleum hydrocarbons, well corings, and materials from deep mine sites; the use of improved techniques has led to the acceptance of the idea that microorganisms exist far below Earth's surface
- Microbial processes take place in different regions
- Shallow subsurface where water flowing from the surface moves below the plant root zone
- Subsurface regions where organic matter has been transformed by chemical and biological processes to yield coal, kerogens, and oil and gas; mobile materials move up into more porous geological structures where microorganisms can be active
- Zones where methane is being synthesized
- Soil Microorganisms and Human Health
- Soils contain a wide variety of pathogenic organisms, which can cause disease if they have an entry point into the human body and find favorable conditions there (e.g., Clostridium, Acanthamoeba, and Cyclospora)
- The growth of soil-related microorganisms in buildings can cause mold problems and "sick building" syndrome
- Understanding Microbial Diversity in the Soil
- It is estimated that only 1 to 10% of microscopically observable organisms in soils can be cultured in the laboratory
- Molecular techniques are now making it possible to assess the range of microbial diversity in soils
- However, to fully understand the physiology of these microorganisms, scientists will ultimately need to culture them
|
|