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    Lecture 1: Introduction to

    Aquatic Environments

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    Aquatic Environments

    Aquatic environments

    Oceans

    Coastlines/Estuaries

    Streams Lakes

    Wetlands: bogs and fens

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    Freshwater Ecosystems

    Only 3% of the worlds water is fresh, and

    99% of this is either frozen in glaciers and

    pack ice or is buried in aquifers. The

    remainder is found in lakes, ponds, rivers

    and streams.

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    Limnology: The study of freshwater

    ecosystems, with anemphasis on

    understanding thephysical, chemical andbiological processes thatare important instructuring andmaintaining freshwatersystems.

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    Science of Limnology

    embraces: Lakes (Lentic waters)

    Rivers (Lotic waters)

    EstuariesOther microhabitats such as springs and

    streams

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    Geology

    By what processes arelakes and rivers formed?

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    Geology

    Physics

    What are theimportantphysical properties of

    water and watermovement?

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    Geology

    Physics Chemistry

    How do thechemical propertiesof water and the

    nutrients it containsaffect whichorganisms can livethere?

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    Chemistry

    Physics Geology

    Biology

    How do organismsinteract with each otherand their aquatic

    environment?

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    Geology

    Origin of Lake Basins

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    Physics

    Physics and ionic content of water

    Light and Heat in Lakes

    Distribution of heat in Lakes

    Water movement in Lakes

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    Chemistry

    Oxygen, carbon, alkalinity and pH

    Nutrient Dynamics (P, N, Si)

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    Biology

    OrganismsPhytoplanktonZooplanktonAquatic Insects

    Fish

    Case StudiesAcid DepositionPaleolimnologyExotic Species

    Cultural EutrophicationKiller Lakes

    ConceptsPopulation dynamicsCompetitionPredationDiseaseLife-histories

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    Freshwater is a major limiting resource

    Why study limnology?

    Lakes and ponds are great laboratories to studyecological and evolutionary processes

    Water in the Biosphere: Volume (thousands of km3) % of totalOceans 1,370,000 97.61Polar ice, glaciers 29,000 2.08Ground Water 4067 0.295Freshwater lakes 126 0.009Saline lakes 104 0.008

    Soil and subsoil moisture 67 0.005Atmospheric water vapor 14 0.0009Rivers 1.2 0.00009

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    UN Report(2002)

    suggeststhat by2025, two-thirds of

    the worldspopulationmay facewater

    shortages

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    Brief History of Limnology

    1600sLakes in Germany and the Netherlands are categorized

    based on the presence/absence of stream inflows

    van Leeuwenhoek describes algae and rotifers from aDutch lake

    1700sH.B. de Saussure determines that deep waters in some

    Swiss lakes are much colder than surface waters

    Early studies of lakes flourished in the glaciated regions of Europe and North America

    1800

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    1800s

    L. Agassiz uses a tin cup to measure turbidity in Lake Superior

    Acid Rain is linked to coal burning in England (1852)

    Development of the Secchi disk

    1900s

    First textbook of Limnology published (1901)

    Linking of sewage inputs to algal blooms (1910)

    First whole-lake manipulationwithout control lake (1938)

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    What Now??

    Shift from observation to experimentation Species interactions and food web dynamics

    Ecosystem ecology

    Acid rainCultural eutrophication

    Exotic Species

    Paleolimnology

    Genomics

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    Francois Alphonse Forel

    1841-1912, Swiss

    Defined limnology as oceanographyof lakes

    1st textbook of Limnology published in 1901 Handbuchder Seenkunde. Allegemeine Limnologie, based on 30years of research on Lac Lman (Lake Geneva).

