Samenvatting Ecohydrology Afkortingen: GPP: Gross Primary Productivity. GPP is the rate at which ecosystems use energy (from photosynthesis), either for respiration or the creation of biomass. If you subtract the respiration from the whole, you get the NPP. NPP: Net Primary Productivity. NPP is the rate at which biomass is created TER: Total Ecosystem Respiration NEP: Net Ecosystem Productivity NEE: Net Ecosystem Exchange (Net CO2 gas exchange). NEE = GPP - TER LAI: Leaf Area Index = m 2 leaf m 2 soil WUE: Water Use Efficiency = mmol O2 fixed / mol H2O lost LUE: Light Use Efficiency NUE: Nutrient Use Efficiency ET: Evapotranspiratie Week 36: Hydro-ecology: The landscape ecological study of ecosystems dependent on groundwater and surface water (Wassen 1994). (Ecologie centraal) Eco-hydrology: The interdisciplinary study of groundwater hydrology as a component of ecosystems and as a determining factor for the pattern, distribution and development of vegetation (Pedroli1994). (Hydrologie centraal) Ecohydrology started in the Netherlands in the 1970s. Photosynthesis: 6CO2 + 6H2O + energy → C6H12O6 + 602 Produces sugars and O2 Solar energy needed Radiation energy in visible spectrum absorbed by pigments Takes place in green leaves (chloroplasts) Nitrogen: Important in photosynthesis: linear correlation between N in leaves (in Rubisco, chlorophyll) and photosynthesis rates. Photosynthetic Pathways: C3 Rubisco in leaves (Less maintainance) C4 Rubisco in bundle sheath cell (More CO2 efficient) CAM o Separated in time: CO2 exchange at night, photosynthesis during day o Stomata stay close during the day o Very water efficient, but slow growth o Mainly succulents (cacti) Respiration: C6H12O6 + 602 → 6CO2 + 6H2O + energy o Autotrophic respiration: ‘burns’ organic matter (sugars) to provide energy for plant functions (for example transport across membranes) Heterotrophic respiration: The conversion of organic matter to CO2 by organisms other than plants (decomposers, animals). Is dependent on: - Temperature (increase respiration exponentially to max, then decrease to zero (enzymatic activity interrupted)). - Moisture - Level of oxygen (low in dry conditions, increases to maximum at intermediate moisture levels, decrease when moisture content excludes oxygen). - Nutrient content (Poor quality more difficult to convert) Water & Plants: 70%-90% of fresh weight of non-woody biomass is water! Involved in chemical reactions (photosynthesis, respiration) Solvent Turgidity (water in vacuoles within protoplasts): Hydrostatic pressure in plant cells. Transport medium Water Movement Bulk flow: water flows from high to low potential Diffusion: from a high to a low concentration Osmosis: water flow through a semi-permeable membrane from a solution with a high water potential (low solute concentration) to a solution with a low water potential (high solute concentration) Water flows from roots to leaves because of the very large negative potential in the canopy due to transpiration. Soil Definitions: Saturation: Maximum water drainage Field Capacity: The maximum possible amount of water remaining in the soil after excess water has drained away. Water cannot drain freely. Water can be taken up by plants Permanent wilting point: Water cannot drain freely. Water cannot be taken up by plants due to the turgor limit of plants. It is defined as the minimal point of soil moisture the plant requires not to wilt. Water Use Efficiency (WUE) WUE = mmol CO2 fixed / mol H2O lost Water Use Efficiency: CAM > C4 > C3 Low stomatal conductance (depressed ET) → high WUE Week 37 Biogeochemistry: Interactions between chemistry of water, soil and biosphere. Hydrochemistry • Three main types of water • Distinct chemical composition: • Amount of dissolved ions (EC) • Relative amounts of Ca and Cl Rainwater chemistry • Fractionation factor: use of chloride as a stable marker X Cl rain FCl