Lecture 1

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Ecosystem Wikipedia definition o Living and nonliving things interacting in a system This class talks about ecosystems at a watershed scale o Like to do this so they can define a boundary Makes it easier to study the inputs Solar radiation, precipitation, dry deposition Exports: gases, water, soil Within boundary, interested in flow of energy and materials o And also internal fluxes (litter production, aka dead leaves, decomposition) Ecosystem ecology sits in the middle of different sciences Related to earth science, hydrology, soil science, physiological ecology, community ecology, social and ecological stewardship History of ecosystem ecology: Starts in 1700s o Platts and Hooke did experiments to show nutrients came from air and water o Priestley put animals in jars Without a plant, animal dies Put animal with a plant, itll live longer (made connection between animal and plant dependence on each other) o MacBride and Priestley- found respiration leads to decomposition Late 1800s- plant systematics o Fredrick Clements- idea that plant communities live like a super organism (little parts making up a bigger whole) o Gleason- individualistic theory on plant communities- plants randomly dispersed to a spot and if conditions were right then they would live, also based on environmental conditions o Tansley- viewed ecosystems as a physical construct- father of ecosystem ecology Chemistry and physics and biology interacting

Tansley ideas: Ecosystems will remain constant or in steady state until disturbed (exogenous events, coming from outside ecosystem, abiotic) Stability and persistence (sustainability)- time compared to biomass graph o Oscillations in graph but if you took the mean, it would basically remain even Founded New Phytologist His stuff wasnt grounded in actual evidence but intellectual theory o We see how science has changed today because he never would be published without the data Late 1800s Lotka- ecosystems as an energy transforming machine o First two laws of thermodynamics Modern day ecology Hutchinson- cool guy, never forgot anyone, met teacher at party and remembered her a year later Lindewan (1915-1942) quantitative paper using ecosystem concepts (using Lake in Minnesota) o Conversion of suns energy o Described succession o Food cycle (nutrient cycle) o Combined abiotic and biotic into one to talk about ecosystem Odums- 1952- first textbook on ecology

Newer things in ecosystem ecology NEON- National science foundation- understanding changes on different scales (global, national and local) o Platform for scientists to do more research without federal grant

Ecosystem concepts: complexity/chaos/scaling1/23/2013 12:02:00 P

Basic ecosystem: Challenge to Ecosystem Ecology is to Define the Fluxes/Rates/Velocities of Mass and Energy Transfer associated with the Arrows and the Size of the Pools The Exchange is made To and From Primary producers: plants, autotrophs o Harvest sunlight to create sugars and feed the rest of the system Soil: o Rich separate ecosystem o Break down complex organic matter into hummus and inorganic matter Ecosystem ecology: the Baldocchi-Biometeorology Perspective Physics Wins, Biology is How its Done o physics sets the limits (how many plants and how big the plants can get) Physics wins: Ecosystems function by capturing solar energy o Only so much Solar Energy can be capture per unit are of ground Plants convert solar energy into high energy carbon compounds for work o growth and maintenance respiration Plants transfer nutrients and water down concentration/potential energy gradients between air, soil and plant pools to sustain their structure and function Ecosystems must maintain a Mass Balance o Plants cant Use More Water or Carbon than has been acquired Ecosystems are Complex Systems

Biology is how its done Species differentiation (via evolution and competition) produces the structure and function of plants, invertebrates and vertebrates

In turn, structure and function provides the mechanisms for competing for and capturing light energy and transferring matter Gases diffuse in and out of active ports on leaves, stomata o Big leaves or small leaves depending on environment (adaptations) Bacteria, fungi and other micro-organisms re-cycle material by exploiting differences in Redox Potential; they are adept at extracting chemical energy by passing electrons; Microbes are pivotal for sustaining ecosystems Reproductive success passes genes for traits through the gene pool.

Corollary to Silvers Rule: Microbes Rule the World They do, given the Energy stored in Carbon Substrate, produced by Plants, eating Sunlight and Consuming CO2 All Biogeochemical Cycles come to a Halt without Microbes and their ability to Recycle Nutrients and Extract Chemical Energy All function, and indeed all life, within an ecosystem depends upon the utilization of an external source of energy, solar radiation, o RL Lindeman, 1942, Ecology microbes may rule the world but they are still dependent on an interconnected system (they couldnt live if they didnt have things to break down) Bottom-Line: Plants and Microbes Work Together as a System, An EcoSystem

Attributes of complex adaptive systems Many Coupled Processes o With Non-Linear Response to Forcing Ex: light, temperature, soil, humidity (abiotic) o And Subject to Feedback o Ex: transpiration, photosynthesis, respiration Hierarchal System Multiple-scales o Power Law Behavior Y = ax^n Plotting log y to log x you get a straight line

Makes it easier to diagnose data because the data you will get will most likely be on that straight line Deterministic/Predictable o Change in some component in ecosystem over change in time Sensitive to Initial Conditions o Path dependency Self-Organization Scale-Emergent Properties o Sum of parts does not equal the whole Resiliency and Robustness o How far can you push an ecosystem before it collapses

System complexity: many interconnected ecosystem processes, with feedbacks Weather drives physiology Physiology is building blocks to what a plant is Growth and allocation of plants leads to biochemistry Once you have individual plants they need to work together in a system and that leads to ecosystem dynamics o Dynamics include: reproduction, dispersal, competition, disturbance, mortality, succession Landscapes form non-linear systems The presence of life causes river systems to build up sediments and be non-linear Without life, the river system will basically be straight Non-linear biophysical processes are ubiquitous in ecology Photosynthesis o Square root graph, levels off at the end Transpiration Respiration Leaf temperature

Why do we worry about non-linear processes

The mean of the non-linear function does not equal the function of the mean

Ecosystems operate across a hierarchy of time and space scales Fast and small gradually to large and slow Hierarchy Hierarchy is defined as the arrangement of some entity (space, time, organism) into a graded series of compartments Hierarchies are nested in that each level is made up of a group of lower levels EP Odum

Biological-ecological hierarchy Organelles, cells, organ, organism, population, ecosystem, landscape, biogeographic region (biome), biosphere Hierarchy of ecologically-relevant space scales Microbes, stomata, leaf, plant, canopy, landscape, biome/continent, globe Super-position of fast and slow fluctuations on carbon flux time series Lots of fluctuations Scale emergent properties a cell is more than its molecules, as an organ is more than its cells, and as an organism is more than its organs, in a food web, new structure emerges at every organizational level up to and including the whole web (Cohen) ex: thought, language, creativity in humans not just a body ex: melting ice cube vs. melting glacier are completely different o melting glaciers are more complex of ecosystems: o ex: the behavior of how photosynthesis of a leaf and canopy respond to light (leaf is square root looking graph and a canopy is linear

complexity is not the same as random, its deterministic Simple sets of Coupled Ecological or Meteorological Differential Equations (dx/dt=f(x)) can produce Complex Behavior o Lord Robert May, Nature, 1974 o Ed Lorenz, J. Atmos Sci, 1963 Complexity: creating order out of chaos and establishing the limits on predictability Chaotic systems are deterministic it all depends on initial state Example of self-organization polygons in the tundra Freezing and thawing and naturally processes led to natural made polygons Sensitivity to initial conditions and path dependence Soccer ball on a hill example, it can roll down one side out one water shed or down the other side and out another way and the difference is a slight difference in initial condition Non-Linearity and Complex Systems are Robust to Perturbations, they but are subject to Regime Shifts, too A Lesson and Warning for Unintended Consequences, when Perturbing Complex Systems Ecosystems like to stay in a state, push them too far and it could collapse Hysteresisanother non-linear response--in Ecology The path taken to collapse an ecosystem isnt the same path reversed to get recovery Know this (graph on powerpoint)

Chaos and complexity in predator-prey dynamics Hair and lynx: populations cycle together Example: periodic crashes in the stock market, another complex system

Managing complex systems forces us to think and act differently

Ecosystems work across a hierarchy of time and space scales that span over 14 orders of magnitude

Discussion 1 1/28

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Hubbard Brook: first watershed scale study where people tried to quantify pools and fluxes of nutrients Cut down trees in one watershed and compared to normal watershed Lots of nitrogen in runoff Power law behavior: ecological behavior doesnt scale linearly

Scaling part 2

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Allometry: Measuring complicated variables with the simple metrics o Measuring trunk of a tree to see how much wood there is/how much it would weigh when its fully grown The study of the Relationship between Size and Shape Non-Destructive o Use tape measure or scale Practical: o Measure easy variables, like tree diameter, can infer difficult to measure quantities, like leaf area, height, and productivity Allometry, a Tool of Ecology and Forest Management You can measure trunk diameter and see the leaf mass o Data follows trends (physics wins, you can only be so big given genes and environment) o Helps with ecological analysis at larger scales Fundamental power laws Y = ax^b log y a b log x o The Exponent, b, equals the Slope of the Log-Log plot of x and y

Overarching Concepts Emergent Features of Complex Systems o Allometries provide useful Rules for Ecological Assessments; they bring order of the complex diversity of ecosystems o Many Biological Allometries have exponents that are multiples of 14 o Are Associated with the Physics of Hierarchal Systems and Fractal Nature of Branching Systems Ecological Power Law Scaling Rules Individual: o Leaf Mass scales w/ Tree Diameter o Leaf Area scales w/ Sapwood Area

o Metabolism scales w/ Mass Community: o Biomass scales w/ Number of Trees per unit Area o Density scales inversely with Size Number of trees per unit area ~ Mass o Biodiversity scales with Area Physical limits to resources or availability

Fundamentals of scaling: Euclidian vs. fractal scaling Euclidian scaling relates to exterior shape o power law exponents are multiples of 1/3 Biological Scaling is Fractal o Associated with internal shapes and networks o Power Law exponents are multiples of 14

Fractal Geometry, Space-Filling Perspective, adds 4th spatial dimension Law Dependency Ex: roots with mycchorizal: root takes up most the space but the mycchorizal have a lot of surface area Fundamentals of scaling Living things are sustained by transport of materials (water, nutrients) through networks of paths. For the network to function, it must be space filling throughout the volume the final branch is scale invariant the energy required to transport material must be minimized o The hydrodynamic resistance must be minimized. Using Scaling Laws to Infer Information on the Properties and Performance of Ecosystems Measure upper leaves in a canopy and scale down from there to see metabolism of a whole ecosystem

Size vs Number Density transcends 9-12 orders of magnitude in an Orderly Manner Physics Wins: You can only be so big and sustain so many individuals for the resources available Corollary 1: You can only grow so Big and So Fast; an Ecological lesson for the Stock Market and the Federal Reserve. Corollary 2: Dont Eat anything Bigger than your Head (Mom) o You can only eat so much because your stomach is only so big and your metabolism only works so fast Yodas Self-Thinning Law Negative slope with density (x) and average total mass (y) o Negative power law mass = a number ^-4/3, number = (mass/a)^ -3/4 o a forest can only sustain a few large trees, or many small trees Kliebers law Metabolic rate (B) of an organism scales to the power of its mass (M) o B = M^3/4 The Metabolic Energy needed to Sustain an organism INCREASES with Mass, to the 34 power Ex: Basal Metabolic Rate (B) of uni-cellular and multi-cellular organisms scales to the 3/4 power of their mass (M)

Metabolic Scaling of Populations of Organisms is Scale Invariant: an Emergent Property of the System Energy flux of a population per unit area (Bt) is invariant with mass of the system (M): When adding scaling laws, add exponents o Helps with looking at entire ecosystem

How weather and climate control ecosystems (and vice versa) 1/23/2013 12:02:00 PM
Weather & Climate Meteors Atmospheric meteors (observable phenomena): o Atmospheric radiation Solar radiation=short-wave Terrestrial radiation=long-wave o Temperature, wind speed and direction, moisture and precipitation, pressure Feedbacks between ecosystem processes and weather & climate All those things listed above control: photosynthesis and respiration, evaporation, decomposition, ecosystem structure, and soil development and nutrients

Weather vs. Climate Weather: day to day variability (chaotic, extreme behavior) Climate: average over a long time (30 year, century) (more stable, predictable) Atmospheric dynamics: Prime example of a dynamic system: many + and feedbacks operating on a range of spatial and temporal scales o Non-linear feedbacks o Sensitivity to initial conditions o Strange (chaotic) attractors o Thresholds/tipping points Complexities with a lot of variables that need to be considered

Edward Lorenz: Butterfly effect Chaotic solutions can arise from a small predictable system of equations o Depending on initial conditions o Small scale things creating disproportionally large effects (interconnection) o Ex: glacial ages, forces shifting between different stable states

Could be normal and then greenhouse gases change and force it to a warmer stable state Teleconnection: atmospheric phenomena at one location can impact dynamics anywhere else in the atmosphere

