• Wetland Ecology • Lectures 14-15-16 • Wetland Biogeochemistry • What is biogeochemical cycling? – • • • Transport & Transformation of chemicals in an ecosystem, involving numerous interrelated physical, chemical, & biological processes Examples of movement include – Water-sediment exchange – Plant Uptake – Organic exports Two major categories of wetland biogeochemistry include: – Intrasystem cycling through various transformation processes – Exchange of chemicals between a wetland & its surroundings Open vs Closed Ecosystems OPEN • BHF + Tidal Marshes have significant exchange of materials with surroundings – River flooding + Tidal Exchange CLOSED • Bogs + Cypress Domes – Little material exchange except for gaseous matter! • Biogeochemically open vs closed • BGC Open – Abundant exchange of materials with surroundings – • BGC Closed – Little movement of materials across the ecosystem boundary – • Tidally driven or riparian Cypress swamps or very stagnant areas So, are wetlands open or closed systems? – They can be both! • Wetland Soil • Wetland soil is: – Medium in which many of the wetland chemical transformations occur – The primary storage of available chemicals for wetland plants • • Two major types of wetland soils – Organic & Organic Mineral Soils – Mineral Organic Soil + Organic Mineral Soils Defined under two saturation conditions • • 1. Saturated with water for long periods – Have > 18% organic carbon – Have > 12% organic carbon if no clay present – Have a proportional content of organic carbon (12 – 18%) if clay content (0 – 60%) 2. Soils are never saturated with water for more than a few days and have > 20% organic carbon – %Corg = %OM/2 – %Corg = percentage of organic carbon – %OM = percentage of organic matter • Bulk density & porosity – have lower bulk densities (dry weight of soil/volume). Low due to high porosity (peat soils ~ 80% air space) • Hydraulic conductivity – Depends on degree of decomposition. Organic soils hold more water (do not necessarily allow more water to pass) • Nutrient availability – Organic soils have more minerals tied up in organic form. • Cation Exchange Capacity – Organic soils have greater CEC (sum of exchangeable ions) • Organic soils (What order?) are classified into four groups; 3 are hydric – Saprists (muck) – > 2/3 of the material is decomposed, < 1/3 of plant fibers are identifiable – Fibrists (peat) - < 1/3 of material is decomposed and > 2/3 of plant fibers are identifiable – Hemists (mucky peat or peaty muck) – Conditions fall between saprist & fibrist soil – Folists – organic soils caused by excessive moisture (precip > evapotranspiration) that accumulate in tropical & boreal mountains; not classified as hydric because saturated conditions are the exception rather than the rule • Mineral Wetland Soil • When flooded for extended periods mineral soils develop certain characteristics that allow for their identification – Redoximorphic features (mediated by microbes) – The rate these features are formed depend on three conditions (all must be present) • Sustained anaerobic conditions • • • • Sufficient soil temp (5°C “biological zero”) • Organic Matter (substrate for microbial activity) Hydric mineral soils are characterized by: – Gleying – Oxidized Rhizosphere – Mottles (aka Redox Concentrations) – Current nomenclature 1. Redox concentrations – Accumulation of Fe & Mn in 3 different structures – Nodules & Concretions (firm extremely firm irregularly shaped bodies with diffuse boundaries) – Masses – formerly called ‘reddish mottles’ – Pore linings – Formerly included ‘oxidized rhizospheres’ 2. Redox depletions – Low-chroma (<2) bodies with high values (>4) including: – Iron depletions: “Gray mottles” or “Gley mottles” – Clay depletions: Contain less Fe, Mn, and clay than adjacent soils • 3. Reduced matrices: Low-chroma soils • REDOX in Wetlands • When mineral or organic soils are flooded anaerobic conditions result. Water fills pore spaces and rate of oxygen diffusion through soil is drastically reduced • Rate of oxygen depletion depends on: – Ambient Temperature – Availability of organic substrates for microbial respiration – Chemical