X Cl seawater • Calculate contributions of sources other than seawater Adsorption • Adherence of ions to the surface of soil particles • Based on electrochemical load (soils often negative load) • Adsorption capacity (CEC) depends on total surface area of soil particles soils with small particles (e.g. clay) and/or high organic matter content (e.g. peat) have high adsorption capacity • Also dependent on chemical composition of the soil particle (load!) Ion exchange • Exchange of adsorbed or absorbed ions and ions in groundwater • Specific sequence of adsorption based on electrochemical charge: Al3+ H+ Ca2+ Mg2+ K+ NH4+ Na+ Main differences aerobic-anaerobic C-cycle • Rates of processes (fast-slow) • End products (CO2 – CH4) • Soil organisms (aerobic – anaerobic; fungi - bacteria) • Mechanisms (OM decomposition – ‘chemical’ oxidation) • Influence of water quality Aerobic N-transformation Nitrogen mineralization: NH2·CO·NH2 + H2O 2NH3 + CO2 NH3 + H20 NH4+ Nitrification (actually ammonium oxidation): NH4+ + O2 NO2- + 4H+ + 2e- (by Nitrosomonas spp.) NO2-+H2O NO3- + 2H+ + 2e- (by Nitrobacter spp.) Anaerobic N-transformation Denitrification (actually nitrate reduction): C6H12O6 + 4NO3- 6CO2 + 6H2O + 2N2 Nitrification not possible NH4+ + O2 NO2- + 4H+ + 2e Significant denitrification only possible when: 1) There is a nitrate stock (e.g. after flooding) 2) At the aerobic – anaerobic interface 3) Rewetting and drying alternate 4) There is an external nitrate source (e.g. nitrogen deposition, fertilizer application or inflow) Phosphorus cycling: Anaerobic conditions P-availability mainly chemically regulated Chemical binding susceptible to redox and pH Low pH: binding to Fe and Al, high pH: binding to Ca Redox effects: increased solubility of reduced metals (dissolution of insoluble metal-phosphate complexes) Plant adaptations to drought Strategies to minimize water loss: - Leaf structure (grootte, vorm en waslaagje op blad) - Type of photosynthesis (C3, C4, CAM) - Timing of activity Strategies to maximize water uptake: - Extensive root systems Water storage Plant growth under wet conditions Stress from being waterlogged - Lack of oxygen for root activity - Toxicity of reduced compounds in the soil - Lowering of nutrient availability - Physical stress from water flow Plant adaptations to being waterlogged Physiological response: anaerobic metabolism Physical responses: - Aerenchyma formation (and Radial Oxygen Loss) - Adventitious root formation - Petiole (bladsteel) elongation and hyponastic growth (bladeren gaan rechtop staan) During short periods of being waterlogged plants can metabolize anaerobe Salt stress Toxicity (inhibition of physiological processes) Physical damage (crystal formation) Osmotic stress Als een plant in zout water staat kan water bij droogte door osmotische stress uit de plant diffunderen. Stress from acidity/alkalinity Toxicity (inhibition of physiological processes) Mobilization of heavy metals (often Al) Access to nutrients blocked Acidity: Organic soils Rainwater accumulation/infiltration Alkalinity (or neutral conditions): Calcium rich soils, Exfiltration of groundwater The nutrient that gives the biggest growth is limiting. Species interactions: Competition o Nutrients: Better access: Root architecture, Symbiosis More efficient use (NUE): More biomass produced per unit of nutrients o Light/Space: Growth form Active eradication of other species: secondary metabolites Facilitation o Species modify environment, other species also profit Week 38 Definitions: Evapotranspiration = Evaporation + Transpiration Interception = water intercepted by vegetation Interception evaporation Througfall = precipitation - interception evaporation Percolation = drainage to groundwater 1 mm rain = 1 liter / m2 Watercontent of soil: fraction or % of total volume Darcy’s Law: q k q k H z h H z = water flux in (m/d) = Hydraulic conductivity in (m/d) = Hydraulic Head = stijghoofte (m) = Elevation Head = hoogte van de water kolom (m) = Pressure Head = H-z (m) o ∂H= -0.2m o ∂z= 0.4m o We measured a flux of 5 m/d o So: q=5=-k · -0.2/0.4 o k=10 m/d k (depends on grain sizes) o Gravel = 100 - 1000 m/d o Sand = 1 - 100 m/d o Clay = 1e-6 - 0.2 m/d Water flows from high to low Hydraulic head Waterflow in pores: v avg q vavg Isohypsen: waterflow in pores Porosity: 0.35 - 0.5, peat: 0.7 Points with equal Hydraulic head. No flow: ∂H = 0 Large densities of Isolines large water flow (also called equipotential lines) Zuigspanning (soil water pressure) pF = log (-Pressure head (in cm)) pF = log (-h) Natter dan de veldcapaciteit kunnen de meeste planten geen water opnemen. Transpiratie van planten is de grootste flux. Why does a plant transpire? 1. Meteorological: cooling optimal T on leaves 2. Plant physiological: Photosynthesis stomata open for CO2 losing H2O 3. Soil Science: Root water uptake of nutrients Modellen Components 1. State variables: amounts, number 2. Rate variables: interactions (per time) 3. Model parameters: system properties N in soil: Initial amount: N(t=0) = 100 kg/ha Rate constant: a = 0.02 d-1 N(t) N(t 1) aN(t 1)dt N(t) N(t 1) aN(t 1) dt If t is very small: dN aN(t 1) dt Leaf cooling model: Warming up: solar radiation Cooling down: ambientair (H) Latent heat (Transpiration) Test on leaf: with gas-chambers Test on canopy level: with for instance sapflow Test on ecosystem: with eddy covariance Week 39: Excursie: The Horstermeerpolder is one of the deeper polders in the area, located between 2.5 and 3.5 m below mean sea level. Its low position in the landscape is problematic, because the polder now attracts water from adjacent areas, including valuable nature reserves such as the Ankeveense – and Kortenhoefse Plassen. Furthermore, it diverts the deeper groundwater flows as well. Water extraction to keep the polder dry is so intense that it not only attracts water from adjacent areas, but also attracts salt water from deeper aquifers. The ditches in the central part of the polder are therefore brackish, and you can find several salt tolerant species there. Water depletion in the adjacent nature reserves used to be solved by the inlet of river water from the Vecht, inlet of water from the Spiegelplas and inlet of water from the ‘boezem’ (which drains the superfluous water from polders). This caused problems because of the different chemical composition of the water and because of pollution of the river water. More recently, another strategy is being tried. In the southern part of the polder, buffer zones are being created to counteract seepage from the adjacent nature reserves. These bufferzones consist of former agricultural land, which is isolated from the hydrological system of the polder, allowing the groundwater level (or actually, the pressure head) to rise. Predicted by Darcy’s law, this should reduce water flow from the adjacent areas. In the literature assignments, you have read about research that was performed here on the ecological impact of these measures. Week 40 SVAT modeling SVAT: Soil-Vegetation-Atmosphere-Transfer Solar radiation is dependent on the location on the earth. Saturation: Maximum water drainage Field Capacity: Water cannot drain freely. Water can be taken up by plants Permanent wilting point: Water cannot drain freely. Water cannot be taken up by plants. Evapotranspiration 1) Energie Budget: Rn E H G 0 Rn = net radiation E = latent heat = evapotranspiration H = sensible heat G = storage heat in ground (= 0 (versimpeling)) 2) Vapor Transport (Fick’s Law): E 3) 'va va rs ra = latent heat of vaporization (J/kg) ’vs = vapor density at the surface (kg/m3) (‘ = verzadigd in ’vs) 3 va = vapor density at the atmosphere (kg/m ) rs = surface resistance = stomatal resistance (s/m) ra = boundary layer or aerodynamic resistance (s/m) Heat Transport (Fouriers Law) Ts Ta H c ra H = Sensible heat loss (W/m2) Ts = Surface temperature (K) Ta = Air temperature (K) ra = boundary layer or aerodynamic resistance (s/m) c = Volume specific heat capacity (J m-3K-1) The temperature at the surface is very hard to measure! 