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Module 9 - 13
Enzymes
Proteins that catalyze biochemical reactions by lowering activation energy and increasing the speed of chemical reactions in cells. It binds to specfic substrates in active site of proteins
Catalase
A common enzyme that breaks down peroxide into water & oxygen - it protects organisms from oxidative damage 2H2O2 → O2 + 2H20
Peroxidases - Intracellular or Extracellular
oxidative stress response and lignin degradation (blank cell walls)
Chitinases - Intracellular or Extracellular?
degradation of chitin polymers (fungal cell walls)
Cellulases - Intracellular or Extracellular?
degradation of cellulose polymers (plant cell walls)
Amylases - Intracellular or Extracellular?
degradation of starch polymers (C cycling)
Proteases - Intracellular or Extracellular?
degradation of protein (C and N cycling)
Phosphates - Intracellular or Extracellular?
removes inorganic P from organic substrates
What does enzyme activity in soil indicate?
change in soil structure and chemistry
Cellular Functions of Extracellular Enzymes:
solubilize/breakdown large polymers
detoxify the environment
reduce/oxidize insoluble minerals/metals
What traits make soil fungi so effective at secreting extracellular enzymes?
Fungi are filamentous, can create a large surface area to search for food and secretes enzymes
Why do fungi act as good extracellular enzymes?
They are primary degraders of plant cell wall material using hydrolases, oxidases, laccases, peroxidases, and phenol oxidases. They also perform phosphorus mineralization, which is when the fungi release enzymes to collect phosphate groups from organic compounds which they then turn into something useful for plants to grow
Methods to measure general microbial activity and specific enzyme activity in soils - know limitations of each method
Soil Respiration - measures the rate of carbon dioxide released from the soil as microbes break down organic matter. Limitations: cannot distinguish the difference between microbial respiration and root respiration; heavily fluctuates with immediate changes in soil moisture & temperature rather than remaining stable
Dehydrogenase Activity Assays - dehydrogenase acts as an intracellular marker of intact, viable oxidative metabolism. Limitations: do not function outside a cell, highly sensitive to air drying and storage
Microbial Ecology
The study of interactions of microorganisms with each other and their environment
Population
All the organisms belonging to a single species with the potential to interact. The spatial scale of a population depends upon the size, mobility, and reach of the species
Community
All the organisms living in one habitat; includes both populations or subpopulations of various species
Community Structure
The numbers and types of organisms present
Neutralism
A type of interaction where organisms occupy the same habitat but are neutral to each other, neither organism benefits nor is harmed by the actions of the other organism in the community
Commensalism
Describes an interaction where organisms occupy the same habitat but at no cost or benefit to another species - very common
Examples of Commensalism
Oxygen utilization by aerobes in surface areas can benefit obligate anaerobes (symbiotic relationship) - obligate anaerobes gain a critical survival advantage, given access to new resources, habitats, & protection against oxygen toxicity (+) whereas aerobes are completely unaffected (0)
Degradation of complex organic matter - specialized microbes secrete extracellular enzymes that break down complex polymers to produce byproducts that are simple nutrients to the surrounding environment (+). In contrast, neighboring microbes that lack the degrading gene will take the free simplified nutrients (0)
Production and release of metabolic byproduct - the first organism creates and releases byproducts naturally (0), while the consumer uses the waste products as a primary source of nutrients, energy, or environmental modification; think of it as a free meal (+)
Toxin Inactivation/Detoxification - one organism breaks down environmental toxins/ metabolic waste products (0); the second organism now has a safe, habitable environment (+)
Auxotroph
An organism that has a nutrient deficit and is unable to synthesize one or more vitamins, growth factors, amino acids, nucleotides, etc. Organisms must obtain the missing compound from its environment.
Prototroph
An organism that can make all intermediates and end-products necessary for growth and metabolism from the basic building blocks
Mutualism
Interactions that benefit both organisms involved
What are 2 types of Mutualism and examples?
Proto-cooperation: a temporary mutualistic association where both partners benefit
ex. cross-feeding nutrients: species A produces an amino acid that species B takes up and uses. Species B produces biotin, which is taken up and used by species A. Both partners benefit when needed, but a species can easily be replaced with another that does the same function
Symbiotic mutualism: an obligatory, non-transitory, mutually beneficial interaction
ex. Mycorrhizal fungi and plant roots: The plant performs photosynthesis and supplies fungi with Carbon, energy-rich sugars, and nitrogen. While the fungi provide a large network that extends far beyond the root, increasing the surface area for plants to absorb water, phosphorus, nitrogen, and other essential minerals
Competition
rivalry over a limiting factor or resource
Interspecies Competition
complex interactions with neighboring organisms from a different species; includes many trophic levels
Intraspecies Competition
competition between parents and progeny in the same species - leads to variation in phenotype and abilities within a population
Carrying Capactiy
the maximum population of a particular species that a given habitat can support over a given period of time
Common Competitive Strategies
High growth rate
Efficient use of nutrients
Metabolic diversity - switches to other metabolisms/pathways with less competition
Motility - leaves one habitat to colonize a new one
Amensalism
Oligotrophs
Organisms that are able to sense and acquire resources that are scarce and highly dilute (low in concentration) and use them more efficiently; slower to metabolize & grow, but can grow at nutrient concentrations that would not be accessible to other organisms
Copiotrophs
Organisms that have high nutrient requirements both to sense, acquire, and utilize those nutrients; usually grow faster when nutrients are abundant; however do not grow well when nutrients are scarce
Amensalism
a type of antagonistic interaction between organisms - one organism is harmed/destroyed while the other is completely unaffected (no benefit)
ex. pH change, antibiotic production, siderophore (iron cage)
What are biotic and abiotic factors that influence the soil bacteria communities?