oxygen demand from reductants such as ferrous Fe • Resultant O2 deficiency prevents plants from normal aerobic root respiration and affect nutrient availability and adds toxic materials in the soil • Usually thin layer formed and is related to: – Rate of O2 transport across the atmosphere-surface water interface – Small population of O2 consuming organisms – Photosynthetic O2 production by algae within water column – Surface mixing by convection currents & wind action • Eh ranges • If DO is present, the redox potential range +400 to + 700mV • If O2 disappears, Eh range from +400 down to -400mV • As organic substrates in a water logged soil are oxidized (donating) the redox potential drops = a sequence of reductions (gains) takes place • 1 reaction to occur after becoming anaerobic is the reduction of NO3- (nitrate) first to NO2(nitrite) and ultimately to N2O or N2 st – Nitrate becomes an acceptor ~ 250mV – At 225mV Mn is transformed from manganic to manganous – At -75 to -150mV Fe is transformed from ferric to ferrous; while sulfates are reduced to sulfides • pH • Soil and overlying waters of wetlands occur over a wide range of pH’s – Organic soils – often more acidic – Mineral soils – often neutral or alkaline • Alkaline soils previously drained decrease in pH because of buildup of CO 2 then carbonic acid • Acid soils previously drained increase in pH because of reduction of ferric iron hydroxide • Nitrogen Transformations • Nitrogen is often the “most limiting nutrient in flooded soils” – Limitations reported in salt marshes, freshwater inland marshes, & freshwater tidal marshes • Involve complex microbial processes • NH4 is the primary form of mineralized N in wetlands • Mineralization • Often referred to as ammonification NH2CONH2 + H2O 2NH3 + CO2 NH3 + H2O NH4 + OH - • Nitrification • Once ammonia has formed, it can take several possible pathways – Aerobic environment – ammonium is oxidized (nitrification) in two steps by Nitrosomonas sp. + - + 2NH4 + 3O2 2NO2 + 2H2O + 4H + energy – - By Nitrobacter sp. - 2NO2 + O2 2NO3 + energy • Denitrification • Denitrification is carried out by facultative bacteria under anaerobic conditions – nitrate is the terminal electron acceptor C6H12O6 + 4NO3 6CO2 + 6H2O + 2N2 • Most significant path of nitrogen loss from wetlands • Usually lost as N2 & N2O • N fixation • Conversion of N2 gas to organic nitrogen – Favored by low oxygen concentrations – Rhizobium species – Cyanobacteria (blue-green algae) are also common in Louisiana, northern bogs, & in rice cultures – Fe & Mn Transformations • Found in reduced forms in wetlands • Readily available Toxic levels • More soluble – Ferrous Fe reduced form of Fe – Ferric Fe oxidized form of Fe • Oxidized form creates barrier; may prevent plant from uptaking other nutrients • Reduced form reacts with P making it unavailable • Sulfur • Rarely limiting to plant or animal growth in wetlands • – Hydrogen sulfide (H2S) is toxic (rotten-egg smell) – Ferrous sulfide is responsible for black color of wetland soils (highly reduced sediment) Negative effects of sulfides on higher plants are attributable to a number of causes – Direct toxicity of free sulfide as it comes into contact with plant roots – Reduced availability of sulfur for plant growth because of its precipitation with trace metals – Immobilization of zinc & copper by sulfide precipitation • Carbon Cycle • Photosynthesis & aerobic respiration dominate the aerobic horizons • Fermentation • Methanogenesis • Occurs when certain bacteria (methanogens) use CO2 as an electron acceptor for the production of gaseous methane (CH4) • Requires extremely reduced conditions – Redox potential range from -250 to -350mV • Phosphorus • One of the most important chemicals in wetland systems • Most limiting in northern bogs, freshwater marshes, & southern deepwater swamps • Retention is one of the most important features for natural & constructed wetlands • Principle inorganic form = orthophosphate 3- – PO4 – H2PO4 (pH 2-7) – HPO4 (pH 8-12) (pH > 13) - — • Predominant form dependent on pH • P is not directly altered by Eh changes – • Affected by association with other elements