4) ’vs - va = ’va - va + s(Ts – Ta) ----> Combine 2 & 4 5) 6a) 6b) E 'va va s(Ts Ta ) rs ra H ra c Rearrange 1: H Rn G E T Ta Rearrange 3: s ----> Combine c T T ra Rn E 7) s a ----> Combine 5 & 7 ( 'va va s 8) E rs ra Rn E ra c Al deze vergelijkingen moet je doen, omdat je Ts (temperatuur blad) moeilijk kunt meten. Penman – Monteith equation: s Rn c ( 'va va) ra E s (1 rs ra ) = psychometric constant = c = 0.495 g m-3 K-1 Hoef je niet uit je hoofd te leren! Relative humidity = va 'va = (Vapor density atmosphere / Saturation density atmosphere) ’va is afhankelijk van temperatuur Aerodynamic component (ac) of Penman – Monteith equation: ----> a c('va va) r 'va va 0 Priestly Tailor equation: If air is saturated ac = 0 E s(Rn G) s Replace aerodynamic component (ac) with constant : E s (Rn G) ( = 1.26 for well watered grass) s Coupling: degree to which a plants responds to its environment (CO2, vapor, etc.) If: leaf/canopy completely coupled: E imp ( 'va va) rs assume --> ra very small --> Ts = Ta rs = surface resistance is belangrijk, dus beter Penman - Monteith gebruiken. If: leaf/canopy completely decoupled: E dec s Rn s dus beter Priestly Taylor gebruiken. assume --> ra very large --> aerodyn. comp. richting 0 E E dec (1 ) E coupling = degree coupling 0 = perfect coupling 1 = perfect decoupling rs most effective when = 0 Veg factor = 1 e ( LAI ) Soil – Vegetation: Darcy’s Law: Jw k <-- naar transpiratie d dz k = hydraulic conducivity Saturated zone ksat ~ pore size kunsat ~ soil water content Richard’s equation: Jw k k g U t z z = density water = volume water content = soil water potential t = time k = hydraulic conductivity z = soil depth g = acceleration of gravity U = source–sink term (root water uptake) Simplification: Tipping-Bucket approach Week 41 In het Kruger National Park in Zuid-Afrika vind je ook venen. Ook al is er weinig neerslag en hoge verdamping. Er is heel warm grondwater. Isohypsen: Lijnen met grondwaterstand van gelijke hoogte. IJzer duidt op een kwelzone Ab Grootjans: Wanneer er een probleem is, analyseer dan mogelijke oorzaken eerst. Als je het systeem niet begrijpt is de kans groot dat je door ‘herstelmaatregelen’ het probleem vergroot. Do not jump to restoration! Kalk slaat neer als CO2 uit water aan de lucht ontsnapt. Dit kun je meten aan de hand van EC-metingen in de beekjes. Stroomafwaarts worden de waarden steeds lager, omdat er steeds minder kalk in het water zit. Our expedition has shown that Peninsula Mitre holds a wide variety of mire types over two exciting gradients: – local geology induced gradients from extremely carbonate rich to extremely acid. – a regional climate induced west – east gradient from Sphagnum magellanicum- in the West, via Caltha dioneaefolia-, to Astelia pumiladominated mires in the East of the Peninsula. This wide variety makes the Peninsula to a globally important mire region that deserves full protection. Overgrazing stimulates surface water erosion. Surface water erosion lowers groundwater tables in the peat. Thufur: lifted peat due to frost. Low water tables promote the disruption of thufurs. Frost upheaving is adding to the problem. Week 42 Stomata: exchange of CO2 and water vapor. But the diffusion rate of water vapour is 1.6 times greater than CO2. Guard cells regulate diffusive conductance of the leaf (gs) between 0 (stomata closed) and maximal (gsmax) (fully open) High CO2 Species develop Low gsmax Low CO2 Species develop High gsmax High gsmax can only reached with high stomatal density (D) Miljoenen jaren geleden was er meer CO2, dus planten hadden lage stomatale conductance. Als er meer CO2 is, gaat de plant efficiënter om