Chapter 2 The Ecosystem Ecosystem: General Model Ecosystem: Internal Dynamics S = Storage A = Autotrophs H = Heterotrophs Ecosystem Trophic Structure • Autotrophic Stratum – ‘Green’ upper level (chlorophyll containing plants) – Fixation of light energy; use of simple inorganic substances complex organic substances • Heterotrophic Stratum – ‘Brown’ lower lever (soil, sediment, decaying matter, etc.) – Use, rearrangement, and decomposition of complex materials Ecosystem Components 1) Inorganic substances – C, N, CO2, H2O, and others involved in material cycles 2) Organic Compounds – proteins, carbohydrates, lipids, humic substances, and others that link the biotic and abiotic components 3) Physical Environment – air, water, substrate, climate regime, other physical factors Ecosystem Components 4) Producers – (autotroph) organisms, usually green plants, that manufacture food from simple inorganic substances 5) Heterotrophs – (phagotroph) organisms that ingest other organisms or particulate organic matter 6) Decomposers – (saprotroph) mostly bacteria and fungi that obtain energy by breaking down dead tissue or absorbing dissolved organic matter 1) Saprophages – organisms that feed on dead organic material; important for nutrient cycles Designations are not species specific. A species may be intermediate between heterotroph and decomposer. Consider decomposition as a process involving several organisms. Organic Detritus • Detritus – all the organic matter involved in the decomposition of dead organisms – Important link between abiotic and biotic components • Products of decomposition – POM – particulate organic matter – DOM – dissolved organic matter – VOM – volatile organic matter Ecosystem Function • Interaction between the autotrophic and heterotrophic components Simple Material (C,H,O) Autotrophs Heterotrophs Complex Material (Carbohydrate) Most vital elements are in a constant state of flux. Others, like ATP, are never found outside of the cell. Humic substances never found inside of a living cell. Gradients and Ecotones • The biosphere is characterized by a series of gradients (zonation) • Temperature: – Equator to the poles, mountaintop to valley. • Moisture: – Wet to dry along major weather systems • Depth: – Shore to bottom in aquatic environments Vegetation based zones Horizintal and vertical zones Metabolic zonation Temperature stratification zonation Ecosystem Boundaries? • It is easy to picture ecosystems as having distinct boundaries. • The area of transition from one ecosystem to another is considered to be an ecotone. • Ecotones have a mixture of species from both ecosystems. – A marsh between a freshwater lake and dry land. – Zone of grasses, shrubs, and scattered small trees between forests and grasslands. Where does one ecosystem end and the other begin? Two examples of ecotones. Species Overlap in Ecotones Land zone Transition zone Number of species Species in land zone Species in aquatic zone Species in transition zone only Aquatic zone Edge Effect • Higher species diversity found on the edge of an ecosystem (ecotone) than in the interior – Marsh and open water (shrimp, crabs, juvenile finfish) • Edge species – those species that are concentrated in ecotones • Sharp edge usually a poor habitat – Clear cut – forest edge Old Field versus Pond • Two systems compared to understand ecosystem structure. • Majority of inorganic and organic compounds are in storage. – New Hampshire Forest nitrogen: 90% in soil organic matter, 9.5% in biomass, 0.5% available in soluble form. • Rate of ecosystem function is controlled by: – Rate of nutrient release from solids, solar input, temperature change, day length, other climate conditions Primary Producers • Pond: macrophytes and phytoplankton – Macrophytes can be important in some cases – Phytoplankton important in oceans • Old Field (Grassland): macrophytes – Rooted plants dominate – Algae, mosses, lichens can be present Consumer Organisms • Herbivores Primary consumers – Pond: zooplankton (animal plankton) or benthos (bottom forms) – Field: Small insects, large hooved animals • Carnivores Secondary and tertiary consumers – Pond: predaceous insects and small fish (nekton) – Field: predatory insects, spiders, birds, mammals Detritivores and Decomposers: • Found throughout ecosystems, but mostly at the mud-water or soil-leaf litter interface. – Nongreen bacteria, fungi, flagellates • Decomposition increases with temperature – Cellulose, lignin, and humus impart a spongy texture to soil Food Webs (chain) and Energy Flow Heat First Trophic Level Second Trophic Level Third Trophic Level Fourth Trophic Level Producers (plants) Primary consumers (herbivores) Secondary consumers (carnivores) Tertiary consumers (top carnivores) Heat Heat Heat Solar energy Heat Heat Detritvores (decomposers and detritus feeders) Heat Community Metabolism • Production:Respiration (P/R ratio) • If > 1, then excess biomass is being produced • If < 1, then more biomass is being consumed than produced • If = 1, then compensation