Define and briefly discuss the following terms in the context of the course:

1: Redfield Ratio

Alfred Redfield, a physiologist at Harvard/Woods Hole Oceanographic, made many observations

of nutrients in seawater. In 1963 he wrote a CLASSIC paper: The influence of organisms on
composition of seawater. His conclusion was that biologically active elements circulate in a very
different fashion from the general water circulation- biochemical circulation.

Redfield measured the elementary composition of the plankton:

REDFIELD RATIO:
****106C:16 N: 1 P****
MEMORIZE THIS!!

This is the average value for plankton of the three major structural elements. It should be
remembered that it can change because of different biological processes (esp. species
composition).

The relationship of the concentrations of N and P (and associated trace elements) are similar in
different regions in proportions, but not concentrations

2: SOFAR

SOFAR Zone (= sound fixing and ranging).

Sound travels at high speed at the bottom of the mixed layer. A thin, high velocity layer occurs
at about 80 m, just above the pycnocline.

Depending on the angle at which sound waves from the surface arrive at the high-velocity layer,
they will sometimes split and refract to the surface or bend to the depths. An object beyond the
area of divergence would be undetectable- it would be in a shadow zone, a region into which
very little sound energy penetrates. A smart sub captain uses the shadow zone to hide. The
ideal "safe depth" is about 125 m. (Tom Clancey: The Sum of All Fears)

These principles are now used for measuring current speeds in the deep ocean and to measure
the change in temperature of the oceans that may be caused by global warming

3: Spring Tides

When the sun and moon are aligned, the resulting tides are called spring tides- which give the highest
highs and lowest lows of the month.

4: Adaptation

Adaptations: special inherited features that enable a species to function in its niche

Organisms rarely have fixed tolerance limits.

5: Carbonate Compensation Depth

Reverse Weathering: (remove ions from seawater)

Ions in solution precipitate onto solid surfaces by adsorption or co-precipitation of minerals from
seawater. CaCO3 spheres in shallow depths gain weight, but lose weight in deep depths.

The point where the net change in weight of the spheres is zero is the carbonate compensation depth.

6: Isostasy

During the ice ages, the mass moved from the ocean to the land, continents sank and the floors of the
ocean rose about 40 m. When the ice melted, the mass moved and the continents rebounded. This
crustal balance is ISOSTASY. The pressures equalize geophysically

7: Warm core rings

Gulf Stream Eddies: very large eddies of the Gulf Stream form quite frequently to the north and south
of the stream as meanders pinch off. North of the stream, the eddies are warm and rotate clockwise
(warm core rings). South of the stream, the eddies are cold and rotate counterclockwise (cold core
rings).

8: I(z) + Io e^-kz

Attenuation coefficient

k = attenuation coefficient

(wave-length specific)

-different colors attenuate at different rates

-red light is attenuated more rapidly than blue light

-blue light penetrates deepest in pure water

-when particulate and dissolved organic matter are present, green light penetrates.

-a red starfish at depth looks black!

3. Attenuation

way the amount and spectral composition change with depth

9: Pyncocline

Physical Properties of the Ocean

 Relationships with depth in the ocean:

Halocline = gradient in salinity

Thermocline = gradient in temperature

Pycnocline = gradient in density

10: anticyclonic circulation

-in Northern Hemisphere: there is a cyclonic circulation around a low pressure system
(counter-clockwise)

- there is anticyclonic circulation around a high (clockwise).

-these processes are responsible for much of the global water circulation.

11: non-conservative ions in seawater

Major ions have similar proportions in world's oceans (behave conservatively)

Minor & trace elements behave non-conservatively. The salinity does NOT indicate the concentrations
of these elements because they are generally governed by organisms. (e.g., phosphate and nitrate

12: oligotrophic

Vast areas of the ocean surface are downwelling regions, with very slow velocities. These
anticyclonic gyres are very oligotrophic, and are sometimes called the deserts of the ocean.
Examples are the Sargasso Sea and the North Pacific Gyre.