Soil pH, Organic carbon quality & quantity, Soil (O2) and redox status
Predators vs. Parasites
Predators = feed on prey, actively hunts/kills them to survive
Parasites = live on/inside a host organism for an extended period of time, feeding off its resources without killing it immediately
Scramble Competition
Deprive another of a resource by depleting that resource first
ex. 2 different species of microbes colonize a new microhabitat in soil, and both use the same energy source, which is initially abundant. Organism A has a slower growth rate than Organism B
Contest Competition
actively harms another competitor by “germ warfare” - production of toxins and antimicrobials
Competitive Exclusion Principle
2 species competing for the same exact limited resources cannot stably coexist at a constant population sizes in the same ecological niche
Mycorrhizae
A symbiotic association between plant roots and fungi - in most terrestrial plant and all woody plants, relatively non selective
What are the important roles of mycorrhizae?
Nutrient acquisition
Water acquisition
Sequestration of heavy metals
Defense against plant pathogens
Arbuscular Mycorrhizae (AM)
A very common type of mycorrhizal association, mostly herbaceous and some trees. Limited range in taxa but not in the plant.
Describe arbuscular mycorrhiazae and its location.
It is the place of contact between the plant and fungi. The fungal hyphae penetrates epidermis of root and cortex of plant cell walls, produces spores within or outside the root. *The arbuscules are the site of nutrient exchange
Benefits of arbuscular mycorrhizae.
enhances uptake of nutrients to plant (inorganic phosphorus and some nitrogen)
improves plant & and health
can encourage plant diversity under stressful conditions
exhibits limited extracellular enzymatic activity - depends upon plant & soil bacteria for carbon, which minimizes metabolic energy waste
Vesicular-Arbuscular Mycorrhizae
Similar to arbuscular mycorrhizae, but they also form vesicles. The vesicles almost completely fill host plant cortical cells and contain lipids and phosphorus (provides storage for both Carbon and Phosporus)
Ectomycorrhizae
Relatively few in plant species; more dominant in trees and some shrubs. They do not penetrate host cells; instead, they wedge between them. They form an extensive hyphal network (Hartig net) and a sheath on the surface or root. Spores produced in specialized structures away from the root (ex. mushrooms, truffles) *Hartig Net = site of nutrient exchange. It also produces more extracellular enzymes
Ectomycorrhizae extracellular enzymatic activity.
phosphatase - liberate phosphorus from organic matter
Proteinase - liberates nitrogen & carbon (some sulfur) from proteins
Benefits to the fungal partner.
fungi are heterotrophs, the plant provides a ready source of carbon, energy, and growth factors
improves fungal competitiveness with bacteria for shared resources so no competition for food because the plants provide carbon
fungi are heterotrophs (cannot produce their own food); mycorrhizae can demand 20-40% total fixed carbon produced by the plant
Benefits to the plant partner.
increases root contact with nutrient pool - AM mainly supply of inorganic P to plants, ECM provide P & N to plant through proteinase & phosphatase
ECM important in seedlings to help with tree growth
promotes plant-to-plant communication & resource sharing
increased tolerance to stress
protection from plant pathogens
Rhizosphere
the portion of soil under the direct influence of the roots - it is extremely diverse and microbial colonization is patchy along the root
Rhizoplane
the physical surface of the plant root
How does organic carbon affect the rhizosphere?
causes changes in the physical, chemical, and biological nature of soil
can account for about 50% of rhizosphere respiration
high & low molecular weight compounds
High molecular weight organics
the majority of carbon released from roots
complex organic matter ex. mucliage & cellulose
turnover time: days
Mucliage
insoluble high molecular weight polysaccharide-rich material secreted from the root cap and border cells - a large amount of carbon that goes out is found here
Low molecular weight organics
highly diverse suite of molecules
turnover time: minutes-hours
soluble and gaseous compounds like O2
not easily moved, mostly stays at root - in order to secrete the plant might use a gas to transport
Functions & impact of rhizosphere microbiome on plant
Good
growth & development, nutrient acquisition, tolerance to abiotic stress, protection against pathogens, physiology/metabolism, immune response
Bad
plant diseases & food contamination
Beneficial Microbe-Plant interactions
Beneficial rhizobacteria provide nutrients - plants need N,P and trace metals (iron): nitrogen fixation, degradation & mineralization of organic matter, production of bacterial siderophores
Rhizobacteria support plants under abiotic stresses: flooding, salinity, cool temperatures, low pH, soil contaminants/toxins
Provide the first line of defense against plant pathogens: competition for limited resources (food, nutrients, space), parasitism (some beneficial bacteria parsitize pathogens), amensalism
Prime plant for systemic resistance response: induce plants to activate cellular defenses to mitigate pathogen attack, establish disease-suppressive soils, induce plant cell wall reinforcements, enhance production of plant defense metabolites
Replication
capture natural variance: distinguish “true” biological signals from background soil heterogeneity
avoid pseudo-replication: samples should come from separate, independent experimental units
budget constraints: need to balance fine-scale detail with statical power