point Watershed Concept • Ponds and grasslands are actually open systems • Watershed = catchment basin • Often, the entire drainage basin must be considered as the unit of management You will be required to draw a map of the major rivers of the Mississippi River Basin as part of exam 1. Ecosystem Diversity • We can look at genetic diversity, species diversity, habitat diversity, and diversity of functional properties • Two components of interest: – The richness (total # of species) – Relative abundance of each species (evenness) Diversity Indices • A mathematical measure of species diversity in a community. • Reveals important information regarding rarity and commonness of species in a community. Shannon-Wiener Diversity Index (H) H = - pi(lnpi) Larger H = more diversity • Variables associated with the ShannonWeiner Diversity index: S – total number of species in the community (richness) pi – proportion of S made up of the ith species Hmax = ln(S) EH – equitability (evenness; b/t 0 and 1) = H / Hmax Species 1 2 3 4 1.386294 # 12 562 8 1 583 pi 0.020583 0.963979 0.013722 0.001715 ln(pi ) (pi )(lnpi ) -3.88328 -0.07993 -0.03669 -0.03536 -4.28875 -0.05885 -6.36819 -0.01092 -0.18507 H= 0.18507 E = 0.18507/1.386297 = 0.1335 # 12 12 12 12 48 pi 0.25 0.25 0.25 0.25 ln(pi 2) -1.38629 -1.38629 -1.38629 -1.38629 -0.34657 Species Richness = 4 -0.34657 ln (species richness) = Hmax = 1.386294 -0.34657 H = - (sum of (pi)(lnpi)) = 1.38629 -0.34657 E = H/Hmax 0.999997 -1.38629 # pi 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 12 0.05 Species Richness = 4 12 0.05 ln (species richness) = Hmax = 1.386294 12 0.05 H = - (sum of (pi)(lnpi)) = 0.69881 12 0.05 E = H/Hmax 0.504085 240 Species Richness = 20 ln (species richness) = Hmax = 2.995732 H = - (sum of (pi)(lnpi)) = 2.99573 E = H/Hmax 0.999999 Species richness and equitability affect the Shannon Wiener index. # 3 38 1 6 48 pi ln(pi 2) 0.0625 -2.77259 0.791667 -0.23361 0.020833 -3.8712 0.125 -2.07944 -0.17329 -0.18495 -0.08065 -0.25993 -0.69881 Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 ln(pi 2) -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -2.99573 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -0.14979 -2.99573 29 species, fairly evenly distributed 11 species, dominated by 1 species Relative Abundance Global Production and Decomposition • Approximately 1017 grams of organic matter produced by photosynthesis annually – Approximate equivalent oxidized back to CO2 and H2O (but not exactly). • Since Precambian, a small fraction of photosynthetic material was incompletely decomposed and sequestered fossil fuels – Led to decrease in atmospheric CO2 and increase in O2. – Release of the sequestered CO2 has led to increased atmospheric levels Production Within an Ecosystem • Allochthonous input – organic material transferred into the ecosystem from an outside source • Autochthonous input – organic material produced within the ecosystem Photosynthesis: Solar energy converted to chemical energy CO2 converted to Carbohydrate Solar energy + 6CO2 + 6H2O → C6H12O6 + 6O2 Happy Rays of Sunshine You need to know this CO2 O2 (from air) (to air) C6H12O6 H2O Radiant Energy • Photosynthesis converts solar energy into the chemical energy of a carbohydrate by two sets of reactions: • Solar energy + 6CO2 + 6H2O → C6H12O6 + 6O2 Reduced Oxidized Electrons from H2O are energized by the sun. Carbohydrate (glucose) Oxidation-Reduction • Oxidation is the loss of electrons (energy) and reduction is the gain of electrons (energy). • In covalent rxn’s, oxidation also refers to the loss of hydrogen atoms, and reduction refers to the gain of hydrogen atoms. Bacterial Photosynthesis CO2 + 2H2A + light energy (CH2O) + 2A A could be Sulfur (2H2S 4H + 2S; not oxygen) or an organic compound. Photosynthetic bacteria generally play a minor role in the production of organic matter, but are important in nutrient cycling. Photosynthetic Bacteria • Photosynthetic bacteria that release oxygen are largely aquatic cyanobacteria. • Obligate anaerobes – function only in the absence of oxygen (green and purple sulfur bacteria). – Occur between the reduced and oxidized boundary layer in sediments or water where the light intensity is low. – Important for sulfur cycle • Facultative anaerobes – able to function with or without oxygen. – Generally are non-sulfur photosynthetic bacteria Photosynthesis Overview • Composed of light-dependent and lightindependent reactions • Light-dependent reactions – Capture solar energy and excite electrons – Water molecule is split and electrons and H+ enter the electron transport system – O2, NADPH, and ATP are produced • Light-independent reactions – CO2 is reduced to a carbohydrate – NADPH and ATP are consumed Light-dependent Reactions Solar energy is used excite electrons (increases potential energy). ADP and NADP+ are reduced to ATP and NADPH. Water is split H+, e-, and O2 *Considered an electron donor* ATP and NADPH are then used to power the lightindependent reactions. Light-independent