What controls atmosphere dynamics? Solar constant = 1366 J/m^2s or W/m^2 Energy balance: driven by interactions between the sun and earths atmosphere and surface Global energy balance Earth-sun orientation controls seasonal climate throughout the year Long term changes in solar radiation o Milankovic cycles: Eccentricity: shape of earths orbit around the sun Eccentric orbit or a more round orbit Obliquity: change in the angle or Earths axis tilt Axial precession: change in orientation between Earths axis and the sun Earths axis wobbles like a top Happen on very long time scales Glacial cycles: depend on the integrated amount of solar radiation received during the northern hemisphere summer (Milankovic theory) o If the north has many years of cold summers itll kick start a new glacial age o Combination of the precession, obliquity, and eccentricity control glacial cycles Much of income solar radiation is reflected back to space by clouds or earths surface (albedo), energy not absorbed o Albedo: the percentage of reflected short-wave radiation from surfaces Clouds and white things have highest albedo Energy emitted by sun vs. earth o Sun emits much shorter wavelengths (visible) because its hot Short-wave energy

o Earth emits longer wavelengths in IR Long-wave energy o Result of this is from Planks law IR radiation from earth is absorbed in atmosphere and re-radiated back to earth o Absorption/re-emission of radiation in atmosphere depends on chemistry N- 78%, O2- 21%, Ar-1%, water vapor and CO2 trace gasses Trace gases (<1% of atmosphere) are greenhouse gasses o Atmospheric window: window of wavelengths of IR radiation that does make it into space Rayleigh scattering: why sky is blue, particles scattered at high wavelengths and preventing them from hitting earths surface o Works in shorter wavelengths along with oxygen and ozone The left over net energy used to do work at the surface o Conduct heat (sensible heat) into atmosphere or to evaporate water (latent heat) into the atmosphere o Net radiation = sensible heat + latent heat + storage o Net energy is from income solar and outgoing IR that isnt used

Law of partial pressure: total atmospheric pressure is the sum of the partial pressures of individual gases Pressure at sea level = 101.3 kPa Total pressure = mass of overhead atmosphere x acceleration due to gravity Pressure decreases with height since there is less mass overhead at higher altitudes

Vertical structure in atmosphere Temperature decreases in troposphere, hits tropopause and increases temperature in stratosphere because the ozone particles heat up atmosphere Lapse rate: decrease in temperature with height

Adiabatic lapse rate: decrease in temperature on a rising parcel of air with no heat exchange between the parcel and surroundings o Balloon sent up into air: volume increases as it rises, as volume expands the temperature decreases because particles arent hitting each other as much o Dry rate- 9.8 C/km, Moist rate- 6 C/km At dew point temperature water condenses and forms a cloud, lapse rate slows and a cloud forms Orographic precipitation: topography causes air to rise and cool, forming rain (rain shadow) o Dry desert conditions on leeward side because air let out its moisture already on the windward side of the mountain

Cold/warm fronts and precipitation Warm front- cold air sitting and warm system moves in Low pressure systems: Convergence: air rises, temperature decreases, vapor condenses, clouds form, rain possible o Low pressure because molecules of air are moving up into atmosphere o Air moves into low pressure systems High pressure systems: Divergence: air sinks, temperature warms, cloud formation is suppressed o Air moves away from high pressure system o In California, we have high pressure systems off coast in the summer and thats why we have nice summer with no rain Global circulation Lots of upward movement at equator because it receives a lot of radiation o Air rises at equator due to solar heating Superimposed on earths rotation: coriolis force determines wind direction

Determines global precipitation patters

La Nina Trade winds are strong in the pacific ocean o So strong they make the water higher on the west side of the pacific (Australia) El nino Water built up in Australia sloshes back across ocean towards south America Because pacific ocean is so huge it changes global patterns Trade winds are weak in the pacific ocean as water sloshes back to S.A.

Temperature and precipitation sets the limits on what ecosystems can develop in what regions Feedbacks between ecosystems and climate Climate ecosystems: o Greenhouse gases: Take up CO2 through photosynthesis, release CO2 through respiration, CH4 and N2O cycling o Change in land surface changes albedo Reflective desert vs. absorptive rain forest o Ecosystems have different evaporation rates depending on plant type & structure Evaporation from central valley is causing more rain in Colorado Arctic permafrost melt causes increased decomposition and CO2 and CH4 release

Climate ecosystem interactions

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Plants are associated with Climate (idea goes back thousands of years, Theophratus and Linneaus) Plant communities were integrative with numerous emergent properties (Tansley and Clements) Ecosystems are a random assemblages of adapted species Biogeography: How Does Climate affect the Global Distribution, Structure and Function of Vegetation? o The roles of rainfall, evaporation, potential evaporation and minimum and maximum temperature Vegetation and Climate: Holdridge Classification Scheme 3-d graph of biomes based on temperature, rainfall, and humidity tundra- more water as rain than evaporates so water will seep into ground water or sit on the surface Pros and Cons of Holdredge Classification Pro: o Simple o Driven with Easy to measure Climate Variables Con: o Annual Mean Climate Variables can be poor surrogates for functional biophysical variables, like sunlight, evaporation and soil moisture e.g. annual rainfall does not equal available water in soil e.g. annual mean temperature does not reflect growing season temperature or length of growing season ex: C3 grasses are based on temperature not growing season because in California we have a warm summer and would expect a nice growing season but there is no water so everything dies and those grasses are adapted to winter temperatures

Relation with plant functional types and water balance in North America

Because we are moving into cities, there is woody encroachment on grasslands because fires arent burning like they used to

Mean air temp: Eastern side of Asia similar to eastern side of US London is more similar to Seattle than New York because they are both on the west coast of a big continent Precipitation data: Canada isnt as cold as Eurasia because its not as big and it also has the Hudson bay Potential vegetation: Canada has evergreen needle leaf trees where as the east side of Eurasia has deciduous needle tree o It gets so cold in eastern Eurasia another life form had to evolve Virtual field trip: Forest: broad leaf evergreen and broad leaf deciduous o Evergreens in the tropics o Deciduous in the temperate zone Needle leaf evergreen, needle leaf deciduous o More varied in locations Tropical evergreen broadleaved rainforests o Lots of rain, high evaporation, high potential evaporation (lots of sunshine), high minimum temperature (no frost), high density, lots of biomass Temperate deciduous broadleaved forests o Less rain, colder and cooler so less evaporation, Boreal conifer evergreen forests o Canada, Scandinavia, Siberia o Lower rainfall, shorter growing season, Need 500 mm + of precipitation to sustain evaporation and life of a big tree Biogeochemical cycling

o Big and fast in tropics and gets smaller and slower as you more north into temperate forests and then smaller into boreal forests Temperate conifer forests o Year-round growing season in snow free zone o Nor cal, washington, Oregon, Tasmania, new Zealand, Chile o Tall and short trees, cold, good amount of rain Mediterranean Woodlands: deciduous and evergreen broadleaved o So Cal, France, 500-800 mm rain, warmer temperature, lots of sun (lots of potential evaporation), plants adapt to inner annual variability in weather, not very tall trees Tropical/semi-tropical Savanna o Drought deciduous and evergreen broadleaved woodland with C4 grass understory o Hydrological growing season, wet summer o Frequent fires (every 1-2 years) Grasslands: o Middle of continents o Perennial grasses, summer growing season (C3, C4) o Annual grasses, winter-spring growing season (C3, C4) o 200-500 mm rain o exceptions: grasslands with high rainfall (Llanos, Venezuela) seasonal flooded tropical grassland, with sparse trees highly leached, shallow root systems limit tree growth (unique soils) man and fires push back tree growth and sustain grasses

Wetlands

Dominant environmental controls on Net primary productivity Temperature, sunlight, water Map on powerpoint

February 4

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Deterministic- no randomness in a system In an ideal situation you can still get chaotic behavior Scaling laws: Fractal behavior- using smaller and smaller rulers to measure something o Explains the 4 in D^1/4 o Ex: measuring a coast line or the surface area of lungs Yodas self thinning law: o Mass = aN^-4/3 You can have a lot of little trees or a few big trees Kleibers law: o As your size increases, you are going to burn more energy but its not linear and rate of increase tends to decrease when you get to larger masses

Weather and Climate: Weather: short time scales Climate: long term time scales, average trends Sun emits light in Ultra violet and visible light spectra Earth emits in infrared light

Atmospheric window: certain wavelengths that arent absorbed where long wave energy can escape into space Adiabatic lapse rate Why does it vary with moisture? o As moisture cools and condenses it gives off heat and doesnt cool as much as dry air

Rocks and Soil

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Roots congregate in organic rich areas because of the nutrients and organic rich stuff holds water better Lower elevation has higher temperatures leading to more water loss and less vegetation (scrubby savanna) Less precipitation also Large herbivores like elephants can cause big disturbances to ecosystems Soils are the basis for life Stand on, supply goods and services we depend on Hold most of the worlds biomass Have majority of worlds biodiversity Source of food and water for most plants and microbes o Indirectly become nutrients for other animals as well Contribute to composition (gasses) of atmosphere o N2, CO2, N2O, CH4 (methane) CH4- anaerobic decomposition Methane can also be taken out of atmosphere by microbes Repository of net primary productivity o Soils hold a lot of organic matter (carbon) Store 1550-2000 PgC (peta = p = 10^15) Atmosphere stores 750 PgC, Plants store 550 PgC Oceans store 38,000 PgC Soils are dynamic, always changing o Continuous, makes it difficult to study o Heterogeneous and poorly mixed Products of inputs, internal reactions, and exports Cant understand ecosystems without studying soils

What is soil? Definition: unconsolidated mineral, or organic material on the surface of the earth that has been subjected to and shows the effects of genetic and environmental factors Differs from the materials in which it was derived

o Derived from rocks Where does soil come from? Most soil comes from rocks arriving from plate tectonics Exposed rock subjected to weathering o Weathering- chemical (acid rain), physical (glaciers and streams, wind) , and biological (microbes) forces that break down rocks Parent rock (weathered rock) is called parent material o Different rocks have different chemistry and physical characteristics that weather into different types of soils Weathering: Generally slow (over geological time, over hundreds of thousands of years) but not always slow Hot spots of locations on landscape that because of forces it weathers faster (over decades to centuries) Three main types of rock types that form parent materials: Sedimentary: formed by deposition of mineral particles that settle out in lakes, rives and seas and then are reformed (compressed) into rock Igneous: formed when magma slowly cools o Made up of iron (Fe) and aluminum (Al) silicates Common in magma, depends on rate in which it cools Metamorphic: transformed through heat and pressure and remelted into magma

Rock cycle: Clouds and rain drives erosion of sedimentary material into lakes and oceans Through burial and lithification (pressure) makes sedimentary rocks Add heat and pressure to melt and create metamorphic rocks Melts again into magma and magma cools to form igneous rocks Igneous rocks lifted to surface to go through cycle again

o Metamorphic and sedimentary rocks can also be lifted to the surface Properties of soils: Reflect original chemistry of rock Temperature plays role in driving chemical reactions o Warmer environments have faster chemical reactions, cooler is slower and that affects chemical composition of soil Moisture o Dry areas have less acidic soils than wetter areas o Wet areas have water passing through soil constantly and carries away nutrients in the soil (leaching) Biology o Biological processes produce acids to break down rock o Plants animals and humans store nutrients and prevent them from being lost from the ecosystem Rainforests have most weather soils o High water and biological processes, time as well because they have avoided disturbances for a long time so the rock layer is really deep down in earth State factor approach: Hans Yenning Soil forming factors S = f (Cl,O,R,P,T) Soil is a function of climate, organisms (humans), relief (topography), parent material, and time o recognizes other factors can influence soils but those are the main ones

1/23/2013 12:02:00 PM State factor approach: Hans Yenny Climate- temperature, moisture o Warmer climates leads to faster growth of organisms and more organic matter Warm climates with too much water is bad, need to have the right combinations of temperature and moisture o Cooler climates restrict growth, less biomass produced but it may accumulate because of the slow rate of decomposition o Temperature- warmer faster Chemical reactions happen faster in warmer conditions Not always though o Water- transport mechanism Precipitation coming into ecosystems, water goes into soils and roots and out through transpiration or runoff Water integrates across all aspects of ecosystems Organisms- potential biota (used to be there, there now, could be there in the future) (past, present, future) o Morphology- rooting structures of plants, impacts soil Taproots- big deep roots creates heterogeneity in soil Pockets of different soil characteristics o Chemical properties: conifers produce acidic leaf litter Slows down microbial rates and affects elemental chemistry of soil o Gas exchange- wetland plants push oxygen out of roots which allows form a little pocket to let roots function more In rich iron soils, plants oxidize iron to change Fe(II) into Fe(III) which is less toxic to roots o Myrica Faya- invasive nitrogen fixing plant has affect on structure and function of ecosystem o Besides microbes having a huge impact on soils, humans are the second most impacting Knocking down hills, dumping chemicals into soils, water soils, put waste in soil, compact soils by driving heavy machinery Relief- topography- biggest affect through gravity

o Water and materials move down slope o Ridge to valley = catena (toposequence) Soil temperatures vary along cantenas (depends on aspect and slope) Steepness of slope matters- erosion potential and how fast water moves o Accumulation of small particles in valleys and small particles have more potential to hold water so valleys have rich ecosystems o Water accumulates in valleys and if too much, itll become anoxic and slow down biological processes o Texture- size class of particles Fine particles increase in valleys Parent material- rock that underlays ecosystem and stuff soil is derived from o Earths crust made up of primarily- oxygen, silica, Aluminum, Iron, Calcium, Magnesium, Potassium (makes up most rocks) In decreasing order. More oxygen to less potassium O, Si, Al, Fe in highest concentrations, Fe is only nutrient and needed only in small quantities o Organisms need to survive: nitrogen, phosphorus, potassium, sulfur, Calcium, Magnesium Called macronutrients, needed in greatest quantity by most plants and animals Know this! Limiting nutrients Micronutrients needed in smaller quantities, list in book o Aluminum is not a nutrient people with Alzheimers had high aluminum in their brains some plants take up aluminum to make it toxic or distasteful to herbivores (coffee and tea) time- long term = geologic time (weathering, mountains are built, seas rise) o or short term = days, months, years, decades short term- disturbances

humans impact frequency and severity of disturbances climate change- creates novel conditions ecosystems have never experienced