especially Fe P is rendered relatively unavailable to plants & microconsumers by: – 1. Precipitation of insoluble phosphates with ferric Fe, Ca, & Al under aerobic conditions – 2. Adsorption (particle surface) of phosphate onto clay particles, organic peat and ferric & aluminum hydroxides & oxides – 3. Binding of phosphorous in OM as a result of its incorporation into the living biomass of bacteria, algae, & vascular macrophytes • Chemical Transport into Wetlands • Geologic inputs – from weathering of rock – Type of soil direct reflection of parent material • Biologic inputs – Photosynthetic uptake of carbon, nitrogen fixation, & biotic transport of materials by animals (birds) • Hydrologic inputs – Major inputs into wetlands • Precipitation • – Burning of fossil fuels – Increased [ ]’s of sulfates & nitrates in atmosphere Streams, Rivers, & Groundwater • – As precip reaches the ground it will: – Infiltrate into the ground – Return to atmosphere via evapotranspiration – Flow on surface as runoff Groundwater influence: – • • Chemical characteristics of streams & rivers depend on the degree to which the water has previously come into contact with underground formations & types of minerals present in those formations Climate: – Balance of precipitation & evapotranspiration – Type of vegetation present Geographic effects: amount of dissolved & suspended materials that enter streams, rivers, & wetlands depend on: – Size of watershed – Steepness or slop of landscape – Soil texture • Streamflow/Ecosystem effects: The water quality of surface water runoff, streams, & rivers varies seasonally • Human effects: water that has been modified by humans through sewage effluent, urbanization, & runoff from farms alters the chemical composition of streamflow & groundwater that enters wetlands • Drainage from agriculture fields: – • Higher [ ]’s of sediments, nutrients, herbicides, & pesticides might be expected Drainage from urban & suburban areas: – May have high [ ]’s of trace organics, oxygen demanding substances, & some toxic materials • Estuaries • Quality differs from that of rivers • Seawater chemical composition is fairly constant worldwide – 33 0/00 to 37 0/00 – Mass balances • A quantitative account of the inputs, outputs, & internal cycling of materials in an ecosystem • Mass balances help determine: • – Ecosystem functions – Determine the importance of wetlands as sources, sinks, and transformers of chemicals Inputs primarily through: – – • • Hydrologic • Precipitation • Surface & groundwater inflow • Tidal Exchange Biotic • Atmospheric carbon fixation (photosynthesis) • Atmospheric nitrogen (nitrogen fixation) Exports: – Surface water & groundwater – Long-term burial of chemical in the sediments Intrasystem cycling involves exchanges among various pools, or standing stocks of chemicals in within a wetland. Involves pathways such as: – Litter production – Remineralization – Chemical transformations – Translocation of nutrients through plants • Generalizations • 1. Wetlands serve as sources, sinks, or transformers of chemicals, depending on the wetland type, hydrologic condition, & length of time the wetland has been subjected to chemical loadings • 2. Seasonal patterns of nutrient uptake & release – • Temperate regions retention is higher in growing season • Increased microbial activity • Higher macrophyte activity 3. Wetlands are frequently coupled to adjacent ecosystems through chemical exchanges that affect both systems – Downstream ecosystems benefit from retention or from exportation • 4. Wetlands are either highly productive (eutrophic) or low productivity (oligotrophic) • 5. Nutrient cycling in wetlands differs from aquatic & terrestrial systems – More nutrients in sediment & peat – Aquatic systems have autotrophic activity more dependent on nutrients in water column than in sediments. – Wetland plants obtain nutrients from sediment • 6. Anthropogenic changes have led to changes in nutrient cycling in many wetlands • The capacity of wetlands to assimilate anthropogenic wastes from the atmosphere or hydrosphere is not unlimited!
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