Reactions • Calvin Cycle – three stages – CO2 fixation, CO2 reduction, RuBP regeneration – Reactions require energy, which is supplied by ATP and NADPH Light-independent Reactions -Calvin Cycle Fixation of CO2 From lightdependant reactions From lightdependant reactions C3, C4, and CAM plants • Carbon fixation so far has been described as C3. – Initial carbon fixation and Kreb’s cycle occur at the same time in the same place. – Rubisco oxidizes RuBP in the presence of a high oxygen concentration – High rates of photosynthesis also lead to high rates of photorespiration • C4 and CAM plants have adapted the photosynthesis process to reduce photorespiration Rubisco is the enzyme that carboxylates RuBP with CO2. However, in the presence of high O2, it will oxidize RuBp and release a CO2. Because Carbon is fixed as a 3 carbon molecule, this is called C3 photosynthesis. This represents a loss of CO2 that was already ‘fixed’ – this is called photorespiration. In C4 plants, bundle sheath cells also contain chloroplasts, and mesophyll cells are arranged concentrically around bundle sheath cells. Oxygen is produced in the Mesophyll cells, so it does not accumulate in the bundle sheath cells when the stomata are closed. High concentration of CO2 in the bundle-sheath cell Reduces photorespiration Partition by space Partition by time Decomposition (respiration) • Type 1. Aerobic respiration – gaseous oxygen is the electron acceptor (oxidant) • Type 2. Anaerobic respiration – gaseous oxygen is not the electron acceptor • Type 3. Fermentation – anaerobic, but the organic compound oxidized is also the electron acceptor Aerobic Respiration C6H12O6 + 6O2 6CO2 + 6H2O + energy (ATP) Glycolysis and Kreb’s Cycle lead to complete breakdown of carbohydrate to CO2 and H2O Overview of Glycolysis Glucose (6-C sugar) 2 ATP 2 ADP 6-C sugar diphosphate 3-C sugar-phosphate 2 ADP 3-C sugar-phosphate 2 ADP 2 ATP 2 NAD+ 2 ATP 2 NAD+ 2 NADH 2 NADH 3-C pyruvate 3-C pyruvate The NAD+ cycle Remember: When NAD+ NADH it has been reduced. Remember: When NADH NAD+ it has been oxidized. Pyruvate (Oxygen present) Cellular Respiration (Oxygen not present) Fermentation Pyruvate oxidation: if oxygen is present 2 NAD+ 2Pyruvate + 2CoA 2 NADH + H+ 2 Acetyl-CoA + 2CO2 Pyruvate is converted to a C2 acetyl group attached to coenzyme A (CoA), and CO2 is released. This occurs in the cytoplasm if oxygen is present. Acetyl-CoA (2 C) C6 C4 NAD+ NADH CO2 NADH NAD+ Krebs cycle C5 FADH2 NAD+ NADH FADH C4 CO2 ATP ADP +P Oxygen receives energyspent electrons at the end of the electron transport system then combines with hydrogen to form water: ½ O2 + 2 e - + 2 H+ → H 2O Glycolysis NAD+ Transition Electron Reaction Transport NADH Chain Krebs Cycle Remember: Electrons = Energy Anaerobic Respiration • Usually saprophages (bacteria, yeasts, molds, and protozoa) – Can occur in some muscle tissue • Methane bacteria – obligate anaerobes; produce methane by decomposing organic matter – Marsh gas • Desulfovibrio – important sulfur reducing bacteria that reduce SO4 and produce H2S. Chapter 2 Continued…………. Decomposition • The breakdown of large molecules to it’s basic components – Abiotic (forest fires) and biotic process – Organic material is an energy source for decomposer organisms • Decomposition is physical and chemical – Leaf shredders particulate organic matter • Increase surface area – Bacteria and fungi use enzymes to break apart large molecules • Left over nutrients are reabsorbed by primary producers Decomposition Rate • Composition of organic material – For example: Lignin vs. protein, lipid, carbohydrates • Presence of macroinvertebrates such as shredders Decomposition Rate • Can be affected by temperature and water – Remember: enzymes work faster at high temperatures Decomposition Rate • Depends on a variety of organisms – Bacteria, fungi, insects, nematodes Ecosystem Function • A combination of production, respiration, and decomposition • What are the anthropogenic impacts on ecosystem function? • Ecological Footprint – a measure of the anthropogenic effect on the environment Ecological Footprint • • Ecological footprint – amount of land needed to produce the resources needed by the average person in a country Methods: 1. Correct consumption data for trade imports and exports Consumptionwheat= production + imports – exports 2. Convert to land area needed to produce the item Awheat = Cwheat / ywheat A=total area needed, C=consumed, Y=yield 3. Obtain per capita ecological footprint by dividing by population size fwheat = awheat/population size Ecological footprint in relation to available ecological capacity. It would take about 3 times the current land area of Earth if all 6.1 billion people consumed the same as the 276 million people in the US Per Captia Ecological Footprint (Hectares of land per person) Country 10.9 United States 5.9 The Netherlands India Country 1.0 Total Ecological Footprint (Hectares) 3 billion hectares United States The Netherlands India 94 million hectares 1 billion hectares