Rocks weathering into soils Primary minerals- young soils are rich in these, come straight from rock o minerals come from rock secondary minerals- minerals that have been reformed o derived from reformed rocks/minerals o primary minerals break down and recombine into secondary minerals o dominated by Fe + Al oxides or Si oxides warm environments- more Fe + Al oxides cold environments- Fe + Al leach down and we are left with more Si oxides

Soil Biology

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Soil biology and the landscape Bacteria and bacterial colonies make nests (shelters) for themselves o For food or water, or protection, same reason larger organisms do Flora and Fauna found in soil We know nothing about viruses in soil Inverse relationship between size and biomass o Larger size, less biomass you contribute to ecosystem Lots of bacteria are pathogens that make chemicals to do chemical warfare with other bacteria o Gives smell to soil Fungi important in forests- decomposition of plant litter (leaves or roots) Algae: make thin films that cover desert surface, photosynthesis and need light Protozoa: big genomes, hard to understand metabolic processes Nematodes: microscopic worms, predators Earthworms: ecosystem engineers, wholesale movement of soil particles o Take surface soil and push it down deep Soil food web: Archea decompose organic matter Fungi and bacteria eat organic matter and they become food for the higher trophic levels Complexity of soil ecosystems Ecosystems go from the least complex in caves and deserts and grass up to deciduous conifer and rain forests

Typical numbers of soil organisms Forests dominated on a microbial level by fungi In grasslands they are dominated by bacteria Ecosystem roles

Ecosystem engineers Decomposition/mineralization (organic inorganic) Nutrient reservoir Soil structure Soil pH Soil redox

When you have earthworms, forest floor mass (leaf litter) is substantially less than when you dont have earthworms Physical effects- making tunnels Geochemical effects- influence weathering and mixing of nutrients Biological effects- decomposition

Arthropods- break down dead tissue (root, leaf) Make chitin Some are predators (scorpions) Nematodes- important predators Nematode trappers- fungal hyphal rings constrict when a nematode swims through Low amount of root herbivory by nematodes can increase allocation of photoassimilate carbon to roots, leading to increase root exudation and microbial activity in rhizosphere o Microbial population growth (more decomposition, nitrogen fixation, biological processes) o Promotes plant growth even though its grazing on plant roots Essentially causes carbon cycle to spin faster

Protists- live in tight symbiosis with bacteria (bacteria live on them) Live in the gut of soil insects (arthropods) o Help insects eat wood and decompose cellulose Vampyrellids are a group of amoeba that eat fungi Mycorrhizae Fungi types: ectomycorrhizal (exterior of roots, oaks, conifers), o Increase root surface area

o N and P acquisition o Glomalin (soil glue) o Many are obligate and live with only one plant Arbuscular mycorrhizal o Invade roots and exchange carbon and nitrogen Saprotrophic o Decomposing dead plant litter o Food source (mushrooms)

Bacteria and archaea Difficult to culture but ubiquitous > 20% of marine microbial cells, 5% terrestrial cells

Microbial habitats Rhizosphere Aggregates Microbial biomass decreases with depth Microbial functional roles Nutrient reservoirs Microbes and soil organic matter Decomposition o Breakdown of dead animals, plant residues, and xenobiotic compounds Mineralization and immobilization (make new biomass) Soil pH o Soil redox- exchanging electrons Microbes can use over 50 different redox couples to drive their energy generation needs

Energy and ecosystem ecology

Our bodies are stardust; Our lives are sunlight

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Eating the Sun: Converting Solar Energy to Biomass on an Ideal Summer Day Not the Annual Efficiency Graph: wavelength (x) energy per wavelength (y), parabola down Energy loss: o Wavelength outsides photosynthetic spectrum o Reflected o Photochemical inefficiency o Photorespiration C4= 0 C3= 6.1

Potential photosynthetic efficiencies 8 photons per CO2 molecule fixed, 4 e- released with each water molecule split 2%, typical maximum efficiency observed in the field Chemotroph Energy Heterotroph (organic) o Anaerobic, aerobic Lithotroph (inorganic) = work Units: joule = newton meter = N-m = kg^2m^2/S^2 Work = force x distance Force = mass x acceleration Power = dwork/dtime o Watt = J/s Energy flux density = W/m^2 = j/m^2s = kg/s^3

First law of thermodynamics The change in internal energy in the amount of heat amount of work

Energy cannot be created, nor destroyed; it can only be transferred from one state to another. The total amount of energy in a closed system is constant Life and Ecosystems are Open Systems

Photosynthesis has 2 photosystems Photosystem 1 deals with light at 700 nm Photosystem 2 deals with light at 680 nm o Sunlight splits water and gives off O2 and 4 eo Electron transport chain (ADP ATP) Excess energy is lost as heat or fluorescence Photosystem 2 comes before photosystem 1

Energy from redox, microbial battery Reduction, gain of e- (GER) (RIG) Oxidation, loss of e- (LEO) (OIL) R: gas constant: 8.314 T: absolute temperature Energy yields from carbohydrates Aerobic respiration 125 kj/e-, 3000 kj/mol C6H12O6 Solar facts: Solar constant: 1366 W/m^2 o Get this because sun is black body and emits energy according to planks law Solar radiative temperature: 5770 K As surface gets hotter, spectral peak is at shorter and shorter wavelengths Co-evolution of atmosphere and life created ozone to protect us from UV radiation Sun: produces short wave radiation (200-3000nm) (UV, visible, PAR, Near infrared radiation NIR)

Are under the curve of Plancks law is long wave energy lost Terrestrial long wave radiation allows us to have water and life because without atmosphere temperature would be about 255 K Lamberts Cosine Law: The radiation normal to a surface is a function of the projection of area normal to incident rays on a flat surface Picture on powerpoint What happens to photons hitting a leaf: Some reflected, some absorbed, some transmitted through leaves Dont absorb green light radiation Reflect as much near infrared as possible radiation as possible so they dont overheat Albedo highest in deserts and snow Vegetation albedo differs with canopy structure As forests get taller, they trap more light

Snow reflectance differs with the presence and absence of trees

Structure and function of leaves, plants and ecosystems 1/23/2013 12:02:00 PM

Form follows function: Leaves have certain sizes based on environment o Smaller in hot climates, larger in cool climate Redwood trees are so tall because they have a big base 300,000 plant species hard to deal with them all so they try to deal with function (convergent) attributes Diversity in functional characteristics/traits better explains differences in the acquisition of mass and energy (light, water, net primary productivity, nutrients) rather than species diversity The contribution of each species to ecosystem performance is not equal Ecosystem function is controlled by a combination of factors Dominant & Keystone Species, Ecological Engineers and Species Interactions (competition, facilitation, mutualism, predation) Multiple Resource Limitation in Ecosystems Plants adjust resource acquisition to maximize capture of the most limiting resource o Changes in Root/shoot allocation occurs Changes in the environment change the relative abundance of resources, so different factors limit NPP at different times Plants exhibit mechanisms that increase the supply of the most limiting resource o E.g. Symbiotic relation with N fixers Different resources limit different species in an ecosystem

Roles of Structure and Function Supporting Photosynthetic organs o Leaves and plants Intercepting sunlight o Leaf size, shape, thickness, orientation and phenology Mining soil for water and nutrients

o Roots Increasing Reproductive Success o Flower color, type Seed size and shape Phenology, timing of flowering and leafing Facilitating the diffusion of CO2 to Chloroplasts o Leaf thickness, Photosynthetic pathway, stomatal geometry,

Important attributes Physiognomy Grass, Herb, Shrub, Tree Broad-leaved vs. Needle-leaved Tree Fibrous vs. tap roots Life Span Annual, Biennial, Deciduous, Evergreen Pollination and Seed dispersal Insect,Wind Photosynthetic pathway C3, C4, CAM Mycorrhizal Association Endo/Ecto Tolerance Features Drought, shade, fire, nutrient, frost, freeze, heat

Leaf and Plant traits Leaf Form o Size, Thickness and Shape, Longevity, Angle, Photosynthetic Pathway, Specific Leaf Area, Carbon and Nitrogen Content, Photosynthetic Capacity Plant Growth Form o Leaf area density, height, longevity, crown size, tolerance to stress

Shape and size of leaves needles vs planar vs shoots o different surface areas projected vs surface area of needles projected to total needle area Big vs Small

Leaf size, shape and orientation affect the properties of the leaf boundary layer the reflectance and transmittance of light

leafs energy balance o emission of hydrocarbons vs temperature is exponential graph hydrocarbons have unknown function but may be defensive compounds

form follows function needle leaves at the top of canopy are small thin shoots that are good at giving off heat o almost like a desert on the top of a redwood forest Photosynthetic pathway C3- most plants on earth o CO2 interacts with rubisco and forms C3 compound before it forms sugar Trees, dicots C4- tropical grasses (corn, soybean), higher efficiencies o Go through C3 step but use different enzyme in first step CAM- desert species (cacti), open stomata at night o Suck in light during the day and take up CO2 at night Leaf Structure and Function: Emerging Ecological Rules 85% of variation in photosynthesis (per unit mass) is explained by variations in leaf mass per unit area and leaf Nitrogen per unit mass The Thicker the Leaf, More Chlorophyll for Photosynthesis; Too Thick And All Light is Intercepted and CO2 Diffusion through the Mesophyll is Inhibiting, causing a Draw-Down in CO2 and Negative Feedback on Photosynthesis Highest Photosynthesis is with Short living, Thin leaves with high N. Lowest Photosynthesis is with Old Thick Leaves with low N Photosynthesis increase with nitrogen Photosynthesis per unit mass decrease with leaf thickness Photosynthesis decreases with age Evergreens tend to be longer living with lower photosynthesis and nitrogen levels

Thick leaves have more photosynthesis than thin leaves based on surface area but not based on mass area Physical Attributes of Plant Canopies Leaf area index- amount of leaf area per ground area (m2 m-2) o Plantations have high leaf area, Mediterranean have low leaf area woody biomass area index silhouette woody biomass per unit area Beers law> 500 mm/y of precipitation is a threshold for forming a closed canopy > 500 mm/y will allow you to have closed canopy if you dont youll have less leaf density Trade-offs between shade and drought tolerance When light comes through, low shade tolerance. In darker forest, species are more shade tolerant The plants that are shade intolerant tend to be drought tolerant and those shade tolerant are drought intolerant A plant that can photosynthesize at high rates and grow rapidly under conditions of high light is unable to Survive at Low-Light levels (i.e., it is shade-intolerant). A plant that is able to grow in low light (shade- tolerant plant) has a low maximum rate of growth and photosynthesis even under high light conditions

Water and ecosystem ecology

Aluminum is not a nutrient Plants acquire it to lower herbivory

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Hydrological Cycle and Ecosystems Inputs: fog interception, rain, snow o Fog helps redwoods survive drought seasons Stored: soil, surface water Outputs: evaporation, transpiration Coupling between water cycle and nutrient cycles Chemical Potential of Water Chemical potential quantifies the driving force for movement of water between two locations The chemical potential of water is related to the change in the Gibbs free energy of the system, subjected by pressure, gravity, temperature and minor constituents Units in pressure Water potential The total water potential of a system consists of the sum of water potentials forces per unit area associated o Pressure (+/-) Osmosis (-) Soil and Plant Matrix (-) Gravitation (+/-) Turgor (pressure) potential is related to the hydrostatic pressure, as when someone is blow on or sucking on straw that is inserted in a reservoir of water. Its sign can be positive or negative. osmotic potential The presence of solutes reduces the activity of water. Beaker with membrane. Pure water on one side and salt water on the other. Pure water has potential of 0 and water will flow into the salt water matric potential interactions between water and solid surfaces act to reduce the activity of water.