met water. Phenotipic plasticity: aanpassing binnen de genetische grenzen. CO2 gaat via diffusie de plant in (dus door concentratie gradiënt). Stomatal adaptation at 3 timescales: Relevant for anthropogenic climate change! • Dynamic adaptation (seconds ~ hours) plants open/close stomata • Structural adaptation (> years ~ decades) plants grow leaves different number and size stomata within phenotypic plasticity • Genetic adaptation (> ~centuries) natural selection alters ranges of phenotypic plasticity Canopy transpiration decrease at double CO2 College Max Rietkerk (Semi-)arid ecosystems • Yearly potential evaporation exceeds yearly rainfall • Plant growth water limited • 40% land surface • Main land-use is grazing Infiltration rate increases with vegetation cover Run-off and soil loss decrease with vegetation cover Catastrophic shifts: Are sudden, abrupt, as compared to gradual environmental change Show hysteretic loops (difficult to reverse) Have different threshold values associated with this There are no early warning signals Result from positive feedback In mathematical terms: Spacial bifurcation Self-organized patchiness! Outcome of internal dynamics only, starting from random initialization As a result of plant-soil characteristics Concentration of soil water under vegetated patches This is due to the fact that higher biomass leads to higher infiltration rates outbalancing higher transpiration rates Stefan Dekker Positive feedbacks: Transpiration – precipitation feedbacks Albedo feedbacks Stability analysis doe je door een systeem te verstoren (pertubations) en kijken wat er gebeurt. Multiple equilibria are possible due to: different strengths, history and interplay of feedbacks at disparate scales Week 43 Op kleine schaal: cm*s-1 Op grote schaal: 104 km*103yr-1 Het is dus inefficiënt om deze samen te modelleren. What are the major drivers of photosynthesis? Energy o Light Water availability Plant N content (chloroplasts) Photosynthesis is highly related to transpiration Ways to model transpiration: decrease complexity Penman – Monteith Priestly – Taylor Function of radiation alone Function of min and max temperature. Transpiration cools the leaf due to evaporative cooling. Soil moisture is ook beperkende factor bij evapotranspiratie en dus ook bij photosynthese (zit bij WUE inbegrepen). Nitrogen Taken up from the atmosphere through symbiosis by nitrogen fixating bacteria. Ammonification: Bacteria convert dead matter into NH4+. Kan alleen onder aërobe omstandigheden. Nitrification: Bacteria convert other N-forms into NO3-. Kan alleen onder aërobe omstandigheden. Elke plant heeft een bepaalde C:N-ratio. Detritus = Litterfall (canopy) + Root Turnover Micro-organisms (small animals, bacteria and fungi) decompose detritus, releasing CO2, water, nutrients and more resistant organic matter (humus). Decomposition depends on: Temperature (exponential because of enzymes) Soil moisture (slow in dry soils) Soil acidity (slower in acid soils) Litter composition (nitrogen rich leaves decompose fast, lignins (wood) and polyphenols slow) Oxygen availability (aerobic vs anaerobic decomposition) Autocorrelatie: Memory soil moisture (regenbui) Flip Witte Een overschot aan regenwater zorgt voor verzuring. Three feedbacks of vegetation to drought: 1) Closing of stomata 2) CO2 transpiration T 3) Reduction of vegetation cover Tact = function of soil water suction (Feddes, 1978) Soil water suction depends on: P and ETref vegetation type (Tpot) exposure to the sun soil texture (k(Θ) and h(Θ)) groundwater depth Versterking van de verschillen: Wat droog is, wordt droger Wat nat is, wordt natter Shortcommings of ecological models: Verkeerd gebruik van data. Output GIGO: Garbage in Garbage out. Als je input waarden slecht zijn, krijg je ook slechte waarden eruit. Er is vaak geen random sampling. Je moet altijd kijken of de output van je model realistisch is.