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    Stephen Alfred Forbes

    1844 - 1930, American

    Chief of the Illinois Natural History Survey

    Published Lake as a Microcosm in 1887

    concept of a lake as a superorganism

    www.inhs.uiuc.edu/cae/ ltrm/station.html

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    August Thienemann

    1882-1960, German

    With C.L. Naumann started the International

    Association for Theoretical and appliedLimnology (SIL) in 1922

    Influential ideas on nutrient cycling and foodweb structure

    http://www.limnology.org/news/25/thienemann.jpg

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    Edward A. Birge and Chancey Juday

    1851-1950 (Birge), 1871-1944 (Juday)

    Founded Wisconsin School ofLimnology

    Comparative limnological surveys ofhundreds of lakes

    Major weaknesses of Birge & Juday andcontemporarieslack of a clearlyarticulated hypothesis, few statistics,lack of replication and controls

    State Historical Society of Wisconsin

    http://www.library.wisc.edu/etext/WIReader/Images/WER0032.html

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    G. EvelynHutchinson

    1903-1991, professor at Yale.

    Never earned a PhD.

    Published four volume Treatise of

    Limnology(1957-1993)

    Responsible for the introduction

    of numerous ecological andlimnological concepts

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    S f

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    American Society of

    Limnology and

    Oceanographyhttp://aslo.org/

    International

    Association ofTheoretical andApplied Limnology

    http://www.limnology.org/

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    Lakes vs. Reservoirs

    Lakes Natural Reservoirs Man made

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    Streams

    Open systems, constant input of water andnutrients

    Precipitation flows into streams via 2 routes: Overland flow through surface runoff

    Infiltrating soil surface, then flowingunderground and into streams as

    groundwater

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    Stream Classification

    Based on flow

    Permanent: constant above-ground flow

    year-round

    Intermittent/Ephemeral: flow aboveground

    for parts of the year, not all (temporal)

    Interrupted: flow aboveground for parts of

    the stream, not all (spatial)

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    Stream Classification

    Based on order

    1st: no streams

    flowing into it

    2nd: two 1st-order

    streams joining

    3rd: two 2nd-order

    streams joining

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    Physical features

    Channel shape and pattern Changes with age

    Pools and riffles

    Velocities, microclimate differ

    Rivers age

    Young: little meanders, small floodplain,

    fast velocity, V cross-sectional profile

    Mature: many meanders, slower velocity,

    oxbows form, U profile

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    Watershed - the area that a stream drains,

    a.k.a, drainage basin, or catchment area

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    River Continuum Hypothesis

    Predictable structure of river (physical

    features, dominant organisms) from

    upstream headwaters to downstream

    high-order stream

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    Headwaters/upstream:

    Riffles/rapids predominant

    Heavily shaded by riparian vegetation

    Energy importedallochthonous material

    High diversity of benthic fauna

    Downstream

    Pools of slow water dominant

    Only banks shaded by riparian vegetation

    Autochthonous input

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    Lakes

    May be created by a variety of geologic and

    climatic events:

    Movement of tectonic plates

    Volcanic eruptions

    Landslides

    Fluvial processes

    Glaciation

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    Lake Zonation

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    Lake Zonation

    Littoral zone: shallow (

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    Vegetation Zonation

    Shrub &Trees

    Mixe

    dherbaceous

    Grassstage

    Shallow

    emerg.

    Deepwater

    emergents

    Floa

    tingplants

    Submerged

    plan

    ts

    Open waterphytoplankton

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    Lake Stratification

    Different zones or layers due to water

    temperature and water density Epilimnion: layer closest to surface of water; warmed

    by the sun, least dense

    Metalimnion: middle layer with thermocline;

    transitional layer

    Hypolimnion: deepest layer, generally coldest;

    sunlight does not penetrate

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    Lake Stratification

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    Seasonal Changes

    Summer:

    Warm temperatures,

    long days

    Obvious vertical

    stratification

    Epilimnion saturated

    with oxygen Hypolimnion anoxic

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    Fall:

    Air temperatures cool, surface water cools

    fastest and sinks to the bottom

    Complete lake turnover

    Lake no longer stratified

    Lake eventually becomes a uniform 4C

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    Winter:

    Surface water cooler

    than rest of lake

    water

    Ice prevents mixing

    Winter stratification,0C at surface, 4C

    at bottom

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    Spring:

    Ice melts, water surface hits 4C and

    again begins to sink

    Spring turnover, process repeats itself

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    Roach Lake in March

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    Roach Lake in August

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    Nutrients

    Temperature not the only stratified element

    of a lake

    Oxygen: highest concentration near surface

    (photosynthesis)

    Nitrogen: NO3- at surface, NH4

    + at benthos

    Sulfur: SO4-2 at surface, H2S at benthos

    Iron: Fe+3 at surface, Fe+2 at benthos

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    Marsh (Eutrophic)

    Bog (Dystrophic)

    Oligotrophic

    Lake

    Mesotrophic to Eutrophic Lake

    Terrestrial

    Sphagnum

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    Crampton Lake (oligotrophic)

    Bro n Lake

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    Brown Lake

    (mesotrophic - eutrophic)

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    Northgate Bog (dystrophic)

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    Ziesnis Bog (dystrophic)

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    Wetlands: technical definition

    Vegetation presence of hydrophytic (water-loving, flood-

    tolerant) plants

    Soils presence of hydric (flooded, reduced) soils

    Hydrology water table at or near the surface for part of the

    growing season

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    Wetland history

    Historically, wetlands have been drained to: Provide land for agricultural purposes

    Reduce the incidence of mosquito-bornediseases, like malaria, yellow fever

    Wetlands now recognized as havingcommercial, aesthetic, and ecologicalvalue

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    Why are wetlands important?

    Storm and floodwater

    storage

    Improved water quality:

    filtration Rare or endangered

    plants and animals

    Waterfowl nursery

    grounds

    Migration stop-overs

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    Wetland examples

    Marshes

    Swamps

    Glades

    Bogs

    Fens

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    Bogs

    Acidic (pH < 4.1)

    Nutrient-poor soils

    Ombrotrophic:

    precipitation-fed system

    Dominant vegetation:Sphagnummoss,Vaccinium(cranberriesand blueberries), and

    other low-lying species

    Slightly less acidic (pH4.1-6.0)

    Soil more nutrient-rich

    Minerotrophic:

    groundwater-fed system

    Dominant vegetation:

    sedges, rushes, and

    grasses

    Fens

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    Black spruceSwamp alder

    Tamarack Leatherleaf

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    Cotton grassPitcher plant

    Sphagnummoss

    Cattail

    Sundew

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    Long Lake

    Peter Lake

    Paul Lake

    Nutrients added/No Piscivores

    Nutrients added/Piscivores

    No Piscivores

    Piscivores

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    Physical Aspects

    Light, heat, waves, and currents can

    produce physically distinct zones that can

    vary by day and season.

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    Chemical Structure

    Greatest cause of altered water chemistryin natural waters is now pollution fromhuman waste, agriculture and industry.

    Without proper control, the increasedchemical loading can lead toeutrophication of lakes and reservoirs.

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    Chemical Structure

    Chemical composition of natural aquatic

    ecosystems are dependent on rainfall,

    erosion, evaporation and sedimentation.

    Water contains a vast array of inorganicand organic compounds which are present

    as dissolved solids and gases.

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    Biological Structure

    Viruses role usually associated with

    public health problems (i.e.: hepatitis can

    be transmitted by ingestion of natural

    waters contaminated with the feces ofinfected people

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    Biological Structure

    Bacteria unicellular organisms

    responsible for recycling of organic and

    inorganic materials.

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    Biological Structure

    Fungi participate in the decomposition

    and recycling of vegetable and animal

    matter.

    Phytoplankton - algae may be unicellular,colonial or filamentous in form

    (Pediastrum)

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    Biological Structure

    Macrophytes aquatic plants may

    dominate shallow lakes and streams.

    Zooplankton components are

    protozoans, rotifers and crustaceans.