Water interacting with small soil particles

gravity Potential gravitational force is a function of the density of water, the acceleration due to gravity and the height of the water reservoir above or below a reference height: affects limits of how high trees can grow Saturated flow, pure water No matric or osmotic potential Takes Negative Pressure (suction) to lift water against gravity The Gravitational Burden of Water Overhead Imposes a Positive Pressures All about pressure and gravitational potential

Soil-Plant-Atmosphere-Water Continuum Water Moves UPWARD because it flows DOWNHILL Energetically o Electrical analogy: I (current) = voltage/resistance I = water flux density (evaporation rate), voltage difference is water potential difference, resistance is where water is stored so the plant isnt just constantly transpiring water into the atmosphere (soil, roots, xylem, leaves) Trees, Drought and Vulnerability to Embolism 70% of 226 tree species, from 81 sites, operate within 1 MPa of Hydraulic Safety Margin for Injury.. Explains Why Drought Decline is Occurring World-Wide water adhesion breaking leads to embolism and bubbles o a result of drought because pressure of water being pulled up through the plant is more than the water available Relative Humidity: Ratio between ambient, vapor pressure, and saturated vapor pressures, es(T) o water temperature increase saturated vapor pressure

evaporation Evaporation is the physical process by which a liquid or solid is converted to a gaseous state Plant canopies introduce water vapor into the atmosphere via transpiration and the evaporation of water from the soil and free water on the leaves and stems. o Green water- water plants have and use and transfer to the atmosphere Potential vs. actual evaporation Biodiversity affects on biodiversity Depends on types of plants This class is stupid

Soil

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Soil texture: Size class of particles in soil o Sand- 1mm diameter- 62.5 micrometers o Silt- 62.5-3.9 micrometers o Clay- <3.9 Size of particle matters because it determines how much you can pack into a given volume o Comparing silt and clay, packing all silt particles compared to packing all clay particles would lead to more surface area (clay) for the same amount of volume Soil has charged particles in it- opposite charges attract o When charges are opposite, they will hold on to nutrients in the soil + charge is cation, - charge in anion o When they are the same, they will repel and not hold on to nutrients Fertility or plant nutrition- soils that are able to hold on to more nutrients are more fertile o Soils that have more surface area, and hence charge, are

more fertile o Clay usually leads to more fertile soils Sand and silt have much less charge Colloids- <.001mm in size- made up of minerals or organic matter Layered silicate clays o Found most commonly in temperate zone o Crystal structure of tetrahedral sheets held together by oxygen to an octahedral sheet 2:1 layered silicate (2 tetrahedral, 1 octahedral) ions can move in and out of the structure common configurations include: AL3+, Si4+, Mg2+, Fe2+ held together by O^2- and OH one of the elements pops out of the structure and is replaced by something with a particle the same charge or smaller and then the crystal may be charged if the new element doesnt replace the lost charge

o called isomorphs substitution o tend to find more negatively charged soil particles because its easier to take in a smaller charge than a larger one o this is why most macronutrients are cations because if the soil is more negatively charged then its easier to hang on to the positively charged cations o if isomorphs substitution occurs over geological time its called permanent charge because its permanent in terms of biological time 1:1 (1 tetrahedral, 1 octahedral) look like rectangles stacked on top of each other break at the edge to make negative or positive charges at the edges variable charge soils because charge can change over a short period of time reactions happening on edges can change quickly o determined a lot by pH o pH dependent charge soils found in places where they have weathered for a long period of time quick changes in charge can flush macronutrients out of soil depending on charge you can have 2:1 clay exposed to weathering for

Iron o o o o

a long time that changes to 1:1. If you let that weather more you have iron and aluminum oxides and aluminum oxides Crystalline Poor crystalline Amorphous Because they are at the end of the weathering sequence, tend to be poor cation exchangers

Although they do have charge- pH dependent variable charge Red clay soils in tropics Organic matter o Amorphous- no defined structure o Charge dense- holds on to charge well o pH dependent- can be a bad thing but negative above pH 2.5 cation exchanging at pH that occur in nature o sticky and coats other surface Other amorphous material o Allophane, immogalite- materials derived from recent volcanism o High charge density o Weather eventually into iron and aluminum oxides o Important in places like Hawaii or places with recent volcanic activity

Cation Exchange capacity

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Macronutrients- NH4+, NO3-, PO4(3-), K+, SO4(2-), Ca2+, Mg2+ Micronutrients- olluvium?- limiting thing that can slow down nitrogen fixation (needed for nitrogen fixation), Copper, Cobalt, Chloride, and others Cation Exchange Capacity: CEC Soils are like electrically charged billiard balls Temperate soils are relatively young and tend to have negatively charged soil o Negative charge derived from isomorphous substitution Called permanent charged soils (charge properties change slowly) pH dependent charge- variable charge- when pH changes, negative charges will turn into positive charges when you have more negative charges, you can attract more cations o plants have a harder time getting cations and plants need more of them anion exchange capacity can be important too, like for P, and works the same way as anion exchange o N can leach out of systems quickly Little billiard balls are hydrated and covered in a film of water o In the soil there are nutrients floating around and they form an association with the balls o Plant root throws out hydrogen ion into environment and it displaces the cation attached to the ball and the cation is taken up by the plant Or one nutrient can displace another and then with a big rain will be washed out of the system o As the concentration of hydrogen ions increases, it overwhelms the cations on the soil balls and pushes them off Exchange capacity measure in the lab and measured with an index o Use extractions that give an index that give you a scale to relate to something o Index developed in pot growing an onion plant, poured nutrient solution into soil and measured what came out the

bottom (to see how much was retained in the soil and was also taken up by the plant) o Most people use this index around the world o Ammonium acetate pH 7 is standard extraction Flood exchange sights with ammonium and they knock off the cations from the negatively charged balls Collect runoff in a beaker and analyze it for cations and this gives you an index of whats in the soil Flood the soil with another solution and then take another beaker full of ammonium and this tells you how many exchange sights there are Effective cation exchange capacity Used in pH dependent soils (sensitive to pH) o Use neutral salt solution to not fuck up soil pH Same process as first extraction mentioned above o Look for Ca, Mg, Na, H, Al, and K in the solution the sum of those is the effective CEC index of fertility of soils How to calculate CEC 1 mole = atomic mass Ca2+ = 40.08g Cmol = .01 mole, or mmole = .001 moles = mg CEC = cmol +charge/kg soil Put soil in a beaker, shake it and extract it through a funnel K+ = 50mg/kg (concentration of potassium in soil) o 1mmole K+ = 39.01 mg with Al3+ = 26.98, 1 mol = 8.99 because you cut it in thirds o Ca2+ is cut in half when you deal with the charge Atomic mass is 40.08 but you cut it in half because of the charge o Divide the atomic mass by the charge to get the moles

Adsorption- Al>Ca>Mg>K=NH4>Na Soils will preferentially hold onto aluminum

o Al promotes acidity

Nitrogen

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Nitrogen cycling and nutrient cycling used synonymously Because in north temperate zone they were affected by last glacial age and nitrogen is a limiting nutrient to plant growth Humans are drastically altering nitrogen cycle Nitrogen cycle N2 gas in atmosphere o Fluxes into ecosystems through nitrogen fixation (turning it into NH4) o Which gets further nitrified (nitrification) into NO3Lost as NO, and N2O during nitrification or through denitrification Some of that can come back into the process (nitrogen deposition) o Leaching at any step in the process Sources of nitrogen Primary source is nitrogen fixation o 10% happens abiotically, 90% biotically abiotic is lightning biological- symbiotic and asymbiotic o reduction of N2 NH3 NH4 takes a lot of energy have to break triple bond N2 +16ATP + 8H +8e- 2NH3 + H2 + 16ADP +16Pi (inorganic phosphorous) 16 ATP is a lot of energy required energy comes from oxidation of carbon or from photosynthesis plants give up 25% NPP to get nitrogen also requires Mo, and Fe (micronutrients essential to nitrogen fixation) enzyme that catalyzes process is Nitrogenase requires low oxygen or else it will denature legumes are good nitrogen fixers

Who can do it

Symbiotic- bacteria living inside specialized structures in plant roots o Live in nodules on roots o 5-20 gN/m^2/year asymbiotic- free living nitrogen fixers- bacteria that live in litter layer of soil and in soil o have to find own carbon and Mo, and Fe o .1-.5 gN/m^2/year o some are phototrophs and create things to protect process from oxygen o lichens- algae, cyanobacteria produce carbon and do nitrogen fixation with a fungus that provides protection

differences in N fixation rates across ecosystems about 1-5 kgN/ha/year o g/m^2 times 10 = kg/ha highest rates of nitrogen fixation in places with most sunlight, organic matter, where P, Mo, and Fe arent limiting, following disturbance (tree falling) o after disturbance, early succession 20kgN/ha/year isnt sustainable because of energy requirement if N availability increases in an ecosystem, nitrogen fixation will go down

constraints to nitrogen fixation N availability: as N availability increases, N fixation goes down because of energy requirement Energy P, Mo, Fe limiting nutrients Herbivory- herbivores tend to eat high nitrogen tissues and target nitrogen fixing plants

Nitrogen deposition Natural (background) rates are low o Chiloe study- tried to find background rates of deposition when you take people away because people affect nitrogen cycle

.2-.5 gN/m^2/year wet deposition- comes in as rainfall (easier to measure) dry deposition- dust and aerosols, harder to measure o depends on atmosphere, climate, location relative to ocean areas with pollution has more deposition forms of deposition: o NH4+, NO3-, organic N, NH3, HNO3, NO2, NO o NH4 and NH3 come from natural sources Fires, nitrogen fixation, undulates (waste of pigs) o NO, NO2- nitrification, denitrification

Internal nitrogen cycle After you produces NH4, taken up into organisms (plants and microbes) (immobilization) Organisms decomposed by microbes and they take the organic matter and mineralize (ammonification) it into NH4 (part of immobilization process is the production of DON) o As it goes from organic matter to NH4, molecules have to be soluble so it can get into the cell Small concentration of DON in ecosystems, limiting factor, rapid flux Some organisms can take up DON (dissolved organic nitrogen), like in the arctic where the organic matter decomposes slowly and there isnt much nitrogen fixation happening NH4 nitrification (nitrified) into NO3 o NH4 + 2O2 NO3- + 2H+ + H2O o Chemoautotrophic bacteria do this Two steps: NH4 NO2- by one certain group of organisms (requires oxygen) (redox reaction, energy gaining step) NO2- NO3- by another group of organisms o NO3 can be immobilized and taken up by organisms Smaller molecule, more soluble, anion, more competitive as a plant if you can use NH4 and NO3

Nitrogen cycling part 2

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Microbes secrete extracellular enzymes into environment to help break down complex molecules into smaller ones so they can pass into the cell Enzymes are costly to make, spend nitrogen to get nitrogen, spend energy to get energy Carbon nitrogen ratio controls immobilization and DON Correlated with ecosystem processes Use ratio of substrate and microbes to understand how much nitrogen is going to be tied up in organic matter and how much is recycling Growth efficiency of microbes is about 40% (deals with NH4 part of N cycle) Assume C:N ratio of 10:1 o Have to maintain ratio, controls what you can and cannot metabolize Say microbes break down 100 ug of C, only incorporate 40 ug into their bodies (immobilization) and respire 60 ug carbon o For the 40 ug carbon, they need 4 ug N (10:1) o For original 100 ug, C:N ratio is 25:1 If there is a 30:1 ratio, the microbes keep nitrogen to themselves because they need it If its 20:1 there will be nitrogen in the environment because they wont need all of it C:N ratios help us understand rates of mineralization

In N cycle, NH4+ NO3- (both forms are inorganic or mineral Nitrogen, most commonly used by plants (especially in tropics and temperate) In boreal, its more dissolved nitrogen Measure availability of NH4 and NO3- through net N mineralization assays

To measure net ammonification = NH4 (at time final) NH4 (at time initial) all over time is change in ammonium Same for net nitrification (NO3f-NO3i)/t Net mineralization = net ammonification + net nitrification

o Need both numbers because its switching between NH4 and NO3 so quickly The problem with this is that its graphed linearly but in real life it doesnt happen that way (non linear processes) Net mineralization is a good index of nitrogen over long periods of time Tropical forests tend to not be limited by nitrogen Ammonification and nitrification tend to be the same North temperate systems, NH4 pool is larger than NO3 In dry systems, there are high rates of nitrification compared to ammonification Gross nitrogen mineralization Use N14 (more abundant, N cycling around) and .1% N15 isotopes N15 is isotope tracer, add it to total nitrogen and ecosystem to trace them as they move around o You see 15NH4 15NO3 Process studied over hours to days unlike the other process above which is days to weeks o This is because you have to measure it prior to recycling NO3- can affect ecosystems Acidification- NH4+ uptake 1 H+ o Plants maintain charge balance so they spit out H+ Conversion of NH4 NO3 will yield 2 H+ o 2NH4 + 3O2 2NO3 + 2H2o + 4H+ NO3 uptake net yield of 1 H+ NO3 leaching means there is still 2H+ and you tend to lose a cation Dissimilatory nitrate reduction to ammonium (NO3 NH4) (DNRA) Microbes convert NO3 back to NH4 to conserve nitrogen Low redox process, requires labile C and some NO3o Requirements for denitrification More DNRA means less nitrogen lost during denitrification Keeps nitrogen in ecosystem, anaerobic process