13: ABW

Antarctic Bottom Water (ABW) which is even denser, so the NADW is forced toward the surface. It
finally reaches the surface as a upwelling system rich in nutrients in the Antarctic region, which fuels the
great productivity of that area

The North Atlantic Deep Water (NADW) is cold and salty and because of its density sinks to the bottom
and travels south. Near the equator, it encounters the Antarctic Bottom Water (ABW) which is even
denser, so the NADW is forced toward the surface. It finally reaches the surface as a upwelling system
rich in nutrients in the Antarctic region, which fuels the great productivity of that area

14:neretic

A small number are responsible for red tides and/or the production of toxins (about 20 spp. out
of > 1000 extant spp.).

Red tides: these are not always dinoflagellates, and when they are, they don't always make toxins
simultaneously. All coasts of North America have reported outbreaks. All species that make toxins are
photosynthetic, all are estuarine or neretic, and all produce benthic stages (hypnozygotes) by sexual
reproduction. Most of these species are capable of producing monospecific blooms, suggesting
competitive displacement may be occuring. The toxins are water soluble or lipid soluble that are
hemolytic or neurotoxic in activity, depending on the structure, dose, or susceptibility of the consumer

15: hypoosmotic

Marine fish: hypoosmotic: must drink seawater, secrete salt, excrete small amts of
concentrated urine

16epifuana

Plankton -drift with ocean currents

Classed by size (see Fig 15.2 in book)

pico- (0.02-2 m), nano- (2.0-20 m), micro-(20-200 m), meso- (0.2-2 mm), macro- (2-20 cm),
mega- (20-200 cm)

Nekton - organisms that move independently of currents

fish, squid, mammals, turtles

Benthos -live on (epifauna) or in (infauna) the bottom sediments

17: adaptation

18: Carles Wyville Thompson

Charles Wyville Thompson HMS Lightning (1868) & HMS Porcupine (1870); wrote first
oceanography book (1873): The Depths of the Sea. Thompson and John Murray were the
leaders of the Challenger Expedition aboard HMS Challenger 1872-1876- a worldwide
exploration of the sea.

He wrote the first general reference: The Depths of the Sea.

Thompsons greatest achievement was to persuade the British Admiralty to outfit a ship and
sponsor an expedition for the worldwide study of the ocean floor

19: Meroplaktonic

Meroplanktonic: organisms that spend part of their lives in the plankton and part in the
benthos (bottom). Also, species that require benthic resting stages

Neritic: species found in water above the continental shelf.

Tychoplanktonic: organisms normally attached to substrate, but they may be occassionally

broken off and thus float in the plankton.

20: Paradox of the plankton

 How can so many species occupy the seemingly homogeneous mixed layer of a lake or ocean if
the principle of competitive exclusion is correct? (G.E. Hutchinson).

Many limiting resources or disequilibrium

The seven major ions of seawater

sodium ion (Na+), chloride (Cl-), sulfate (SO42-), magnesium ion (Mg2+), calcium ion
(Ca2+), and potassium ion (K+), Bi carbonate HCO3

Three factors which most influence the desnit of seawater

Temperature, salinity and pressure affect density

What does ENSO mean? Explain the major features that characterize an enso event

El Nio-Southern Oscillation

First identified as a change in surface currents along the coast of Chile and Peru and often begins
near Christmas time- thus El Nio- The Child- for birth of Jesus.

Southern Oscillation is a long-distance linkage of the atmosphere barometric pressure over the
Pacific/Indian Oceans. The low pressure system oscillates between western Pacific/Indian
Ocean and Eastern Pacific.

Who coined the term: This blood of the sea What does it mean?

Victor Hensen: "Father of German Oceanography"

Plankton are the "blood of the sea". They are the food source for the rest of the sea,
sea wouldnt function without them

Continental crust is granite and oceanic crust is basalt. Which is more dense? How do they differ in
thickness?

Continental crust (land) is made up of igneous granite (intrusive)

large crystals; Si, Al, Na, K; density = 2.7 g cm-3; thickness = 35 km

Oceanic crust is made up of igneous basalt (extrusive)

small crystals (cools fast); Si, Mg, Ca, Fe; density >3.0 g cm-3

thickness = 7 km

Mantle molten layer = asthenosphere; density = 3.3 g cm-3.

Diagram the exponential growth curve and the logistic growth curve. What makes the logistic curve
different?