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    Biological Structure

    Zoobenthos bottom feeders; includes

    insect larvae, crustaceans and mollusks.

    Fish

    9-1

    O H bit t

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    There are two major marine provinces: thebenthic (bottom) and the pelagic (water

    column).

    The benthic environment is divided by depth intothe: Intertidal zone, Sublittoral zone, Bathyal

    zone, Abyssal zone, and the Hadal zone

    The pelagic environment is divided into the

    Neritic Zone and the Oceanic Zone

    Ocean Habitats

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    The five major kingdoms in the ocean are:9-2

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    The five major kingdoms in the ocean are:

    Monera, Protista, Fungi, Metaphyta and

    Metazoa. Monera are the bacteria and blue-green algae.

    Protista are single-celled organisms with a

    nucleus. Fungi are abundant in the intertidal zone and are

    important in decomposition.

    Metaphyta are the plants that grow attached tothe sea floor.

    Metazoa include all multicellular animals in the

    ocean.

    Marine organisms can also

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    be classified by lifestyle.

    Plankton are the organisms which float in the water andhave no ability to propel themselves against a current.

    They can be divided into phytoplankton (plants) and

    zooplankton (animals).

    Nekton are active swimmers and include marine fish,

    reptiles, mammals, birds and others.

    Benthos are the organisms which live on the bottom

    (epifauna) or within the bottom sediments (infauna).

    Some organisms cross from one lifestyle to another

    during their life, being pelagic early in life and benthoniclater.

    E i t l f t i9-4

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    Environmental factors in

    the marine environment include:

    Temperature

    Salinity

    Pressure

    Nutrients Dissolved gases

    Currents

    Light

    Suspended sediments

    Substrate (bottom material) River inflow

    Tides and waves

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    Ecosystem is the total environment including

    the biota (all living organisms) and non-living

    physical and chemical aspects. Temperature can control distribution, degree of

    activity and reproduction of an organism.

    Salinity can control the distribution of organisms

    and force them to migrate in response tochanges in salinity.

    Hydrostatic pressure is the pressures exerted

    by a column of water surrounding an organism.

    More than 90% of marine plants are algae

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    More than 90% of marine plants are algae

    and most are unicellular and microscopic.

    To photosynthesize (produce organic material

    from inorganic matter and sunlight) plants must

    remain within the photic zone. Diatoms are single cells enclosed in a siliceous

    frustrule (shell) that is shaped as a pillbox.

    Dinoflagellates are single cells with two whip-like

    tails (flagella).

    Zooplankton include the

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    Zooplankton include the

    copepods and foraminifera.

    Copepods are small herbivores (plant-

    eating organisms) that filter diatoms fromthe water.

    Foraminifera are single-celled,

    microscopic organisms which build shells

    of calcium carbonate.

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    Marine Ecosystems

    Ecosystems (or ecological systems) are systems ofcommunities in a large geographical area.

    In order for an ecosystem to be successful, fourthings are required:

    An energy source

    Organisms capable of capturing this energy in the

    form of organic molecules

    .Organic material must be available to all other

    organisms

    Cycling of nutrients must occur between the abiotic

    and biotic portions of the system

    A ti E l A li ti

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    Aquatic Ecology Application

    Compliance with water quality standardsand permits

    Fish population and bioaccumulation

    studies

    A ti E l A li ti

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    Aquatic Ecology Application

    Biomonitoring studies for toxicity ofeffluent

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    Lecture 2: Major Ions and Nutrients

    Water chemistry:

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    y

    Gases, major ions & nutrients

    Gases

    Oxygen (O2)

    Carbon dioxide (CO2)

    Nitrogen (N2)

    Hydrogen sulfide (H2S)

    Major ions (anions and cations)

    Nutrients (phosphorus and nitrogen)

    W t h i t

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    Water chemistry: gases

    What are the ecologically most importantgases ?

