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Institute of Food and Agricultural Sciences (IFAS) . Biogeochemistry of Wetlands Science and Applications. Carbon Cycling Processes. Wetland Biogeochemistry Laboratory Soil and Water Science Department University of Florida. Instructor K. Ramesh Reddy krr@ufl.edu. 11/17/2014. 1.
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Institute of Food and Agricultural Sciences (IFAS) Biogeochemistry of Wetlands Science and Applications Carbon Cycling Processes Wetland Biogeochemistry Laboratory Soil and Water Science Department University of Florida Instructor K. Ramesh Reddy krr@ufl.edu 11/17/2014 1 WBL
Institute of Food and Agricultural Sciences (IFAS) Carbon Cycling Processes CO2 OM CH4 WBL
Carbon Cycling Processes • Lecture Outline • Introduction • Major components of carbon cycle • Organic matter accumulation • Characteristics of organic matter • Decomposition processes • Regulators of organic matter decomposition • Greenhouse gases • Summary WBL
Carbon Cycling Processes • Learning Objectives • Describe major components of carbon cycle • Develop an understanding of the chemical composition of plant litter and soil organic matter • Long-term accumulation of organic matter • Describe the role of enzymes and microbial communities involved in decomposition • Determine organic matter turnover • Indentify the role biogeochemical controls and regulators • Understand the global significance of carbon cycle • Draw a carbon cycle and identify storages and fluxes within and between soil and water column WBL
Oxidation States of Carbon [+4] [0] CO2 C6H12O6 [-4] CH4 WBL
Carbon Reservoirs[1014 kg] • Atmospheric CO2 7 • Biomass 4.8 • Fresh water 2.5 • Marine 5-8 • Soil organic matter 30-50 WBL
Soil Organic Matter [SOM] • Undecayed plant and animal tissues • Partially decomposed material • Soil biomass • Sources of SOM • External: Particulate (inputs) • Internal: detrital material (macrophytes, algal mats, roots) WBL
Detrital Plant Biomass Aerobic Detritus Decomposition Water table Peat Burial Anaerobic Compaction Grazers CO2 microorganisms WBL
Carbon Cycle UV CO2 CO2 CH4 Decomposition/leaching Decomposition/leaching Litter Microbial biomass HCO3- DOC Export Import Peat Microbial biomass HCO3- CH4 DOC Decomposition leaching Decomposition/leaching WBL
Storages Soil organic matter Plant detritus/litter Dissolved organic matter Microbial biomass Transformations Microbial respiration Methanogenesis Outputs Greenhouse gases Nutrient export Ecological/Environmental Significance Carbon sequestration Global warming Water quality Ecosystem productivity Organic Matter WBL
Net Primary Productivity[g/m2 - year] [Craft, 2001] Bog 380-800 Marsh 500 -1100 Riverine 400-1150 Fresh tidal 500-1600 Brackish 600-1600 Salt 950-2000 Mangroves 600-1200 WBL
Carbon Accumulation in Wetlands[g C/m2 year] Alaska - Sphagnum 11-61 Finland - Sphagnum - Carex 20-28 Ontario - Sphagnum bog 30-32 Georgia - Taxodium 23 Florida - Cladium 70-105 WBL
Organic Matter Accumulation 0 Organic matter accumulation 10 Soil Depth [cm] 1964 marker 20 Cs-137 Activity WBL
A. Detritus attached to plant B. Detritus detached from plant C. Decomposed detritus from previous year Water detritus D. Organic matter and nutrient accretion Soil Soil Organic Matter Plant Detritus A B C Decay continuum WBL
Decay Continuum Live plant CO2 CH4 Plant standing dead Litter layer Microbial decomposers Surface peat Buried peat WBL