All soils can do this process

Human Alteration of the Nitrogen Cycle 1/23/2013 12:02:00 PM

Nitrogen losses from the cycle: Next to nitrate leaching, gas loss is the most important Gas loss in forms of NO (nitric oxide) and N2O (nitrous oxide), and N2 (dinitrogen), NO2, and NOx o NO, N2O, N2 are main components Nitrification from NH4 to NO3 is like a hole in the pipe model o As NH4 is nitrified, some of it leaks out in different forms (NO, N2O). This is .1-10% of nitrification NO flux tends to occur in drier environments after rain N2O comes out in dry and wet conditions N2O is most potent greenhouse gas, catalysts for stratospheric ozone loss Ozone in troposphere is pollutant Protective layer in stratosphere Denitrification is an anaerobic process, requires NO3 and C o When its complete it goes to N2 and completes the cycle o Primarily a microbial process and can happen anywhere (extreme deserts) Facultative anaerobes- can breathe oxygen when its there but after it rains and there is no oxygen they breathe nitrate More NO3 you have, the more probability of N2O Nitrous oxide reductase? gas also lost from ecosystems NH4+ + OH NH3 Short half life in atmosphere and gets rediposited shortly after going up into the atmosphere Stimulates nitrification when it goes into the soil and you are

o NH3 o o o

going to acidify the soils Anamox o Anaerobic, ammonium oxidation o Take NH4 + NO2 N2 NO2 comes from the middle step in nitrification Important in marine ecosystems

Human alterations

Human activity has doubled rate of N fixed in the environment o N2 fixed nitrogen Causes of N fixation o Electric power plants (burning fossil fuels), automobiles (fossil fuels), agriculture (synthetic fertilizer and leguminous crops) Fixed n- lightning < free living bacteria < symbiotic N fixation Human doubling n fixation- fossil fuel combustion < fertilizer production < cultivation of N fixing crops o Haber came up with the process to break N2 bond (also boshe too) Much of this process went into bomb making (militaryindustrial process), and agent orange Synthetic fertilizer being used in the tropics and this is bad because they are nitrogen rich soils already and adding it leads to more denitrification and more N2O production

Fossil fuels: Formed during the Carboniferous period in the Paleozoic Era (360286 MYA) o Anaerobic conditions, microbes arent breaking down the organic matter Hydrocarbons from remains of plants and animals compacted, hardened and preserved o Preserved as Coal, Oil, and Natural gas Contain nitrogen that oxidize during combustion

Correlations between human population and nitrogen fertilizer use Steady increase in nitrogen use, most increase in developing countries People buy more fertilizer to make more food which sustains more people

Luxury consumption- plants will keep taking up nitrogen if its available even if they dont need it What goes up, must come down

The more nitrogen you add to ecosystems the more that is eventually in the atmosphere o Reactive Nitrogen oxides and NH4

Nitrogen cycle again

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Wet deposition- fog and rain deposition Tropics arent nitrogen limited and so increased deposition means nitrogen leaks out because its not needed Stationary sources: coal fired electric utility plant Mobile sources: automobiles, planes Dry deposition: pollutants suspended as particles or gases Nitrogen deposition monitoring programs NADP/NTN- national atmospheric deposition program/national trends o Monitors wet deposition (NH4, NO3) Urban and rural sites o Puerto rico site- nitrogen deposition increasing (NO3) Coming from across the ocean (US), developing low lands in Puerto Rico (local) CASTNet (EPA)- clean air status and trends network o Monitors dry deposition (S and N) and ozone Areas outside urban setting

Nitrogen deposition Atmospheric nitrogen deposition initially stimulates (especially in nitrogen limiting environment) growth since N is usually limiting nutrient o Acts as fertilizer Negative consequences of too much nitrogen o Plants use all the nitrogen they can use and it leaks out Called nitrogen saturation Leads to an increase in N2O from denitrification and nitrification o Leaching of NO3- toxic to life and humans Seeps into ground water, blue baby syndrome Kills fish and wildlife o Loss of essential cations (K, Ca, Mg) Increase H+ ions and causes other cations to leach out and declines soil fertility

o Reduced plant productivity and biodiversity We are seeing losses of species in US because of high N deposition for a long time Ex: sugar maple tree (starving, more sensitive to cold winters, more susceptible to pest attack) N2O levels increasing with industrialization, global warming potential Greenhouse gas, N2O has 290 times higher warming potential compared to CO2 3 sages of nitrogen saturation in terrestrial ecosystems early stages: most N used by plants increased productivity later stages: N leaching leads to leaching of other nutrients nutrient imbalances Al toxicity (harder to leach) and Ca and Mg limitation decreased productivity graph: foliar N keeps increasing, NPP increases but then it decreases, Ca:Al and Mg:N ratios decrease nitrate export and acidification of rivers linked to fish kills in mountain lakes waters most affected during early season high flow events o nitrogen builds up before plants take it out and then the water flushes it all out

examples of nitrogen saturation- eutrophied waters algae blooms in Baltic Sea, Chesapeake Bay, Long Island sound Nitrogen concentration in rivers is direct correlation to people living in water sheds From fertilizer use, sewage, runoff, N deposition from precipitation Mississippi river: doubled since 1965 Increased in northeast rivers as well

Effects of nitrogen deposition in N-rich environment Puerto Rico site o Add 50 kg N/ha/year since 2002

This is a ton of N but because it already has a complex N cycle and other cycles, they need to add a lot to detect change o Soil carbon increased with increased N in surface soils Trees didnt grow more so where is it coming from Maybe trees grew more roots Maybe a decrease in carbon respired Labile carbon decreased (easy to break down), while recalcitrant carbon (hard to break down) increased o Turns out root biomass actually decreased So leading to the conclusion that soil respiration decreased It could be decreased root biomass lead to decrease in soil respiration (but this is a small impact) Microbial respiration declines (lab) As it gets warmer, impact is larger Changes in plant and microbial communities o Impacted enzymes breaking down C, N, P o Impacted oxidative enzymes breaking down wood and more complex things

Phosphorous
Nitrogen and phosphorous are linked Both limiting nutrients

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Stoichiometry- elemental ratios C:N, N:P o N:P ratios vary widely across ecosystems Phosphorous sources Primarily comes from weathering of parent material o Apatite minerals (Ca-PO4 mineral), Ca breaks down and releases P o Spike of P in young soils and less over time Especially when you dont have more primary minerals to weather No real gas phase Atmospheric deposition- redistribution of P across ecosystems o Can be pretty significant but usually at low concentrations o Comes in as dust In low amounts but accumulates over time

P cycling Unlike N which is mobile, P is immobile o P is anion reacting with cations and forms strong bonds P + Ca hydroxyapatite with low solubility (secondary minerals that form in the soil) o As it gets more basic, it becomes less soluble Forms under basic conditions In highly weathered Fe Al soils, p reacts with Fe and Al o Form under acidic conditions Occulusion- when P goes into the internal part of mineral o Weathered soils, P migrates into inner crystalline structure and doesnt go out until its weathered away Sorption potential- ability of soils to grab on to P and hold it strongly

Strategies to get P

Morphological adaptations: o Root hairs Biochemical adaptations o Exudates that increase solubility of P in vicinity of root Citrate, oxalate, phosphatase enzymes (increase when they are looking for P or when they are taking it up) Physiological o Retranslocation- Plants pull P back into plant before dropping leaves and losing P Mycorrhizal associations o Myco = fungi, rhizae = root o Symbiotic relationship between fungi and root o Fungi gets carboydrates, root gets nutrients (P, N) Lots of mycorrhizal in low P soils o Fungi put out lots of hyphae (thin root like structures) and increase surface area that root has access to to take up nutrients

Tropical forests on highly weathered soils are P limited Lots of Fe and Al, hot weather with a lot of rain P should be really low High NPP o How do they grow so fast and big with lots of organic matter with highly weathered soils? Direct nutrient cycling- when organic matter dies and is deposited on ground, nutrients go right back into plants and not into soils o Roots invade organic matter and catch nutrients before they get into soil o Grow roots from branches and stems and not just the bottom and these take up dead organic matter (arboreal roots) o But this isnt enough to sustain the forests over long periods of time Fe2+ doesnt bond as strongly to P as Fe3+ so if you have more Fe2+ its easier to get P o When we test soils we turn Fe2+ into Fe3+ so we think there is way less P than there really is

Nutrients into plants

Three main ways nutrients get into plants Diffusion Mass flow Not as important, direct contact

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Diffusion: most important As potassium moves into root, its creates a zone of depletion around the root where there is no potassium o This zone creates a chemical gradient where there is a lot of K outside the zone and none inside o The K moves from high concentration outside the zone to a low concentration inside the zone and is taken up by the plant Ca2+ and Mg2+ have more charge and will move more slowly o K+ and NO3- move faster Bigger sized ions move slower than smaller ones Coulombs law- F (electrostatic force) = charge/radius squared o Force between two point charges Zone of depletion depends on charge o Mg2+ will have a smaller zone of depletion o K+ will have larger zone of depletion Mass flow: important Nutrients moving through water (water stream) Transpiration- movement of water through soil to roots through plants to atmosphere Important when there are high concentrations of nutrients around the root and there is a small concentration gradient Important in micronutrients Works best when there is saturated flow- water flow driven by gravity o More than transpiration alone, completely connected water column in soil

Direct contact: byproduct of nutrient uptake Mass flow and diffusion arent possible without direct contact but direct contact doesnt bring nutrients near/in roots

Mycorrhizae- fungi roots Symbiosis Balanced parasitism o Plant keeps fungi in check so it doesnt damage it and it can extract nutrients from fungi. Fungi extract carbohydrates from plant without killing it Increase soil exploration/exploitation o More surface area to take up nutrients o Important for immobile nutrients (P) Nutrient uptake: Supply rate- nutrient concentrations in soils o Important under steady state conditions (when there isnt a disturbance) Root length- root length able to absorb o When roots lignify and become woody, they are worse at absorbing nutrients o This deals with newer roots Apogeotropic roots- grow towards light, against gravity o Grow up, out of the soil o Environments that are very humid so the root doesnt desiccate Root activity

How nutrients get inside root Nutrients are higher in roots than surrounding environment o Need to transport it away into plant so it doesnt leak out Carrier and transport proteins- helps move nutrients across soilroot interface o When root takes up nutrient, they need to keep chemical balance by spitting out H+ o Nutrient uptake/cation uptake acidifies the soil Soil is not just constantly acidified because they take up anions as well and nutrients can leach out

Area right around root isnt super acidic as you would think, microbial communities help balance pH Root exudates- stimulate microbial activity so roots can take up what the microbes dont

a tree in a nutrient/water rich environment will grow quickly and drop leaves and slough off roots into soils new tissues have higher photosynthetic ability so keeping newer leaves present and dropping the older ones will allow it to grow faster roots lignify when they get older, new roots take up nutrients better as growth rate falls, litter fall decreases as well where nutrients are limiting, this isnt a good strategy o retranslocation- before they drop the leaf, the plant removes the nutrients from the leaf N,P,K These relative to Ca because Ca doesnt get retranslocated as much/not at all (because it is found in cell walls)

Resource use efficiency Light use efficiency- amount of growth per unit light Water use efficiency- growth per unit water taken up Nutrient use efficiency- growth per unit nutrient taken up o NUE- short lived plants, An= nutrient productivity * residence time = Tr An * Tr = NUE Grams biomass/grams nutrient o Long lived plants- look at the litter fall because these are proportional to rate of growth NUE = litter fall mass/litter fall nutrient content

Terrestrial carbon cycle

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General carbon cycle Carbon in atmosphere taken up by photosynthesis (GPP, Gross primary productivity) o Used for growth and plant respiration Plant respiration fluxes carbon back into atmosphere About 50% o From growth biomass- mortality lost into litter Harvest lost to wood products o Litter supports heterotrophic respiration- microbes CO2 atmosphere Some carbon goes into organic matter which leaks CO2 out later on o Litter burns in fire and leads to black carbon CO2 in atmosphere eventually Terms and units tend to be annual on larger time scales 1 m^2 = about 70g GPP: Vc = carboxylation = CO2 carbohydrates (C6H12O6) Photorespiration = Vo = oxygenation o CO2 + rubisco C3 Rubisco likes oxygen and CO2 so in the presence of oxygen in will take that up and release CO2

NPP: NEP: GPP minus autotrophic respiration (R auto) o R (mass, growth, T) NPP = GPP Rauto Autotrophic respiration is respiration of the self-feeders, the plants (leaves, stems and roots) Rauto is a function of growth rate, temperature, mass of the organism Ra/GPP = .5 for a wide range of mass, growth, T

Net ecosystem production = NPP heterotrophic respiration = -NEE (net ecosystem exchange) o NEE = fluxes of carbon in and out of ecosystem Office hours this Heterotrophic respiration is respiration of fungi, aerobic bacteria, invertebrates and vertebrates in the soil It is a function of temperature, soil moisture, carbon content, its lability, and priming from recent photosynthesis o Heterotrophic respiration = Rh Rh compared to T, exponentially increases Drier soil reduces Rh Increase in leaf area index increase Rh

Net Biome production: NBP NBP = NEP carbon loss via disturbance (Fc) o Fire, herbivory, disturbance Mauna Loa observatory measuring atmospheric CO2 Isolated site, measure of CO2 in atmosphere Annual fluxuations of CO2, general increasing trend o CO2 goes up in atmosphere in winter time, more respiration than photosynthesis o CO2 goes down in summer because more photosynthesis than respiration