    O2

    CO2

    N2

    H2S

    Gas sol bilit

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    Gas solubility

    The maximum amount of gas that can be dissolved inwater (100% saturation) is determined by temperature,

    dissolved ion concentration, and elevation

    solubility decreases with temperature

    warm beer goes flat

    solubility decreases with higher dissolved ion content

    (TDS, salinity)

    DO saturation is lower in saltwater than freshwater

    (for the same temperature, solids drive out gases)

    Water chemistry: O

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    Water chemistry: O2

    ~ 21% of air

    Very soluble (DO)

    Highly reactive and concentration is dynamic

    Involved in metabolic energy transfers

    Major regulator of metabolism (oxic-anoxic)

    Aerobes (fish) vs anaerobes (no-fish, no zoops)

    Types of fish

    Salmonids = high DO (also coldwater because ofDO)

    Sunfish, carp, catfish = low DO (also warmwater)

    Major sources of O2

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    j 2 Sources

    Photosynthesis (phytoplankton, periphyton,macrophytes)

    Air from wind mixing

    Inflows tributaries may have higher or lower DO

    groundwater may have higher or lower DO

    Diffusion (epilimnion to hypolimnion and viceversa)

    Major sinks of O

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    Major sinks of O2

    Sinks Respiration

    bacteria, plants, animals; water and sediments

    Diffusion to sediment respiration Outflow (tributary or groundwater)

    Gases: N

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    Gases: N2

    ~ 78% of air Concentrations in water usually saturated because it is

    nearly inert

    Supersaturation (>100 %) can occur in reservoirtailwaters from high turbulence

    May be toxic to fish (they get the bends) N2 -fixing bacteria and cyanobacteria (blue-green

    algae) convert it to bio-available NH4+

    Denitrifying heterotrophic bacteria convert NO3- to N2

    and/orN2O under anoxic conditions

    Gases: CO2

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    2

    Only about 0.035% of air (~ 350 ppm)

    Concentration in H2O higher than expected based onlow atmospheric partial pressure because of its high

    solubilityGas

    (at 10oC)

    Concentration @

    1 atm (mg/L)

    Concentration @

    normal pressure (mg/L)

    N2 23.3 18.2

    O2 55.0 11.3

    CO2 2319 0.81

    How long does your soda pop fizz after shaking it?

    CO reactions in water

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    CO2 reactions in water

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    Inorganic - C equilibria

    Note 100% CO2 for pH< ~ 4.5; 100% bicarbonate for pH ~ 8

    and 100% carbonate for pH > ~12

    H2CO3 HCO3- CO3-2

    pH

    Inorganic - C: Major sources andsinks

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    sinksSources:

    Atmospheric CO2

    Respiration and other aerobic and anaerobic

    decomposition pathways in the water and sediments

    Groundwater from soil decomposition products

    Groundwater from volcanic seeps

    Sinks:

    pH dependent conversions to bicarbonate and carbonate

    Precipitation of CaCO3 and MgCO3 at high pH

    Photosynthesis

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    Soda pop chemistry

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    www.saddleback.cc.ca.us/faculty/thuntley/ms20/seawaterprops2/sld013.htm

    Soda pop chemistry

    CO2 and the inorganic carbon

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    2

    system

    Carbon dioxide diffuses from the atmosphereinto water bodies and can then be incorporated

    into plant and animal tissue

    It is also recycled within the water with some

    being tied up in sediments and some ultimately

    diffusing back into the atmosphere

    Fixed carbon also enter the water as

    allocthonous particulate and dissolved material

    CO2 and the inorganic carbon

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    2

    system - 2

    Alkalinity, acid neutralizing capacity(ANC), acidity, carbon dioxide (CO2), pH,

    total inorganic carbon, and hardness are

    all related and are part of the inorganiccarbon complex

    CO2 chemistry: Alkalinity

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    2

    Alkalinity the ability of water to neutralize acid; a measure of buffering

    capacity or acid neutralizing capacity (ANC)