Carbon Accumulation in Wetlands • Potential energy source (reduced carbon, electron donor • Long-term storage of nutrients, heavy metals, and toxic organic compounds • Major component of global carbon cycles WBL
Carbon Forms • Particulate organic carbon (POC) • Microbial biomass carbon (MBC) • Dissolved organic carbon (DOC) • Dissolved inorganic carbon (DIC) • CO2 + H2O = H2CO3 • H2CO3 = HCO3- + H+ • HCO3- = CO32- + H+ WBL
Chemical constituents of organic matter • Non Humic compounds: • Carbohydrates (Simple sugars) • Monosaccharides: glucose. • Polysaccharides: Starch, Cellulose, and Hemicellulose • Proteins • Lipids etc • Phenolic compounds: • Lignin (branched random polymer of phenyl propanoid unit) • Tannins (heterogeneous groups of phenolic compounds) WBL
Organic Matter (Plant and Soil) • Water soluble components [<10%] • Sugars, amino acids and fatty acids • Cellulose [15-60%] • Hemicellulose [10-30%] • Lignin [5-30%] • Proteins [2-15%] • Lipids and Waxes [1-8%] • Ash (mineral) [1-13%] WBL
Cellulose b-D-glucosidic bond OH H CH2OH CH2OH O O H H H O H H H H H OH H OH OH O H H H O OH OH CH2OH H H WBL
Lignin WBL
Soil Organic Matter [SOM] SOM Extract with Alkali [alkali-soluble] Humin [alkali-insoluble] Treat with Acid Humic Acid [acid-insoluble] Fulvic Acid [acid-soluble] WBL
Fulvic Acid • More ‘O’ and less ‘C’. • MW 1000 -30,000. • Less advanced stage of decomposition. • More COOH group per unit mass. • Functional group acidity (11.2 mol/kg). • Alkali and acid soluble. WBL
Humic Acid • More ‘C’ and less ‘O’. • MW 10,000 -100,000. • Advanced stage of decomposition. • Less COOH group per unit mass. • Functional group acidity (6.7 mol/kg). • Alkali soluble. WBL
Available Carbon Pool • Represents small but biologically active fraction of DOC • Immediately available for microbial utilization • Extremely small in C-limited system • Rapid turnover • May not be directly measurable • Affects short-term community metabolism WBL
70% water Macromolecules 15% protein 3% polysaccharide 2% lipids 5% RNA 1 % DNA 1 % Inorganic ions 3 % others Total weight of actively growing cell of Escherichia coli Wet wt = 9.5 x 10-13 g Dry wt = 2.8 x 10-13 g Microorganisms[Percent wet weight] WBL
Microbial Decomposers • Typically 1-5% of total C mass in soil • Process most of the ecosystem net production • Principal transformers of organic carbon • Recycle carbon and nutrients in recalcitrant biopolymers • Regulate energy flow and nutrient retention WBL
Techniques to Measure MICROBIAL BIOMASS Direct cell count : abundance Lipid based : live microbial biomass CHCl3 Fumigation-extraction based: estimate of Carbon Metabolic activity based: Enzyme activities WBL
MICROBIAL COMMUNITY STRUCTURE • Pure culture approach • Microscopy • Community level physiological profile (CLPP): Substrate utilization: BIOLOG • Measurement of cellular component (physiological status, functional groups):PLFA • Methods based on nucleic acids analysis (abundance, diversity and phylogeny of organisms): gene specific analysis (16S rDNA, DGGE, TGGE, Trflp) WBL
MICROBIAL BIOMASS [Site = WCA-2A - Everglades] 10 9 8 7 LITTER 6 % of Total Carbon 0-10 cm 5 4 3 10-30 cm 2 1 0 0 2 4 6 8 10 Distance from Inflow, km WBL
MICROBIAL NUMBERS [MPN/g soil] [Site = WCA-2A - Everglades] Substrate Eutrophic Oligotrophic Lactate 9.3 x 105 9.2 x 103 Acetate 2.3 x 105 3.6 x 103 Propionate 4.3 x 105 9.2 x 103 Butyrate 4.3 x 105 < 3.0 x 103 Formate 2.3 x 105 < 3.0 x 103 Hector et al. 2003 WBL