Density Independent Growth: dN/dt = rN = exponential growth

Density Dependent Growth: Logistic Equation dN/dt = r N (K-N/K)

K is carrying capacity
 r= maximum capacity for growth

What is the difference between the fundamental niche and the realized niche?

G.E. Hutchinson: n-dimensional hypervolume

-fundamental niche: n-dimensional space an organism theoretically occupies.

-realized niche: actual space occupied- usually restricted by biological interactions

Would you expect a more r-selected species or a more K-selected species to become dominant in an
upwelling area? Why? Indicate which life history traits are likely to characterize this species.

r-selection: selection for traits that favor rapid population growth at low densities (= fugitive,
opportunistic, zymogenous...)

K-selection: selection for traits that favor competitive ability at densities near the carrying
capacity (= equilibrium, autochthonous,...)

There is a CONTINUUM of tradeoffs between reproductive capacity and efficiency.

ANTARCTIC (SOUTHERN): interfaces with three major oceans; westward circulation of water;
upwelling of deep water = nutrient rich; highly productive (ca. 50% of total ocean).

I would expect more r selected species to become more dominant since upwelling area have so
much nutrients there is essentially no limit to how much you could grow thus species that can
growth at more exponential growth rate would flourish more.

What are four of the six processes that remove ions from seawater. (8 points)

Ion Exchange

Carbonate formation

Reverse Weathering

Opal Formation

Sulfate Reduction

Evaporite Formation
 ADDS SEAWATER:

Chemical Weathering, Cyclic Salts

Draw a diagram of how each of the factors below change with depth in the open ocean during
the summer (10 points)

(temperature, salinity, oxygen, sound transmission, light intensity)

Extra credit: name three marginal seas of the Atlantic ocean (3 points)

Meditterranian

Carribean

Hudson Bay
ALL RED TEXT:

Captain James Cook HMS Endeavor 1758-1779 Pacific Ocean

Alexander von Humboldt 1779-1804 Central & South America

Edward Forbes Shetland Sea Benthos 1830's-40's

Charles Darwin HMS Beagle 1831-1836 So. America & Pacific

In 1831 he was appointed naturalist on the HMS Beagle, a ship commissioned to map
and explore around South America and the Pacific

Charles Wyville Thompson HMS Lightning (1868) & HMS Porcupine (1870); wrote first
oceanography book (1873): The Depths of the Sea. Thompson and John Murray were the
leaders of the Challenger Expedition aboard HMS Challenger 1872-1876- a worldwide
exploration of the sea.

Thompson wanted to know about life at great depths (Forbes had said it did not exist).
In summers of 1868 and 1870, he dredged and measured temperatures in deepwater
off Europe and discovered globigerina oozes. He wrote the first general reference: The
Depths of the Sea.

Victor Hensen: "Father of German Oceanography"

Plankton are the "blood of the sea"

Plymouth Laboratory- studied plankton biology in the English Channel

: Louis & Alexander Agassiz

-They did more to promote marine science than anyone else of 19th and early 20th
centuries

Gordon A. Riley 1930's-1960's (G.E. Hutchinson, Yale). Tremendous impact on modern

plankton studies;

Bruce Heezen: seafloor mapping Plate Tectonics

OCEANS: PACIFIC, ATLANTIC, INDIAN, ARTIC, ANTARTIC

continental shelf: big changes in sea level; goes out to ca. 100-200 m depth

continental slope: ca. 4 angle; submarine canyons; turbidity currents

 abyssal plain: ca. 5,000 m

three major types of sediment:

1. sediments from land -lithogenous

2. sediments of biological origin - biogenous: CaCO3 + SiO2 oozes.

3. wind blown sediments: aeolian: red clay

-there are also hydrogenous manganese nodules

Continental crust (land) is made up of igneous granite (intrusive)

large crystals; Si, Al, Na, K; density = 2.7 g cm-3; thickness = 35 km

Oceanic crust is made up of igneous basalt (extrusive)

small crystals (cools fast); Si, Mg, Ca, Fe; density >3.0 g cm-3

thickness = 7 km

Mantle molten layer = asthenosphere; density = 3.3 g cm-3.

This crustal balance is ISOSTASY. Rocks in ocean are about 160-180 million years old

At the edges of the continents the oceanic crust moves down (it is more dense!) into the mantle
= subduction.