Global carbon cycle: Gross fluxes and Pools Units: peta grams Pg = 10^15 g = 1 GtC (gigaton) o Equivalent to a 10 micron layer of water per meter-squared across the terrestrial globe o 1km^3 = 1 Gt Pools- 843 PgC @ 385 ppm in atmosphere o Vegetation is 650 PgC o Oceans = 38,000 PgC o Soil = 3194 PgC Big stores of carbon in permafrost

If you melt permafrost, C goes into atmosphere

Fluxes: o Annual basis, global GPP = 120 PgC o Global scale long term NPP = 60Pg Rh = 60 Pg Balance between photosynthesis and respiration Disturbed by deforestation and fossil fuel burning o Deforestation = 1.5 PgC/year o Fossil fuel = 8 PgC/year

Mass of atmosphere: F = m*a = P*a Mass = 5.3*10^21 g air o C in atmosphere @ 393 ppm (393*10^-6) 860*10^15gC o 2.19 Pg/ppm Evidence of fossil fuel combustion: 13C isotope record Plant based Carbon has a 13C signature ~ -25 per mil o This is in fossil fuel because it was taken up by plants long ago o Combustion of Fossil Fuels Dilutes the Atmospheric Background so the graph is negative and dropping Ecosystem Service: Only ~45% of CO2 emitted into the atmosphere remains there More carbon going into oceans, making it more acidic Fx (fraction?)PgC/2.19 Pg/ppm * 47% = ppm The amount of fuel we burn in 1 year took 175,000 years to sequester Most Coal Deposited during Carboniferous, 300 Ma Reservoirs containing the highest concentrations of N per mass are: petroleum (100-20,000 mg kg-1) coals (2000-30,000 mg kg-1) modern marine sediment (1772 mg kg-1 77 )

shales (600 mg kg-1), limestone (73 mg kg-1 78 )

glacial periods low CO2 (180 ppm), low temp o carbon lost from atmosphere to cause ice age interglacial higher CO2 (280 ppm), higher temp

Carbon cycle part 2 (ecophysiology of leaves)1/23/2013 12:02:00 PM

Basic photosynthesis stoichiometry 6CO2 + 6H2O + (light) (C6H12O6) + 6O2 photosynthetic pathway C3 o Trees o California grasses are C3 because they are Mediterranean C4 o Grasses, tropical/subtropical settings (Maze, sugar cane) o C4 is favored under low CO2 conditions and high temperature conditions Glacial periods with low CO2, C4 grasses expand CAM o Desert species and Pineapple

Photosynthesis- balance between supply and demand Biology drives demand: Biochemical limitation: carboxylation rate o Light limitation o Enzyme limitation Physical limitation: delivery of CO2 to leaf o Diffusion through Leaf Boundary Layer o Diffusion through Stomatal Pores o Potential Gradient between free Atmosphere and substomatal Cavity

C3- Calvin-Benson cycle Light reactions o Chlorophyll, in chloroplasts, captures photons o Photons split water molecules and release electrons Light energy is used to produce chemical energy in the forms of NADPH and ATP ATP is energy source, NADPH is electron source Oxygen is produced reactions Bind CO2 with RUBP (C5 compound) C6 splits to form 2 C3 sugars

o Dark o

o Reaction catalyzed by enzyme Rubisco Rubisco likes oxygen so its less efficient than C4 photosynthesis It may take up oxygen and leak CO2 Rubisco accounts for about 9-25% of the N in the leaf o Chemical energy (NADPH, ATP) used to generate RUBP Graph on powerpoint

Using solar energy for life Visible solar energy (400 to 700 nm) is absorbed by pigments This energy is converted into high energy compounds, ATP and NADPH, by photosystems II and I (PS II and I) o Photosystem II uses 680 nm energy to generate ATP (noncyclic electron transport) o PS I uses 700 nm solar energy to generate NADPH (cyclic electron transport). o Excess energy is lost as heat or fluorescence. 8 Photons per CO2 molecule fixed o ideally- not in the real world because energy lost as heat

C4 leaf Contain bundle sheaths in mesophyll C4 photosynthesis The enzyme PEP Carboxylase catalyzes a reaction between CO2 and phosophenolpyruvate (PEP) to form a C4 compound The C4 compound is transported into the specializes cells, the bundle sheaths, and is decarboxylated o When it is transported, there is no oxygen in bundle sheaths o Extra step but more efficient than C3 CO2 is released into an environment and photosynthesis is completed via the C3 cycle Photorespiration is low; RUBISCO favors CO2 in this environment because the ratio between CO2:O2 is high

C3 vs C4

Diets reflect- Americans have C4 signatures (corn), Italians have C3 signatures (pasta) Ci/Ca = .7 for C3 o Higher concentration of CO2 internally for C3 o Stomatal conductance for C3 is more than C4 Ci/Ca = .4 for C4 o Water retention is greater in C4 than C3 o Ca?? Light use efficiency, CO2 and temperature o C4> gunnutum yield- for every photon they absorb they take up more CO2 Light vs photosynthesis graph. Linear correlation Steeper line is C4, flatter line in C3

Comparative ecophysiology chart Study this Krantz anatomy- C4 plants have one, because they have bundle sheaths CO2 compensation- where photosynthesis and respiration are balances o Graph of CO2 vs photosynthesis- square root curve Higher rates in C4, compensation point is <10 ppm C3 point is 30-80 ppm (where is crosses x axis) More carbon uptake per photons absorbed in C4 PEP grabs all the CO2 it sees and pumps it into bundle sheaths

Ca

Has to fuse through boundary layer Has to fuse through stomata o Between Stomata and mesophyll, Ci = .7Ca Fuse through mesophyll And then it reaches chloroplast

Gross photosynthesis Vc-Vo Net photosynthesis includes dark respiration

Dark respiration vs temperature Temp on x axis Relatively flat until it dark respiration exponentially increases at high temperatures Leaf photosynthesis vs temperature Temp on x axis Increases until around 20 C where it flattens out and then starts to decrease at really high temperatures Biochemical limitation is not enough rubisco to drive reaction Vcmax scales with maximum rate of electron transport Capacity of leaf to do photosynthesis varies throughout the year Live fast, die young in stressed environments Positive relationship between photosynthesis and leaf nitrogen o Rubisco has a lot of N As soil gets drier, water potential gets more negative, stomata has less water and they become more flaccid and close Leads to decrease in Vcmax

Midterm review

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Computations, fill in the blank, matching, short answer, multiple choice SI front of each page and name on the back of the last page Calculating CEC, nutrient use efficiency Look at units and unit sheet online Feamox- NH4+ NO2- NO3 NH4 and NO2 produce N2 (anamox) NH4 + FeIII N2 (Feammox) Dark reactions- CO2 reacts with RuBP (C5) and catalyzed with rubisco This forms C6 compound that splits into to C3 compounds Rubisco likes oxygen so it may leak CO2 C3 photosynthesis does have photorespiration, C4 doesnt Kranz anatomy- use PEP to move C4 into interior when oxygen levels are low This gives off CO2 that goes through C3 photosynthesis Light reactions are splitting water and producing ATP and NADPH which helps regenerate RuBP Fractal scaling Use smaller ruler and get a bigger result o Positive log scale Euclidian laws- slopes of fractal measuring are on the orders of 1/3 Metabolism scaling- a function of biomass to ^3/4 power Density of trees vary with mass (^-3/4 power) o Mass (x), number of trees (y) decreases with -3/4 on graph Multiply metabolism and density together and you get a flat line with m (x) and B (y) for the entire ecosystem It helps tell you how individuals function and not how they assemble in the grander scheme of things

Higher nitrogen concentration increases albedo- controversial

Large scale things

1366 W/m^2 mean annual solar content radiation coming from sun plants in tropics- evergreen broadleaf no frost, no freezing hole in pipe model- during nitrification NH4+ NO3 o NO, N2O leaks out during transaction because of activity of microbes o NO2 is middle step between NH4 and NO3 Silicate clays 2:1- charge comes from isomorphs substitution 1:1- as pH changes, the exchange sites are on the edges of the rectangles can change as well if you have too much nitrate is inhibits nitrous oxide reductase enzyme that enzyme is responsible for taking N2O N2 o so the cycle doesnt go to completion (stays as N2O) o under very strongly anaerobic conditions you have more N2 N2O:N2 concentration is controlled by anaerobic conditions and nutrient availability o N2O needs aerobic Butterfly effect: Some variable will change with time in a deterministic way If you change the initial conditions a little bit, over time it will diverge from the deterministic

Nitrogen leaches mostly as NO3 DON can leach as well Variable charged soil, dominate anion exchanging, NH4 will leach as well

Nitrogen is soluble and therefore can leach Low nutrient availability favors evergreen leaves Carbohydrate is electron donor, oxygen is electron accepter Tall forests trap more sunlight

Decomposition

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Carbon cycle Photosynthesis brings carbon into plant material through GPP o Plants respire some carbon off Autotrophic respiration, about 50% carbon that comes in NPP- net carbon fixed Litter fall dissolved into dissolved organic carbon o Can leach out, get eroded, or is taken up by soil, some is respired during heterotrophic respiration Under steady state conditions: Net ecosystem production will equal net primary production heterotrophic respiration NEP = NPP Rh NEP = NPP +/- F-lateral (carbon transported?)- (Rh + F-removal) o F-removal- carbon removed via disturbance (fire) Heterotrophic respiration- respiration that occurs under aerobic conditions and anaerobic conditions Under anaerobic conditions microbes use different electron acceptors o O2>NO3->Mn3+>Fe3+>SO42->CO2>H+ o Anaerobic respiration is much less efficient and slows down and this tends to lead to a big build up of organic matter CH4 (methane) is a byproduct of some anaerobic respiration and we care because its a greenhouse gas

Who does decomposition Mostly microbes Break down macromolecules Hang out on surface and litter layer Good at breaking down wood (lignin) More mobile than bacteria Tentacles send out to move and grab stuff Detritivores- soil invertebrates o Crunch and crush, break down litter and sometimes eat it o o o o

o Dont really decompose it because they dont really break down the macromolecules Start the process so the microbes can break it down Earthworms are important because they are lined with microbes so there poop is decomposing rich Bacteria o Arent very mobile, desiccate easily o Do better in variable redox conditions Facultative bacteria- switch between redox based on conditions o Important for inner soil Fungi arent very good with anaerobic conditions

Controls on decomposition (most focus on top three) Climate o Faster decomposition in warmer places o Warm and wet faster Not too wet because itll become anaerobic o Cold and dry slow o PET = potential evapotranspiration- the amount of evaporation that wouldve occurred if there was sufficient water available o AET = actual et= amount of water removed via evapotranspiration o CDI = climate decomposition index- modeling variable using monthly values of precipitation and temperature Calculates temperature influence on decomposition Shows threshold points of very cold and very hot when microbes turn off Good on large scale Litter quality o Physical and chemical characteristics of the material being decayed o Chemical factors Lignin- complex macromolecular structure Microbes have figured out how to deal with it

Plant compounds- tannins, polyphonolics Make it hard for cells but bacteria can deal with it Make it so bugs dont eat them but its also hard on the microbes so the processes slow down Unfavorable elemental ratios (stoichiometry) EX: Ca:Al, C:N Low nutrient content Ex: N and P limitation Across ecosystems, climate is important. Within ecosystems, litter quality is important Microbes exude enzymes into environment to break it down

Site conditions o Soil conditions Moisture (not precipitation) Temperature (warm faster) Redox If oxygen gets consumed, not a lot of decomposition Nutrient availability Microbes need nutrients just as plants and animals Need nutrients in soil to stimulate decomposition High in Fe, Al, Ni can be toxic to decomposers Texture Finely textured soils (clays)- hold a lot of water so it can increase processes but too much water will slow down the processes More surface area means more CEC Sorb organic matter Aggregates can lock soil organic matter away from microbes Use 14C dating to see how old it is Can use these assays to study C sequestration (carbon storage/flux)

o Organisms

Decomposers Microbes and fungi, detritivores Heterotrophs So many things use carbon so its hard to figure out who does what, how, and how to manipulate them Microbial succession- during decay, you change the quality of the material When the material changes, the microbes change as well and those that eat the microbes change as well The whole food chain cycles in microbial succession

Plants balance needs so thats why they make woody lignin stems roots and branches even through they are worse at absorbing minerals

Decomposition 2
Decomposition = mass loss/time

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Decomposition over time isnt linear Phases of decomposition o <1-6 months artic environment, 1st phase is slower, tropics is much faster leaching of soluble organic matter o 1-5 years break down of cellulose and other easily decomposable compounds o >1-5 years slow loss of recalcitrant (difficult) material time (x), mass loss (y)- linearly decreases in first few weeks/months o then it exponentially decreases from there getting less steep as time goes on o Litter (at time t) = L (at time 0) e^-kt = LnLf/L0 = -KT k = decomposition constant (negative number, unitless) very few assumptions in equation

turnover of litter standing stock Kl = litter fall (g/m^2/y)/litter pool (g/m^2) Requires steady state assumption o Cant measure after a hurricane for months to years because system is in chaos Residence time = 1/K or 1/Kl Take an ecosystem Above ground NPP and below ground NPP o Flux of heterotrophic respiration and leaching of dissolved organic carbon o Heterotrophic respiration is flux of decomposition Carbon exchange (y), time (x)