    Total Alkalinity (AlkT) = [HCO3-] + 2[CO3

    2-] +[OH-] - [H+]

    Typically measured by titration with a strong acid. The units are in mg

    CaCO3/L for reasons relevant to drinking water treatment

    Can be used to estimate the DIC (dissolved inorganic carbon)

    concentration if the [OH-]

    Conversely, direct measurements of DIC by infrared analysis or gas

    chromatography, together with pH and the carbon fractionation

    schematic can be used to estimate alkalinity

    Alkalinity and water treatment

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    Advanced wastewater treatment (domestic sewage) Phosphorus nutrient removal by adding lime (Ca(OH) 2) or

    calcium carbonate (CaCO3)

    As pH increases >9, precipitates adsorbed PO4-3

    Settle and filter the effluent to obtain 90-95% removal

    Used for particle (TSS) removal also

    Drinking water treatment

    For TSS removal prior to disinfection

    Acid-rain mitigation to whole lakes Lime or limestone added as powdered slurry to increase

    impacted lake pH

    Also broadcast aerially to alkalize entire watersheds

    CO2 chemistry: Hardness

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    Hardness - the total concentration of multi-valent (i.e. >2) cations

    Ca+2 + Mg+2 + Fe +3 (when oxic) + Mn+4 (when oxic); all other multivalent

    cations are typically considered to be negligible

    Sources-

    Minerals such as limestone (Ca and Mg) and gypsum (Ca)

    Water softeners and other water treatment processes such asreverse osmosis and ion exchange

    Evaporation can increase hardness concentration

    Drinking water effects (no real health effects)

    Soap scums and water spots on glasses and tableware

    Deposits (scaling) can cause clogging problems in pipes, boilersand cooling towers

    Water chemistry Major ions

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    Water chemistry Major ionsIon balance for typical fresh water

    Anions Percent Cations Percent

    HCO32- 74 Ca2+ 63

    SO42- 16 Mg2+ 17

    C1- 10 Na+ 15

    K+ 4SiO2 < 1

    Note: plant nutrients such as nitrate, ammonium and phosphate that cancause algae and weed overgrowth usually occur at 10s or 100s of parts-

    per-billion and along with other essential micronutrients usually represent

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    Nutrients phosphorus

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    Nutrients phosphorus

    Essential for plant growth Usually the most limitingnutrient in lakes

    Derives from phosphatic rock abiotic, unlike nitrogen

    No gas phase, but can come from atmosphere as

    fugitive dust

    Adsorbs to soils

    Naturally immobile unless soil is eroded or excess

    fertilizer is applied

    Phosphorus moves with sediments

    Nutrients phosphorus

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    Nutrients phosphorus

    Not toxic

    Algae have physical adaptations to acquire

    phosphorus

    High affinity (low k) Alkaline phosphatase activity

    Storage

    Luxury uptake Single redox state

    Phosphorus cycle is closely linked to the iron

    (Fe) cycle

    Phosphorus basic properties

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    Phosphorus basic properties

    No redox or respiration reactions directlyinvolved (organisms are not generating energyfrom P chemistry)

    PO43 highly adsorptive to cationic sites (Al+3,

    Fe+3

    , Ca+2

    ) Concentration strongly affected by iron redoxreactions

    Ferric (+3) insoluble floc, adsorb PO43

    Ferrous (+2) soluble so PO43

    becomessoluble, too, unless Fe2+ reacts with sulfide,causing FeS to precipitate

    Phosphorus levels in the

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    environment

    Major factors affecting phosphorus levels,cycling, and impacts on water quality

    include:

    Soil properties Land use and disturbance

    Transport associated with runoff

    Where does phosphorus come

    f ?

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    from?