DetritalMatter Leaching Complex Polymers Cellulose; Hemicellulose; Lignin Proteins; Lipids and waxes Extracellular Enzyme (Hydrolysis) End product Monomers Sugars;Amino acids Fatty acids Electron acceptors Bacterial Cell End products + energy WBL
Periplasmic space Bacterial cell Detrital/clay material Extracellular Enzymes • An extracellular enzyme is involved in transformation or degradation of polymeric substances external to cell membrane. • Enzyme can be bound to the cell membrane or are periplasmic (ectoenzyme)(Chrost,1990) • Enzyme occurs free in the water or adsorbed to surface other than its producers e.g., detrital particles or clay material (extracellular enzyme) • Most of these are hydrolases WBL
Enzymes • Cellulose degradation • Exocellulase - Cellulose • B-glucosidase - Cellobiose • Hemicellulose degradation • Exoxylanase - Xylan • B-xylosidase - Xylobiose • Lignin degradation • Phenol oxidase - Lignin and Phenols WBL
Enzyme – Catalyzed Reaction E S E + P E + S S = Substrate E = Enzyme P = Product All enzymes are proteins – amino acid polymers WBL
Reactions of Enzymes R-O-PO32- + H2O R-OH + HO-PO32- alkaline phosphatase R-O-SO3- + H2O R-OH + H+ + SO42- arylsulfatase R-O-glucose + H2O R-OH + glucose b-glucosidase casein + H2O tyrosine protease phenolics + O2 quinones phenol oxidase WBL
Ca2+ Ca2+ Ca2+ Ca2+ Ca2+ Ca2+ Ca2+ Ca2+ Inhibition of enzyme activity Humic acid-Enzyme complex Active Enzyme Humic acid E E + Ca2+ + E + E WBL
P APase MUF-P P MUF Pi Measurement of Enzymes • Spectroscopic • p-nitrophenol phosphate (pNPP) • Fluorescence • Methylumbelliferyl phosphate (MUF) • Enzyme Labeled Fluorescence (ELF) WBL
b Glucosidase Activity 100 ug p-nitrophenol g-1 h-1 50 0 Oxygen Nitrate Sulfate Bicarbonate E h (mV) 618 214 -145 -217 pH 4.5 7.6 7.5 6.5 WBL
impacted transitional unimpacted b Glucosidase Activity [Everglades -WCA-2A] 4 February 2 0 4 May B-D-Glucosidase Activity (mg p- nitrophenol g-1 h-1) 2 0 4 August 2 0 Detritus 0-10 cm 10-30 cm WBL Wright and Reddy, 2001
impacted transitional unimpacted Phenooxidase Activity [Everglades -WCA-2A] Wright and Reddy, 2001 5 4 May 3 2 1 0 Phenol Oxidase Activity (umole [DQC]g-1 min-1) 5 August 4 3 2 1 0 Detritus 0-10 cm 10-30 cm DQC = dihydroindole quinone carboxylate WBL
DetritalMatter Leaching Complex Polymers Cellulose; Hemicellulose; Lignin Proteins; Lipids and waxes Extracellular Enzyme (Hydrolysis) Reduced product Monomers Sugars;Amino acids Fatty acids Electron acceptors Bacterial Cell End products + energy WBL
Decreasing energy yield Organic Matter Decomposition SOIL DEPTH WBL
Metabolism • Catabolism • Anabolism • Types of energy source • Light … Phototrophs • Inorganic … Lithotrophs • Organic …. Heterotrophs • Oxidation of organic compounds • Fermentation • Respiration WBL
Chemolithotrophy • Inorganic compound as energy source • eg. H2S, Hydrogen gas, Fe(II), and NH3 • Source of carbon for biosynthesis cannot be organic therefore use CO2 and hence are autotrophs • Hydrogen oxidation • Sulfur oxidation • Ferrous iron oxidation • Annamox • Nitrification WBL
Carbon CO2 H2O ADP hu 1/2O2 ATP (CH2O)n Phototrophy • Photosynthesis is conversion of light energy into chemical energy. • Most phototrophs are autotrophs ( use CO2 as sole Carbon source). OXYGENIC PHOTOTROPHS ANOXYGENIC PHOTOTROPHS Carbon CO2 H2S ADP S0 hu SO42- ATP (CH2O)n WBL
Catabolism Energy sources: Organic, inorganic, light Waste products: Organic, inorganic Cell biomass Nutrients: N, P, K, S, Fe, Mg, ... Carbon sources: Organic, CO2 Anabolism Metabolism WBL