Alfred Wegener proposed the Theory of Continental Drift,

seafloor mapping being done by Bruce Heezen and others.

Rock magnetism: Sea floor spreading: . Ocean Ridge System: PLATE TECHTONICS

Lecture 2:

IONS IN SEAWATER:

ANIONS CATIONS

Chloride, Cl- 18.980 Sodium, Na+ 10.556

Sulfate, SO42- 2.649 Magnesium, Mg2+ 1.272

Bicarbonate HCO3- 0.140 Calcium, Ca2+ 0.400

Potassium, K+ 0.380

. Eleven major ions make up 99.99% of salinity. They are the ions left behind after a variety of complex
reactions occur, such as the formation of organic matter, skeletons, sediments. There is biological and
geochemical processing.

, the chloride in the ocean is an excess volatile

Processes that ADD ions to seawater:

Chemical Weathering

Cyclic Salts

Rainwater composition looks very much like seawater, dominated by sodium and chloride

The chloride entering the ocean came primarily from the ocean.

Seawater is in a long-term steady state for chloride.

Processes that REMOVE ions to seawater:

1 Ion Exchange,2 Carbonate Formation, 3Reveres Weathering

Reverse Weathering: The point where the net change in weight of the spheres is zero is the carbonate
compensation depth

4 Opal Formation

. Si is part of the long-term buffering system of the ocean.

5 Sulfate Reduction

6 Evaporite Formation

--

Minor Constituents of Seawater:

Major ions have similar proportions in world's oceans (behave conservatively)

Minor & trace elements behave non-conservatively. The salinity does NOT indicate the concentrations
of these elements because they are generally governed by organisms.

Alfred Redfield, a physiologist at Harvard/Woods Hole Oceanographic, made many observations

of nutrients in seawater. In 1963 he wrote a CLASSIC paper: The influence of organisms on
composition of seawater. His conclusion was that biologically active elements circulate in a very
different fashion from the general water circulation- biochemical circulation.

The elements required for biological production are drawn out of seawater in particular proportions
during the formation of organic matter. The twin processes of synthesis and regeneration can be
separated in space

REDFIELD RATIO:
****106C:16 N: 1 P****
MEMORIZE THIS! average value for plankton

Halocline = gradient in salinity

Thermocline = gradient in temperature

Pycnocline = gradient in density

Temperature, salinity and pressure affect density

 AIW = Antarctic Intermediate Water S = 33.8

T= 2.2 C

ABW Antarctic Bottom Water S = 34.62

T = -1.9 C

Sound Propagation of seawater:

-velocity decreases with decreased temperature

-velocity increases with increased pressure

Temperature effects dominate in shallow water

Pressure effects dominate in deep water

SOFAR Zone (= sound fixing and ranging).

Sound travels at high speed at the bottom of the mixed layer A thin, high velocity layer occurs at about
80 m, just above the pycnocline.

Marine fish: hypoosmotic: must drink seawater, secrete salt, excrete small amts of
concentrated urine

Incident light: light at surface of seadirect or diffuse (scattered)

2. Reflection influenced by sea state, waves, foam patches, bubbles

3. Attenuation way the amount and spectral composition change with depth

4. Absorption and Scattering: reason light decreases with depth

LIGHT DECREASES EXPONENTIALLY WITH DEPTH

Attenuation Coefficient

I (z) = Io e-kz

k = attenuation coefficient
 (wave-length specific)

-different colors attenuate at different rates

Lecture 3:

Coriolis Effect

Points at different latitudes on the surface rotate at different velocities,

Eckman Spiral

The wind drives the surface water in a direction of 45 to the right of its path

-in Northern Hemisphere: there is a cyclonic circulation around a low pressure system
(counter-clockwise)

- there is anticyclonic circulation around a high (clockwise).

Upwellings under lows

Downwellings under highs

These create geostrophic currents

Water piles up in the center of the mid-ocean gyres because of Coriolis effect, and forms a hill

Upwelling regions bring nutrients to the surface and are therefore areas of high primary
productivity.

Downwelling regions restrict the resupply of nutrients and are therefore regions of very low
productivity.