    Phosphorus external sources

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    Phosphorus external sources

    Nonpoint sources Watershed discharge from tributaries

    Atmospheric deposition

    Point sources Wastewater

    Industrial discharges

    Phosphorus nonpoint sources

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    Phosphorus nonpoint sources

    Watershed discharges from tributaries Strongly tied to erosion (land use

    management)

    Stormwater runoff (urban and rural)

    Agricultural and feedlot runoff

    Atmospheric deposition

    Often an issue in more pristine areas

    Arises from dust, soil particles, waterfowl On-site domestic sewage

    Phosphorus point sources

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    Phosphorus point sources

    Wastewater Municipal treated wastewater

    Combined sewer overflows (CSOs)

    Sanitary sewer overflows (SSOs)

    Industrial discharges

    Phosphorus internal sources

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    Phosphorus internal sources

    Mixing from anoxic bottom waters with high phosphatelevels is closely tied to iron redox reactions

    O2 > 1 mg/L Insoluble ferric (+3) salts form that

    precipitate and settle out, adsorbing PO4-3

    O2 < 1 mg/L (anoxic) ferric ion reduced to solubleferrous ion (Fe+2) allowing sediment phosphate to

    diffuse up into the water

    Wind mixing (storms and fall de-stratification) can re-

    inject high P water to the surface, causing algal blooms

    Phosphorus Lake budget

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    Phosphorus Lake budget

    Nutrients phosphorus cycle

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    Nutrients phosphorus cycle

    Major pools andsources of P inlakes Natural inputs are

    mostly associated with

    particles P as ortho-p, Pi, soluble

    reactive P

    Wastewater is mostlydissolved phosphate

    P is rapidly removedfrom solution by algal-bacterial uptake or byadsorption to sediments

    Phosphorus cycling major

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    sources Sewage

    Dissolved

    Tributaries and

    deposition

    Particulate

    Erosion

    Particulate

    Sediments

    Particulate and

    dissolved

    Phosphorus cycling internal

    li

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    recycling

    Rapid PO4-3recycling

    Bacterial uptake

    Algal uptake

    Adsorption to

    particles

    Detritus

    mineralization

    Zooplankton

    excretion

    Fish excretion

    Phosphorus cycle major

    t f ti

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    transformations

    Modified from Horne and

    Goldman, 1994.

    Limnology. McGraw Hill.

    The whole

    phosphoruscycle

    Nitrogen basic properties

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    Nitrogen basic properties

    Nitrogen is relatively scarce in somewatersheds and therefore can be a limiting

    nutrient in aquatic systems

    Essential nutrient (e.g., amino acids,nucleic acids, proteins, chlorophyll)

    Differences from phosphorus

    Not geological in origin Unlike phosphorus, there are many oxidation

    states

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    Nitrogen general properties

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    t oge ge e a p ope t es

    Essential for plant growthNot typically limiting but can be in:

    Highly enriched lakes

    Pristine, unproductive lakes located inwatersheds with nitrogen-poor soils

    Estuaries, open ocean

    Lots of input from the atmosphere Combustion of NO2, fertilizer dust

    Nitrogen general properties

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    g g p p

    Mobile in the form of nitrate (soluble), itgoes wherever water flows

    Ammonium (NH4+) adsorbs to soil particles

    Blue green algae can fix nitrogen (N2) fromthe atmosphere

    Nitrogen has many redox states and is

    involved in many bacterial transformations

    Nitrogen sources

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    g

    Atmospheric deposition Wet and dry deposition (NO3

    - and NH4+)

    Combustion gases (power plants, vehicle

    exhaust, acid rain), dust, fertilizers Streams and groundwater (mostly NO3

    -)

    Sewage and feedlots (NO3- and NH4

    +)

    Agricultural runoff (NO3- and NH4+)Regeneration from aquatic sediments and

    the hypoliminion (NH4+)