GPP is graphed and looks like a hill. Goes up, stays flat for a little and then rolls down NEP is shorter hill Graph in book, know relationship between these

Rsoil = R roots + Rnet

Death destruction and rebirth (disturbance and succession) 1/23/2013 12:02:00 PM

Disturbance Important for turnover of resources and nutrients Makes room for new organisms Disturbance severity (x), different levels

top one would be herbivory because its a small disturbance going down they could be fire (sort of severe), agricultural clearing, flooding, mining and war, glaciers and volcanoes (very severe) o more severe has more organic matter removed as an index

succession- changes in plant community composition following disturbance primary- biological colonization and changes in systems that have no live plant material o products of ecosystems are completely removed or buried o no organic matter only a little bit of microbes secondary- changes in organisms and the environment following a disturbance where there are still organisms alive o still life and organic matter (may not be a lot)

primary succession disturbances- glaciers, volcanoes, meteorites (potentially), landslides (small scale), large scale flooding, human construction (roads) stages: o dispersal- getting to the site small seeds, spores generally get dispersed via wind cyanobacteria (fix nitrogen), lichens, algae (form crusts which creates an environment that captures dust and nutrients and stabilizes ground) o colonization

most of the nutrients come from atmospheric deposition early colonizers are often nitrogen fixers combination of free living and symbiotic ones improve environment for later organisms to invade shade, nutrients, water, safe spots (facilitation) facilitation ex: Fern in the tropics makes stem that captures seeds that grow sometimes early colonizers make it hard for later organisms to colonize producing a lot of leaf litter will block seeds from making it into the soil arrested succession- when succession proceeds for a while and then kind of stops Hawaii- Myrica faya, non-native species that colonizes areas Brought by birds, nitrogen fixer, grows well in early succession Creates a lot of organic matter (because its a tree) and rapidly increases pace of primary

succession o Establishment Primary succession environments (in regards to nutrients) Time (x axis), carbon flux (y)- graph is of NPP (green), GPP (red), biomass (blue), decomposition (yellow), NEP (black) o Peaks in the mid because fresh greens and new growth and towards the end its older tissues that dont sequester as much carbon In the beginning, not a lot of nutrients available so it takes time for NPP to gear up At some point, nutrients light and water becomes limiting

in the desert or arctic, it would look similar but over a longer period of time yellow is decomposition- heterotrophic respiration. Lags behind NPP because there needs to be organic material available to break down o at first there is hardly any organic matter NEP is going to be balance between heterotrophic respiration and NPP

Time (x), soil carbon(y) (red), CEC (blue)

slow at beginning because there arent a lot of nutrients or organic matter growth

early succession- good at holding nutrients, retranslocation inputs = outputs mid succession- dont hold onto nutrients as much, more leaky old succession- outputs > inputs If it was primary succession, there would be no plant carbon. After disturbance during early secondary succession, ecosystem looses carbon because wood decomposes and there is a pulse of heterotrophic respiration

Slope of K (decomposition) should be negative Leaves tend to be more nitrogen rich because of RuBP and this makes them easier to break down compared to roots

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Carbon 3

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carbon into an ecosystem is NEE auto respiration fluxes out (from plants and soils) heterotrophic respiration out from soils NEE = GPP Recosystem o Recosystem = Rroot + R heterotroph

something about photosynthesis during the day

cant have negative photosynthesis, not much going on towards the lower limits of photosynthesis maybe some moss or something, not many organisms Tropical forests Growing season January to December- take up light 365 days a year 2% light use efficiency forests are somewhat transparent o amount of sunlight getting to soil from canopy decreases dramatically in a forest have to take albedo into account

seasonality effect day (x), carbon flux (y)

below the line is in the growing season (photosynthesis dominating, sink), above the line respiration dominates (source) tropical forest (red), California (green)

temperature (x), respiration (y)

increased global warming means increased respiration year to year change in photosynthesis (delta GPP) (x), year to year change in respiration (delta Respiration) (y)

decrease in photosynthesis doesnt mean increase in respiration. Things are in balance

flux of carbon largely depends on time since last disturbance after disturbance. Below line is source, above is sink

varies depending on where you are (bog vs chaparral) o bog would take a long time to get to the zero point, chaparral would fluctuate a low because of fires

temperature (x), respiration (y)

lot of variability once the temperature warms, have to consider the other factors o ex: soil moisture, phenology, pulses of respiration from rain (microbes)

ecosystem dynamics
time (x), carbon (y)

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slope is deltaC/deltaT derive deltaC/c = r o ln C = rt take e and C(t) = C(0)e^rt

r (growth rate, y), number (x)

r = rinitial(1-N/K) o as population increases, reduces r initial to get r

concentration of organisms (y), time (x)

C(t) = (K*Co*e^rt)/K + Co*e^rt-1 Law of .69 .69/%growth rate (r) = when the population will double N(t) = N(o) e^rt, divide by N(o) N(t)/N(o) = 2 for doubling time 2 = e^rt, take natural log and .69/r = t r vs. k selection r- small adults, small offspring, many offspring at a time, reproduce early in life k- reproduce late in life, produce few offspring, produce large offspring, large adults pulses

 switches

ex: denitirfication huge flux of N2O

ex: period of dormancy to period of rapid growth (spring), fire, phenology, landslide

Temperature of soil is constant deep into ground black (2cm) blue (5cm) red (16cm) temp (y), time (x)

soil temperature lags with depth

weather physiology (respiration, photosynthesis, transpiration) biogeochemistry (light interception allocation, growth) ecosystem dynamics reproduction (dispersal) recruitment (baby plants succeeding in a new niche) competition (light, nutrients, water) facilitation- ponderosa pine growing under tree because it thrives in shade mortality- 1-2% mortality/year disturbance and succession

Global ecology and remote sensing

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Global ecology: goal: to understand fluxes everywhere all of the time started in 1969 with images of earth from Apollo mission Lieth- broke up earth into grid of latitude and longitude and measured temperature and precipitation Launch of Terra satellite working on MODIS Helps us see things from space Land cover photos from space helps us understand plant functional types Dark and seasonal deciduous forest Incoming PAR- visible sunlight .4-.7 um Use light use efficiency based of others things like temperature and vapor pressure deficit Photosynthesis (y), PAR (x) Sun leaves blue, shade leaves red

1 meter resolution from space satellites clouds make it harder for us to see from space Pros to satellite processing Repeated sampling, able to detect change o Disturbance, phenology, Fire Interannual data

Wavelength (x), reflectance (y)

first bulge at green, dips at blue and red and the rises in the NIR

NDVI = Rnir Rred/Rnir + Rred Reflectance of different types of vegetation and ecosystems GPP = Ipar*Fpar*epsilon

Species effects on ecosystems

specific traits: morphological o ways species show their differences biogeochemical processes o nitrogen fixation C3 vs C4 photosynthesis Deep roots vs. shallow roots Browsers, grazers, pathogens Mangaver indica (mangos)

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Increased soil carbon wherever it was found Species that tends to flush out leaves (drop leaves) and can go through two or three flushes throughout a year

Species good at retranslocating nutrients: Ex: phosphorus. Soil will be phosphorus depleted around the plant o Leading to patches of depletion around a forest Species impact soil temperature Insulating soil at surface because of root mats

Albedo: Fire regimes: Eucalyptus burns easily so when you introduce them into an ecosystem the ecosystem becomes more fire susceptible Diversity in ecosystem function If number of species (biodiversity) is related to ecosystem function then maintaining diversity can be justified for the conservation of ecosystems (so we can get ecosystem services) o This isnt easy (see scale below) Species richness- number of species per unit area

Scale

Scale of ecosystem function and diversity differ o Species are discrete units

o Functioning processes (nutrient cycling, succession) are measured with indexes and a rate of process Continuous and difficult to measure Other difficulties o Complex need to simplify o Detecting changes in function Most species of plants participate in most functions in fundamentally the same way (photosynthesis, nutrient uptake) (microbes however dont do things the same way) Exceptions: nitrogen fixers, C4, CAM More focus on these because they are different to deal with detecting problem- functional groups Deep roots vs non deep roots Nitrogen fixers vs non nitrogen fixers Comparing one group against another Problem is functional groups vary in time and space

How

o Redundancy: Are there redundancy between species? Can we lose some without negative side effects? Appear to be redundant but could be important in some unknown way Theories to look at biodiversity and system function Rate process (y), number of species (x)

black (no change in number) blue (linear, higher NPP with more species) green (asymptotic, at some point species become redundant and adding species doesnt impact function) competition- more species, more they divide up the niche, more resources they use and more competition o adaptations evolve overtime to deal with or avoid competition in niches o life form diversity: different canopy heights lead to different life forms living at the different heights (ground, palm, tall tree, canopy) different phenologies to exploit the environment number of traits doesnt correlate with number of species competitive exclusion- competition leads to decrease in species numbers especially when one is superior o invasive species out compete native intraspecific competition- competition within species can decrease numbers complimentary- more species more resource utilization o facilitation- one species helping another species trees that fix nitrogen and allow other species to come in palm in tropics catch seeds and its a good place for seeds to grow o niche differentiation- sum is greater than whole species working together gives you more ecosystem function than the species alone sampling effect- probability of getting a certain (special) species as you increase number of species in sample o when you have more species, your probability of having a special species is higher special species could be nitrogen fixer, deep rooter, species that has a disproportionate impact on function

experiment design for biodiversity and ecosystem function

Dave Tillman- planted tons (200+) of small plots (1-2 meters) with different species mixtures and followed that over time o Had to weed to maintain species richness Weeding causes disturbances of soil o Results- analyzed data before and after drought Nitrate concentrations in root zone was weakly correlated to number of species More diverse plots, more nitrogen fixers (sampling effect) Experimental microcosms- people create ecosystems in controlled environments o Lets people look at a different range of processes

Look at data Professor Silver posts

Landscape patterns in ecosystem structure and function 1/23/2013 12:02:00 PM

Causes of landscape heterogeneity State factors o Climate: light, temperature, rainfall, soil moisture o Geology: parent material and soils o Topography: Exposure, erosion o Water: rivers, lakes, tidal action (estuaries, coast) Community factors o Founder effects Species influence ecosystem processes May reflect stochastic patterns of colonization o Historical land-use patterns o Disturbance Natural disturbances: fire, hurricane, landslide, diseases Creates patches of different ages and sizes Human activity, logging, agricultural conversion, roads, urbanization, fire suppression/prescribed burning Extensification/intensification Interactions among sources of heterogeneity o Self-Organized Criticality Scaling laws, power laws o Random Dispersal events Sunol Hills: Roles of Topography, Water, Grazing, Fire Lateral Transfer of Water from Ridges to Draws Meets Greater Evaporative Demand of Trees o Need more water in tree area compared to grass area Grass areas drain into tree areas Grazing and Fire Sustain Patterns Patterns in Semi-Arid Systems Alleopathy- plants exute toxic materials to prevent other things from growing close to them Resource islands- water, nutrients Facilitation/competition

o Low vegetation density soil is too hot and dry low rates of colonization and recruitment low vegetation density Shading soil is important because it can account for 3050% water loss o High vegetation density shading lowers soil evaporation higher vegetation density o Higher vegetation density organic matter in soil promotes higher water infiltration higher vegetation density Guano deposition of birds create patches of P rich area where trees can form Boreal forests are a mosaic of time since last disturbance (fires, glaciers) Clumped forests are resilient to fire spread Arctic polygons: formed naturally Freezing and thawing over and over cause shapes Wetland- cold, little evaporation, positive water balance Want to minimize energy needed to get from point a to point b and this affects landscape design and drainage network Land fragmentation Altered environmental conditions Edge effects Increased abundance and incidence of invasive species Changed disturbance regimes Altered species interactions o Pollination Seed dispersal Predation Herbivory competition Genetic deterioration

Fragments have poorer species richness Habitat patches connected by corridors Retained more native plants than isolated patches

Did not promote invasion by exotic species

Landscapes exhibit complex patterns that can be Geometrical and Natural Driven by limitations of resources (water, nutrients), tending to form most in arid and flooded environment Rivers form dendritic networks to minimize the energy used to transfer of water Meandering Rivers, vs Braided Rivers, is a signal of life Fragmentation of Landscapes Reduces Connectivity and Decreases Biodiversity

Ecosystem modeling
Why model Diagnose and understand complex interactions Cross scales o Leaf and soil up to canopy and landscape Look at conditions of the past and future o Ex: CO2, T, N

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Light drives photosynthesis, water Models can consider disturbance, land use change, nutrient, carbon Model parameters Set of equations (y = ax) Driving input PAR, ppt, T Spatial scale Take a landscape and break it up into a grid o How do you put Fine scale information (microbes) into larger scale models Time step Century, or what time scale Validate Field measurements or calculations o Sometimes models are better than field information Models can help understand weaknesses in measurements Hierarchy of models Big leaf model o Assume ecosystem is a huge green leaf Bucket model

Michaelic Menten equation Rate or velocity (y) is function of Vmac * C/ Km+C Square root looking graph Deals with metabolism? 1-D turbis medium

box with dots (leaves) randomly distributed helps model multilayer stuff (canopy)

mean of function doesnt = function of the mean stochastic statistically and randomly predict where a disturbance may occur biophysical model SIB from nasa CLM from NCAR

Biogeochemical model Century BGC- cycling CASA Pnet Biogeography Dissociate plant functional groups and climate

Orchidaee (French) look at ice ages and see how ecosystems move and how species distributions will vary Test models and validate them and they are pretty good models but not as good as they could be

Global Warming

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Policy and science People dont want to change and its going to cost us a lot in the long run Internalize the externalities Fallacious logic ClimateGate scandal (Al Gore) Oil lobby and organizations Global temperature record Fact CO2 is increasing (Mauna Loa) o Oscillating increasing since 1960 Dips in northern hemisphere summer because of photosynthesis Raises in winter because of respiration CO2 absorbs infrared radiation in open IR windows o Radiation vibrates the CO2 molecules and they re admit it randomly o Peak absorption at 4.25 um, 7.2 um, 14.99 um The window where radiation escapes is the absorption area of CO2 so when we add CO2, we make the window for escaping radiation smaller From 1880 to 1960, slight increase After 1980, pretty big increase

Evidence of global warming Sea level rise Tree rings Etc..

