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Ecosystem Function
primary production forms the base of marine food webs so understanding the variability of primary production in the ocean allows for a better understanding of all marine organisms
biogeochemistry
understand how life in the ocean affects global elemental cycles
global carbon cuycle
big topic because it is closely related to our global warming system
photosynthesis
consumes carbon dioxide gas to form the particulate carbon of algae
difference b/w photosynthesis and respiration
what sinks to the ocean floor
phyloplankton
vast majority of production in the ocean is carried out by chlorophyll containing single-celled organisms
three main groups of phyloplankton
-diatoms: require silica
-flagellates: motile so they are able to avoid sinking in calm waters
-photosynthetic bacteria: able to grow at very low nutrient concentrations
respiration
collectively the generation of CO2 by reactions needed to construct new complex molecules, and to provide basic metabolic needs, consume oxygen. (opposite of CO2)
net primary production (NPP)
difference between the amount of CO2 consumed by photosynthesis and the amount of CO2 produced by respiration. Equivalently, it is the new gain or net loss of carbon within the cell
compensation depth
the depth at which ambient light intensity is equal to the compensation light intensity
dominant cell diameters of phytoplankton assemblage
1.0- 10.0- 100.0
The 4 Phytoplankton Nutrients of Interest to
Oceanographers
1) Nitrogen, 2) Phosphorous, 3) Silica (for diatoms) and 4) Iron because at any given place or time in the ocean it is one of these
four nutrients that is in short supply and can limit the growth of phytoplankton
The Main Source of Nitrogen, Phosphorous and
Silica to the Surface Ocean
The main source of nitrogen, phosphorous and silica
to the surface layer of the ocean is by vertically mixing
or upwelling of nutrient-rich deep-water to the surface
Main source of iron input
dust blowing off of the continents. Southern Ocean
Surface convergence of the Ekman Layer in the subtropics
(forced by the Trade and Westerly Winds) forms a mound/lens of warm (low-nutrient) water (and associated gyre rotation) and an associated downward surface layer velocity into the deeper ocean. Taken together, this makes it difficult for nutrients to move upward to the the surface ocean and so primary production of exceptionally low year-round in the subtropical gyres
subtropical gyres
exhibit low primary production on a per meter square basis and very little seasonal variation
Equatorial Upwelling of Cold Nutrient-Rich Deep
Water in the Eastern Equatorial Pacific and Atlantic
1. Easterly Trade Winds Cause Surface Waters to Pile Up in the West
2. Themocline is Deep in the West and Shallow in the East
3. Proximity of Thermocline Near the Surface in the East Enhances Upwelling of
Cold and Nutrient-Rich Deep-Water to the Lighted Region of the Surface
Ocean and Thus Enhances Biological Productivity in this Area
Equatorial Pacific
Exhibits very little seasonal variability in primary production.
Atlantic
exhibits modest seasonal variability because of sudden seasonal trade wind bursts in the spring
Tidal Mixes
brings nutrients to the surface year-round
Coastal upwelling
seasonally superimposes additional nutrients
The Critical Depth
1. When cells are below the Compensation Depth, they lose carbon because light is too dim to allow for positive net primary production (NPP)
2. The average light level that phytoplankton experience over the course of a day becomes dimmer as mixing depth increases because
cells spend an increasing proportion of the day below the compensation depth in the dark
3. When cells mix below to the Critical Depth they have spent too much of the day below the
compensation depth losing carbon
net losses of carbon experienced while below the compensation depth exceed the net gains of carbon experienced while above the compensation depth
Spring Shoaling of the Thermocline above the ritical Depth
Brings about Positive Net Primary Production (NPP)
1. Changes in the mixing depth
relative to the critical depth
determines if NPP is positive or
negative and thereby determines
if phytoplankton blooms will
occur (i.e., if/when there is
positive NPP) .
2. In winter, mixing is below the
critical depth (due to cold winter
storms) and NPP is negative
3. In spring, mixing is above the
critical depth (due to shallow
thermocline) and NPP is positive
Westerly Wind Region
• Deep vertical mixing in winter
- brings high levels of nutrients to the surface
- causes phytoplankton to mix below the critical depth and so even though
nutrients are plentiful, cells spend too much time in the dark and NPP is
light limited.