These anticyclonic gyres are very oligotrophic, and are sometimes called the deserts of the
ocean

Langmuir circulation -mixes plankton organisms in the surface region, driven by the wind (
 Coastal upwellings are on scales of 100's of km. Along continental margins, Ekman transport
carries surface water away from the continent, and upwelled water replaces the water that has
moved away.

Equatorial upwellings are created by the Coriolis effect acting on the westward flowing equatorial
currents

Gulf Stream Eddies: very large eddies of the Gulf Stream form quite frequently to the north and south
of the stream as meanders pinch off. North of the stream, the eddies are warm and rotate clockwise
(warm core rings). South of the stream, the eddies are cold and rotate counterclockwise (cold core
rings).

The North Atlantic Deep Water (NADW) is cold and salty and because of its density sinks to the bottom
and travels south. Near the equator, it encounters the Antarctic Bottom Water (ABW) which is even
denser, so the NADW is forced toward the surface. It finally reaches the surface as a upwelling system
rich in nutrients in the Antarctic region, which fuels the great productivity of that area

Meridional Overturning Circulation: MOC

Slowing of the thermohaline circulation (MOC) as a result of higher temperatures in polar

regions (especially Arctic) and a decrease in salinity of surface waters due to ice sheet melt

TIDES:

Tides are noticeable features where the ocean reaches land, and form the interesting,
biologically diverse, intertidal zone.

The moon is overhead 24h and 50 min later every day- this is the lunar tidal cycle

When the sun and moon are aligned, the resulting tides are called spring tides- which give the highest
highs and lowest lows of the month.

When the sun and moon are not aligned, the tides are called neap tides- which produce lower
amplitudes.

Semidiurnal tidal cycles- twice each day

Diurnal tidal cycles- typical

Mixed tidal cycles- large land effect

The king tides occur when the Earth, moon and sun are aligned at perigee and perihelion

earth is closest to the sun (perihelion)

moon is closest to the earth (perigee)

Size of waves also depends on fetch- the length of open ocean over which the wind blows.

Crest highest point; Trough- lowest point

Height- vertical distance from crest to trough

 Wavelength- distance between crests

Period- time it takes for wave to go past a point

Capillary waves: light winds over water cause ripples

Gravity waves: stronger winds above 6 km/hr (4 mph) cause large waves to form

Tsunamis: They are seismic sea waves.

ENSO

El Nio-Southern Oscillation

First identified as a change in surface currents along the coast of Chile and Peru and often begins
near Christmas time- thus El Nio- The Child- for birth of Jesus.

Southern Oscillation is a long-distance linkage of the atmosphere barometric pressure over the
Pacific/Indian Oceans. The low pressure system oscillates between western Pacific/Indian
Ocean and Eastern Pacific.

La Nia is the opposite extreme of El Nio, but does not always occur after an ENSO event.

Divisons of Marine Environemnt:

Pelagic: open ocean environment

Benthic: ocean bottom environment

Neritic: from the shore out to 200 m depth

Oceanic: area beyond the 200 m contour

In the mesopelagic zone, there is an oxygen minimum

Plankton -drift with ocean currents

Classed by size (see Fig 15.2 in book)

pico- (0.02-2 m), nano- (2.0-20 m), micro-(20-200 m), meso- (0.2-2 mm), macro- (2-
20 cm), mega- (20-200 cm)

Nekton - organisms that move independently of currents

fish, squid, mammals, turtles

Benthos -live on (epifauna) or in (infauna) the bottom sediments

LECTURE 4:

G.E. Hutchinson: n-dimensional hypervolume

 -fundamental niche: n-dimensional space an organism theoretically occupies.

-realized niche: actual space occupied- usually restricted by biological interactions

No two species in equilibrium can occupy the same niche. (Gause)

Paradox of the Plankton

How can so many species occupy the seemingly homogeneous mixed layer of a lake or ocean
if the principle of competitive exclusion is correct? (G.E. Hutchinson).

Adaptations: special inherited features that enable a species to function in its niche

Organisms rarely have fixed tolerance limits.

Acclimatization: changes in tolerance with seasonal environmental change. This usually

involves several correlated environmental changes.

Acclimation: compensatory process involving a shift in a function following an environmental

change.