Stable isotopes in ice, a proxy for temperature C4 grasses outcompete C3 under low CO2 conditions so they dominate between ice ages

Natural

solar forcing of climate variability Eccentricity Obliquity (tilt) Axial precession

Tree rings- grow better in warm wet years Sea level rise- thermal expansion and melting ice Temperature lags CO2 Feedback: climate warms ice melts CO2 solubility in ocean decreases CO2 rises climate warms More Particles Reduce Daytime Warming from Sun More Greenhouse Gases Reduce Nightime Cooling leafing and flowering earlier in the year

Climate Change

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Things scale from cell to leaf to organism to organisms to ecosystem to landscape to globe 3 main greenhouse gases CO2 CH4 Nitrous oxide (N2O) Other greenhouse gases CFCs, SF6, water vapor, O3, HFC Radiating forcing = energy change per unit area of globe measured at the top of the atmosphere Amount of energy captured per unit area per unit time (for greenhouse gases, longwave from earth) Positive is warming, negative is cooling Global warming potential Relative index of heat trapped by greenhouse gases o Relative because all scaled to CO2 = 1 o GWP (x) = something + H = time horizon ax = radiative efficiency due to increase in abundance x(t) = time dependent decay r = reference gas (CO2) GWP of CO2 = 1 GWP of CH4 over 20 years = 72 o Over 100 years = 25 o Methane decreases because it decays in the atmosphere where N2O doesnt decay as quickly GWP N2O over 20 years = 289 o Over 100 years = 298

Carbon cycle Photosynthesis in, respiration out o Keeling plot (mauna loa) lets us know this is happening

o More wiggle in northern hemisphere because there is more land area and more photosynthesis and respiration happening Carbon in atmosphere = 760 Pg (10^15) o In plants = 650 Pg Turns over every 11 years (average on global scale) o Oceans = 38,000 Pg Surface water = 1,000 Pg Stored as inorganic carbonate Marine biota = 2 Pg but cycle as much carbon as terrestrial vegetation Turnover is 2-3 weeks Biological pump- carbon is fixed and pushed to bottom of the ocean More CO2 added to atmosphere, more CO2 entering the water and this changes acidity of water Impacts shellfish ability to make shells Ocean cant hold as much CO2 under acidic conditions compared to neutral or basic conditions Solubility pump- linked to upwelling

CO2 rich waters get transported down in northern latitudes (thermohaline circulation?) Slower cycle than biological pump o Soils = 1,500-2,500 Pg C Turnover around 25 years 3 pools fast- turnover in months to years intermediate = years to decades slow = decades to millennia can manage the carbon in soil and possibly can push it into the slow pool to store the carbon o Rocks and sediments = 10^7 Pg C Turnover slowly naturally = millions of years But humans are mining fossil fuels and oxidizing it and making that turnover time faster CO2 emissions are 77% greenhouse gas emissions o Sources fossil fuel combustion (70%)

Biomass burning and land use change (17%) Deforestation

Reducing CO2 emissions wont be enough to change climate change Hope Reduce emissions Grow plants that sequester carbon Pushing carbon into rocks (CO2 injection) o Risks with earthquake faults and releases of high concentrations of CO2 Methane (CH4) 20% current global warming natural sources: o wetlands, termites, tropical forests o come from archea (not bacteria) and produce methane under anaerobic conditions human sources o landfills (anaerobic conditions with a lot of organic matter) o enteric fermentation- cows, other live stock produce methane in their stomachs when they break down food (fiber) o rice agriculture- flooded, anaerobic conditions residence time about 10 years (CO2 is hundreds of years)

N2O

6% global warming residence time hundred years Natural sources o Tropical forests (cycling nitrogen at fast rate, denitrification) Human sources o Cow manure (feedlots, dairies) 2.1Tg o Fossil fuel industry 1.3 Tg o Fertilizer and N fixing crops

Ecosystems and environmental change 1/23/2013 12:02:00 PM

More N is fixed Synthetically than Naturally > 50% of accessible fresh water is being depleted ~ 20% Marine fisheries are over exploited Extinction rates of species have accelerated Were experiencing the 6th Great Extinction Period over Earths History, the 1st caused internally. Land Use Change and Extinction Opens Niches for Invasive Species Attributes of Global Environmental Change- Anthropocene Changing things much faster than previously o The time scale of this great extinction is so much smaller than in the past Change is non-linear, hysteretic Change is conditional on antecedent conditions o Down-regulation- photosynthesis response to CO2 will depend on conditions in which they are grown Change is Multi-Factorial (CO2, N, ppt, T, O3, age, invasive species, disturbance)

Ways to study ecological change Mix of observations, models, manipulations, and relate those to past environments (paleo records) CO2 tunnels Help you see how photosynthesis changes with different species with different CO2 levels o From pre industrial levels to ice age levels to projected levels In summer in northern hemisphere, ITCZ shifts slightly northward and so tropical areas are on the edge of the region and wont get as much water Timing of rainfall may be as important as amounts added/subtracted Look at monthly climate drivers over annual (or at least look at both)

 In Principle, Global Warming Should Accelerate and Water Cycle Increase Evaporation Increase Cloud Cover Increase Precipitation Reduce Solar Radiation Negative feedback on Evaporation

Results from global change experiments

Photosynthesis vs CO2 Linear/curved at the top relationship Ci = .7Ca Higher and higher CO2, saturate plants

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Down-Regulation of Photosynthesis based on CO2 Growth Environment Dont achieve as high of rates as you would otherwise Graph on powerpoint As CO2 levels increase, stomatal conductance decreases o Dont have to open up stomata as much More efficient water use under high CO2 conditions

Increased CO2 will increase yield and biomass but less than the theoretical amount Down-regulation Elevated CO2: Direct and Indirect Effects Growth goes up o Compound interest effect- small plants will grow faster with more CO2 but bigger plants grow faster than smaller ones Mineralization decreases Respiration increases

At high CO2, insects eat more leaves to get proper amount of nitrogen Herbivory increases because leaves have less nitrogen Possible shifts with warming Increase in mean (hotter averages) Increase in variance (more extreme weather) Increase in mean and variance

Temperature acclimation Bad effects if the temperature changes over a short time scale because plants dont have time to acclimate so more plants die o Models assume big jump in temperature and this may not be accurate

o Photosynthesis and acclimate in a good way o Respiration will down-regulate for warm adapted places Cold places will up-regulate respiration rates Plants can handle a few days of heat stress but repeated days of high temperatures will cause a drastic increase in mortality Can tolerate some stress but eventually give in Phenology is the study of periodic plant and animal life cycle events and how these are influenced by seasonal and interannual variations in climate. Plants will flower earlier with global warming o What happens when there is an unsynchronized pattern between flowers and pollinators

Increasing trends of growing season since 1980s 177 days to about 187 days disturbance and land use change Fires Deforestation/Logging Urbanization Desertification Woody Encroachment Afforestation/Reforestation Storms Invasive Species Soil Degradation

Deforestation leads to net loss of carbon No trends really but additions to atmosphere

Ecosystem management and climate treaties

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Weve covered- ecosystem structure and function, water and nutrient cycles, light, chemistry and physics of plant soil atmosphere interactions, role of biota Scaling from small to globe Humans as part of ecosystems Consumers Use raw materials, transforming and sometimes recycling them o Some stuff cant be broken down- no oxygen, no light, molecules that are hard to break down (microbes break down the easy stuff first), wont be broken down on human time scales o Everything is recycled on some time scale Tipping point- passing a point where you no longer can recover Ecosystem scientists contributing to policy REDD- reduced emissions from deforestation and degradation o Financial market to incentivize reducing greenhouse gas emissions from deforestation o Intact forest Benefits- biodiversity conservation, poverty reduction, clean water, reduced erosion o Emissions from deforestation and degradation account for 1220% of warming trends o Intact forests sequester carbon and can store them for a long period of time When you remove forest and burn them and increases microbial respiration (both of which release CO2) then less CO2 is sequestered and more in the atmosphere o Challenges Deforestation? Clear cutting? Selective harvesting? How much? Definitions are unclear How large does the area need to be to be a REDD forest? Primary forests? Secondary forests? Plantations? Forest management? Thinning can increase growth

Do we thin if we want to sequester carbon? Fire? Degredation- how do you quantify it? Reforestation- turning old forest land into forests again afforestation- taking non forest land (ag land) and making it forest

additionality- carbon projects, need to establish a gain in carbon above baseline conditions time (x), carbon (y) in a system that is degrading

normal in red. Blue is what you would have to show for a REDD project o you have to show, that even though it is declining, it is still retaining more carbon than normal

leakage- carbon offset projects that result in carbon loss from another environment REDD project- say you want to build a forest on a cow pasture, well those cows are going to go elsewhere where a forest will be cut and burned and thats the leaked carbon Permanence- 100 years (impractical period of time) How do you fallow carbon for 100 years- only way through models Maybe the time period should be 20 years GRASSLANDS: 30% global land surface, 30-50% US surface is range lands

grasslands store 1/3 of worlds soil carbon more water loss than more water input, have to allocate resources in soil and put lots of roots in soil and store carbon relatively stable lands manage the lands by adding manure or compost and led to build up of carbon in ecosystems

Final review
general concepts on decomposition

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sampling effect- applied to concepts of biodiversity ecosystem function probably of finding nitrogen fixing species in a group of species is higher if you have more species decomposition litter chemistry: o nutrient content of litter look at N or P o amount of lignin- difficult to break down o tanins, polyphenolics- distasteful or toxic to organisms secondary compounds for defense or byproducts o cellulose, hemi-cellulose o stoichiometry- elemental ratios- C:N, N:P, Al:Ca primary succession carbon flux (g/m^2/y) (y), time (x) npp (blue), gpp (red) (50% of carbon take up goes out as autotrophic respiration), heterotrophic respiration (green), NEP (yellow)

if it was secondary succession- steeper carbon loss (heterotrophic respiration) (dark green above) NPP would start higher, drop to almost zero and then peak above the other NPP (light blue) NEP (orange)- lots of carbon because it grows like mad after secondary succession o NEP should be negative in the initial stages but I couldnt redraw it in the graph above o Evidence grasslands are in the decline NEP wise and are loosing carbon o Over time, NEP

primary succession- start from scratch, very gradual increase secondary succession- start higher and then there is a disturbance and there is a dip additive models, multiplicative models how its used, why its used how is remote sensing a tool used for understanding ecosystems biodiversity and ecosystem function memorize global warming potentials whendee OH-1-2, Dennis-after 4 less nitrogen to carbon ratio with increase CO2 so herbivory increases Increased CO2 decreased photorespiration

o increased leaf temperature o increased water use efficiency o decreased transpiration increased growth stimulation down regulation photosynthesis from nutrient limitation increased nitrogen limitation (C:N) ration increased herbivory nitrogen cycle increases o plants taking up N faster o insects eating more and they secrete nitrogen in feces and itll cycle faster

tolerance- previous characteristics allow plants to be more tolerant to change waxy leaves biodiversity doesnt matter so much, just that each function is being served in an ecosystem spatial resolution- see bigger picture, less specific spectral- more specific, smaller picture shifts (ball in cup model, dormancy to rapid growth like in the spring), lag (recovery rate, temp and CO2, soil temperature with depth) , pulse (respiration after rain, denitfrication and N20)

things going to be on the test primary and secondary succession o graphs too carbon pools PgC = 10^15, Tg = 10^12 Affects of higher CO2 on plants Remote sensing- pros and cons o NDVI vs. LAI

o Cons- clouds, precision (tradeoff between resolution and scale) o Pros- large scale, easier than field measurements, interannual data, detect disturbance GWP (CO2, CH4, N20) o Atmospheric window of CO2 Ecosystem response to environmental change (on stevens sheet) o Acclimation, movement, adaptation, extinction, tolerance, avoidance o TAAAME How to calculate decay o Flux/pool Residence time = 1/k o Factors influencing it NDVI equation Landscape heterogeneity Hysteresis- plants already acclimated to warm temperatures wont change respiration rates as much with increasing temperature as cold adapted plants o Dependence on past conditions