• Formation of shallow thermocline in spring
- depth of mixing confined above the shallow thermocline and above the
critical depth so phytoplankton spend much of the day high in the water
column where there is lots of sunlight
- Nutrients are still plentiful from winter mixing so cells have lots of
nutrients and lots of sunlight and spring bloom forms
• Continued stratification in summer
- Mixing remains shallow, and above the critical depth, but nutrients are
depleted and NPP is nutrient limited
Global Distribution of Annual Net Primary
Production (NPP)
Oceanic NPP is about 46% of Global NPP
World Ocean Net Primary Production
While the Open Ocean (Trade Winds, Westerly Wind and Polar regions)
exhibit relatively low intensities of primary production (NPP per square meter) relative to coastal regions, they contribute most (71%) as a whole to the global ocean total NPP because of the vast areas comprising these regions
Pelagic
the water column environment
Benthic
the seafloor environment
Holoplankton
Planktonic organisms that live their entire life in fluid suspension. IE: Copepods
Meroplankton:
Planktonic organisms that spend only part of their life in fluid suspension
Autotrophs
Group of organisms whose carbon for growth comes from non-organic sources. For example, phytoplankton are autotrophs because they use CO2 for their carbon needs
Heterotrophs
Group of organisms whose carbon for growth comes from previously formed organic carbon material. For example, herbivorous zooplankton are heterotrophs because they consume phytoplankton for their carbon needs. Carnivores would also be heterotrophs
Trophic Level
Nutritional feeding level within a food chain or food web e.g., primary producer (i.e., autotroph), primary consumer (herbivore), secondary consumer (first carnivore), tertiary consumer (second carnivore),
et
Size
Size determines almost everything about an organism's position/role in
the community of pelagic organisms (except for the possibility of containing or not containing chlorophyll) (1/10) Marine food webs are said to be strongly size structured
Trophic Transfer Efficiency
depends on 1. Exploitation Efficiency and 2. Gross Production Efficiency
Exploitation Efficiency
1. strategies for detecting prey
2. strategies for capturing prey once detected
3. counter strategies to avoid detection in the first place
4. counter strategies to frustrate capture if detected
Diel Vertical Migration (Avoid Detection)
Much of the zooplankton community migrates up to the surface layer of the ocean at night to feed in the dark while also avoiding visual predators like small fish. During the day they migrate down to the safety of the darkness found at depth
Exploitation Efficiencies in Spring Blooms in the temperate north atlantic regions
During long winter periods grazers (copepods mainly) sink into the deep ocean and enter a diapause (i.e.,
hibernation) stage and thereby become decoupled from any variations in primary production above.
2. In spring, phytoplankton standing stock can initially grow to very high density because it is not held in check by strong grazing pressure (phytoplankton growth is decoupled from grazing) until the large grazers have a chance to come out of diapause, grow and reproduce to
the high numbers needed to control increases in phytoplankton abundance.
3. This allows for exceptional phytoplankton blooms during
the decoupled period
**Exploitation efficiency is very low in this case- much of the phyloplankton is not found by the grazers and instead sinks into the deep ocean as dead phytoplankton cells
Exploitation Efficiencies in Tropical Environments
1. Grazers remain active throughout
the year and consume
phytoplankton as fast as it is made.
2. Any increase in production is
quickly met by an increase in
consumption.
3. This leaves standing stock of
phytoplankton nearly constant
throughout the year
**Exploitation efficiency is very high in this case- almost all phyloplankton is found and consumed by grazers
Gross Growth (Production) Efficiency
Amount of CONSUMER
BIOMASS produced divided by amount of PREY INGESTED. This
efficiency ranges between 20% and 60%
Trophic Transfer Efficiency is a function of
Exploitation Efficiency (10% to 90% )
Gross Production Efficiency (20% to 60%)
The combined effect of both exploitation and gross production efficiencies yields an overall
trophic transfer efficiency of about 10% to 20%
The number of trophic levels between phytoplankton and
harvestable fish
smaller in high nutrient regions such as coastal upwelling regions
Highest production of harvestable fish is in the
coastal ocean region
1. The open-ocean region comprises most of the global ocean primary production, but phytoplankton in this region are small and so there are a lot of trophic steps (7 steps) to get to harvestable fish and each trophic step reduces carbon biomass
production by 1/10 so for 7 steps there is a 1/107 reduction altogether.
2. The coastal region has less overall primary production, but it benefits greatly by having just 2 trophic steps to harvestable fish which makes for very efficient transfer of carbon from primary producer to harvestable fish in this region and so it is this region that is the most productive for fish.
Use of Epifluorescent Microscopy and Fluorescent DNA Stains Became Widespread Between 1975 and 1985
• Dramatically increased estimates of bacterial concentrations in the ocean
• Also allowed easy distinction between autotrophic and
heterotrophic flagellate cells (i.e., chlorophyll containing or chlorophyll
lacking)
Bacteria-Sized Autotroph
-Came to be Known as Prochlorococcus
-The Discovery was Made using a New Technique called Analytical Flow Cytometry
-High Abundance, especially in Oligotrophic Regions
oligotrophic (low nutrient) open-ocean environments
the growth advantage goes to the smallest phytoplankton cells which are now recognized to be represented mainly by
Prochlorococcus.
Prochlorococcus
the main contributor to primary
production in open-ocean
environments.