Density Independent Growth

dN/dt = rN or Nt = No ert = exponential growth

Density Dependent Growth Logistic Equation

dN/dt = r N (K-N/K) r= maximum capacity for growth K = carrying capacity

for organisms with one or more generations per year, life tables are constructed.

survivorship curves are constructed as a logarithmic function of lx per unit time.

r-selection: selection for traits that favor rapid population growth at low densities (= fugitive,
opportunistic, zymogenous...)

K-selection: selection for traits that favor competitive ability at densities near the carrying
capacity (= equilibrium, autochthonous,...)

There is a CONTINUUM of tradeoffs between reproductive capacity and efficiency.

intraspecific between individuals of same species

-interspecific between individuals of different spp.

 Interference competition: access to a resource is denied to competitors by the dominant
individual or species. e.g.:

-Exploitative competition: (or scramble competition): the direct use of a resource that
reduces its availability to a competing individual or species, simply by consumption. e.g., taking
up essential resources

coefficient of competition: 12 or 21. 12 = competition coefficient that scales the effect of an

individual of species 2 on species 1

21 = effect of individual of sp 1 on sp 2.

Connell: intertidal Balanus & Chthamalus

Functional Responses to prey density:

Holling curves: as prey increases, predators eat more prey.

1. Rate of Successful Search for Prey

2. Search & Handling Time

3. Hunger Level of Predator

4. Inhibition of Predation by Prey

commensalism: benefits one +/0

2. mutualism: benefits both +/+

3. parasitism: benefit at host's expense

Factors affecting diversity:

a. time: older communities more diverse

b. spatial heterogeneity: more complex = more spp (patchiness)

c. competition: species packing by character displacement

d. environmental stability: more stable- species less likely to go extinct because of sudden
environmental change

e. predation: lowers competition that can lead to exclusion

plankton: Greek for wanderer

coined in 1887 by Victor Hensen

Neuston: organisms attached to the air-sea interface: includes bacteria, protozoa, algae and
even larger animals such as Physalia (Portuguese man-of-war).

Nekton: animals that swim (adult fish, large crustacea, marine mammals). Some small fish and
larvae may be plankton.
 Holoplanktonic: spend entire life in open water; no benthic stages

Meroplanktonic: organisms that spend part of their lives in the plankton and part in the
benthos (bottom). Also, species that require benthic resting stages

Neritic: species found in water above the continental shelf.

Tychoplanktonic: organisms normally attached to substrate, but they may be occassionally

broken off and thus float in the plankton.

some filamentous forms have special cells with thick walls (=heterocysts), which are centers of
N-fixation

may have gliding motility, but no flagellae can function as symbionts,

N-fixation is a very important process in the nitrogen balance of the ocean, and only
cyanobacteria are capable of doing it. Sometimes this is done by symbiotic cyanophytes, or by
Trichodesmium, which is colonial.

Pyramimonas: alternates between flagellate cell and benthic stage.

Micromonas: common in the plankton; very small (1-2 m)

capable of phagotrophy (when they both photosynthesize and engulf cells, they are called
mixotrophs = mixed feeders).

species such as Distephanus have internal Si skeletons

coccolithophores

this class was separated from the previous one

may be either unicellular or multicellular in colonies or filaments

The most important species is Emiliania huxleyi.

Olisthodiscus is cultured as a food for oysters. It lives in brackish waters

Chroomonas is a common species.

(some are colorless = heterotrophs)

zooxanthellae, especially in invertebrates or certain marine ciliates (protozoa), in some cases

causing red tides

most distinctive feature is that the cell walls are made of silica (glass), and are called frustules

The zygote is called an auxospore

dinos also produce non-motile zooxanthellae, especially when in association with hermatypic
corals (reef-building corals)

A small number are responsible for red tides and/or the production of toxins (about 20 spp. out
of > 1000 extant spp.).
 Red tides: these are not always dinoflagellates, and when they are, they don't always make
toxins simultaneously. All coasts of North America have reported outbreaks. All species that
make toxins are photosynthetic, all are estuarine or neretic, and all produce benthic stages
(hypnozygotes)

NSP: Neurotoxic Shellfish Poisoning (Gulf of Mexico)

PSP: Paralytic Shellfish Poisoning (worldwide)

DSP: Diarrhetic Shellfish Poisoning (worldwide)