    Nitrogen - toxicity

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    g y

    Methemoglobinemiablue baby syndrome > 10 mg/L NO3

    --N or > 1 mg/L NO2--N in well water

    Usually related to agricultural contamination of

    groundwater

    NO3- possible cause of stomach/colon cancer

    Un-ionized NH4+ can be toxic to coldwater fish

    NH4OH and NH3 at high pH

    N2O and NOx contribute to smog, haze, ozone layer

    depletion, acid rain

    Nitrogen many oxidation

    t t

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    states

    Unlike P there are many oxidation statesOrganisms have evolved to make use of

    these oxidation-reduction states for energy

    metabolism and biosynthesis

    -3 0 + 1 + 3 + 4 + 5

    NH4+ N2 N2O NO2- NO2 NO3-

    Nitrogen bacterial transformations

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    Organic N NH4+-N Heterotrophic ammonification or

    mineralization. Associated with oxicor anoxic respiration.

    NH4+-N NO3

    - Involves oxygen (oxic). Autotrophicand chemosynthetic ("burn NH4

    +-N

    to fix CO2).

    NO3- N2 (gas) Anoxic process. Heterotrophic.

    ("burn" organic matter and respireNO3

    -, not O2).

    N2 (gas) Organic N Some blue green algae are able to dothis.

    Decomposition

    Nitrification

    Denitrification

    Nitrogen fixation

    Nutrients- The Nitrogen Cycle

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    g y

    modified from Horne and Goldman. 1994. Limnology. McGraw Hill.

    Chemical forms

    of nitrogen in aquaticsystems

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    OrgN

    N2 = largest reservoir

    but cannot be used by

    most organisms

    Fixedoravailable-N

    organism-N + detrital-N+ dissolved organic-N

    NH4+

    NO2-

    NO3-

    -3 +5+4+3+2+10-1-2Oxidation state

    NO2N2ON2

    gases

    y

    NH4+

    NO2-

    NO3-

    Dissolvedinorganic-N (DIN)Ammonium:

    basic unit forbiosynthesis Nitrite:

    usuallytransient

    Nitrate: majorrunoff fraction

    Functionally in the lab using

    filters

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    filters

    Total-N = particulate organic-N + dissolved organic-N

    + particulate inorganic-N + dissolved inorganic-N

    TN = PN + DON + DIN

    Dissolved inorganic-N = [Nitrate + Nitrite]-N + ammonium-N

    DIN = NO3-N + NO2-N + NH4-N

    Notes: Nitrate+nitrite are usually measured together.

    Nitrite is usually negligible.

    Assimilation Assimilation

    Main N-cycle transformations

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    Assimilation

    (algae + bacteria)

    Assimilation

    -3 +5+4+3+2+10-1-2Oxidation state

    Denitrification

    NO2N2ON2

    NH4+

    NO2-

    Mineralization

    Org-N

    N2 - Fixation- Soil bacteria- Cyanobacteria- Industrial activity

    - Sulfur bacteria

    Denitrification(anoxic bacteria)

    Nitrification 1(oxic bacteria)

    Nitrification 2

    NO3-

    Ammonification

    gases

    Anammox(anoxic bacteria)

    N2 AmmoniaTribs, GW, Precip

    Whole lake N-budget

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    Surficial Sediments

    N2

    Algae

    oxicanoxic

    NO3- NH4

    +

    Nitrification

    Assimilation

    Mineralization

    NH4+

    NitrificationNO2

    -, N2ONO

    Sedimentation

    DINPONDON

    Sedimentation

    DeepSediments BurialBurial

    Ammoniavolatilization

    Tribs, GW, PrecipDON, PON, NO3

    -, NH4+

    NO3-

    Outflow

    diffusion

    N2-fixation

    Mixing

    Mineralization

    Nutrients summer vertical profiles

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    Oligotrophic Eutrophic

    TT

    O2 O2

    NO3

    NO3

    PO4

    PO4 NH4

    NH4

    0 0

    anoxia

    anoxia

    Sulfide and iron summer vertical profiles

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    TT

    O2

    O2

    Soluble Fe

    H2SH2S

    Soluble Fe anoxia

    anoxia

    0 0EutrophicOligotrophic