-autotrophic bacterium
-responsible for more than a quarter of the global ocean primary production
Heterotrophic Bacteria
highly abundant in all ocean environments
Oligotrophic:
Pelagic environment (water column) that has naturally very low plant nutrient concentrations
--the vast subtropical gyres are oligotrophic
Eutrophic
Pelagic environment (water column) that has naturally high plant nutrient concentrations
--coastal upwelling zones are eutrophic
Global Carbon Cycle
The pathway that carbon takes from CO2(gas) to particulate organic
carbon (through photosynthesis), and on into particulate organic carbon of
higher trophic levels, varies when nutrient concentration varies
efficient biological carbon pump
when the dominant phytoplankton cells are large, the dominant grazers are large and their large fecal material easily sinks to the deep ocean taking organic carbon down with it
Inefficient biological pump
when the dominant phytoplankton is small and grazers are small and fecal matter is so small that it cannot easily sink and the particulate carbon is instead respired back to CO2
fixed chemical stoichiometry
all living things have a roughly fixed ratio of major elements in their cells
-because of this the pattern of cycling and export to the deep ocean for all of major elements will look quite similar
Nitrogen Cycling
the concept of new and recycled primary production
Total primary production
recycled + new primary production
New Primary production
uses Nitrate (NO3) from the deep ocean for its nitrogen source
Recycled Primary Production
uses Ammonia (NH4) generate by animal excretion in the upper ocean for its nitrogen source
Eutrophic (high nutrients)
conditions are dominated by large cell and most of the primary production is new production
oligotrophic (low nutrients)
conditions are dominated by small cells and most of the primary production is recycled production
As nutrient concentration is reduced
...competitive growth advantage shifts to small phytoplankton cells
Molecular Structure of Water
Water is a highly polar molecule
Hydrogen Bond
electrostatic attraction between partial + and - charges on separate polar molecules
Low Temperature Limit of Water
-Energy of the hydrogen bond is greater than the energy of the thermal bond
-maximum # of H bonds
-maximum order, low thermal motion
-regular lattice structure of ice
-solid
Intermediate Case of Water
-E h-bond is equivalent of E thermal
-clusters of H- bonded water (structural water)
-interspersed non-H bonded water (free water)
-liquid water
High temp limit of water
E h-bond < E thermal
-minimum number of H bonds
-minimum order, rapid thermal motion
-independant, non interacting gas molecules
-water vapor (gas)
Specific Heat Capacity
Amount of heat required to raise 1 gram of liquid water by 1 degree Celsius. This is among the highest of any substance on earth
Latent Heat Vaporization
Amount of heat required to convert 1 gram of liquid water to water vapor
-540 calories per gram
Waters high heat capacity
-means that it takes an exceptionally large amount of heat energy to change ocean temperatures
-all thing being equal- if you have more heat energy coming into the earth than leaving the earth you should observe a steady rise in global temperature
Condensation
latent heat released into the atmosphere by condensation of water vapor to form clouds and rain
Evaporation
Latent heat removed from the ocean and stored in the atmosphere in the form of water vapor
Molecular Properties of Water
-strong polar nature makes it a very good solvent for ionic constituents
-hydrogen bonds are weak, but below 100C they are strong enough to allow individual water molecules to bond temporarily with other water molecules to form liquid water
-high specific heat capacity
-high latent heat of vaporization- allows large amounts of heat to be removed from the ocean, stored at latent heat in the form of water vapor and then transported by winds to other parts of planet where it can then be released to the atmosphere as sensible hear upon precipitation
salinity
a measure of the salt concentration in a sea water example
-often expressed as the number of gramps of salt contained in a thousand grams of seawater
salt in the ocean
the magnitude of input and output rates have been roughly equal for millions of years
Surface ocean salt concentration
the total amount of salt in the ocean does not vary, but the unequal addition/removal of freshwater over the global ocean's surface creates large regional differences in surface ocean salinity
Salinities in the ocean
set at the air-sea interface
-overall, salinity is a direct function of evaporation minus precipitation
Atlantic Ocean Salinity
Once removed from the surface the salinity remains constant unless it mixes with other water masses
Salinity Variation
-while salinity may vary considerably in different regions, the relative proportion of one ion to another does not vary
Geochemical Cycles
keeping track of elemental inputs, chemical transformations and outputs
Conservation Constituents of seawater
those that are only varied by physical exchange processes at the sea surface.
Once the water leaves the surface, salinity, temperature and inert gas concentration are conserved
Non-conservative Constituents
those that are varied by processes (other than mixing) that occur anywhere in the water column
Plant Nutrients
Nonconservative constituents
1- Low in surface layer because of rapid uptake by phytoplankton in the presence of sunlight
2- High at depth because of respiration/ remineralization and no uptake by phytoplankton in the dark
global warming
expected to increase the strength of the thermocline and thereby reduced vertical mixing and diffusion across this boundary and make the oxygen minimum zone even lower
Photosynthesis Consumes
CO2 in the surface ocean to form particulate organic carbon
Respiration Produces
CO2 in the deep ocean