Marine Ecology Flashcards

1
Q

Describe and explain the seasonal cycle of biological activity (e.g., phytoplankton, zooplankton and fish biomass and production) in the pelagic.

A

Other factors influence phytoplankton growth rates, including water temperature and salinity, water depth, wind, and what kinds of predators are grazing on them. Phytoplankton can grow explosively over a few days or weeks. Basically, the seasonal cycle is driven by sea-surface temperature and the onset of the thermocline leading to phytoplankton blooms during spring, the prevalence of thermal stratification leading to exhaustion of nutrients and subsequent demise of phytoplankton during summer-autumn, and remixing and regeneration of nutrients during winter.

Plankton predominantly comprises short-lived organisms. As a rule, these reproduce so rapidly that several generations may be produced within a single year. The development of planktonic organisms generally follows a regular annual cycle that begins with a spring bloom of the phytoplankton.

Fish biomass:
LOOK AT SLIDES

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2
Q

Explain how environmental factors and the species diversity and biomass of benthic communities vary from the continental shelf to the deep sea.

A

There are a lot of factors that can affect benthic biodiversity along a depth gradient like water temperature, light availability, oxygen (and other elements) concentration and pressure.

Interactions between pelagic and benthic environments are related to a variety of abiotic and biotic processes that have a major influence on the structure and dynamics of marine ecosystems. Transport of particulate and dissolved materials, gases, as well as living organisms, and also sedimentation and erosion are subsumed under these processes that induce a shifting of materials between benthic and pelagic material pools and vice versa. Imbalances in these transactions result in a change of biotic structures and have far-reaching consequences for the development of the communities. Exchange processes are either directed from water to the bottom sediment (termed as pelagic–benthic) or reversed (termed as benthic–pelagic), and impact on abiotic material pools as well as on the biota, such as producers and consumers, or can be related to the exchange between abiotic and biotic material components.

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3
Q

Describe and explain intertidal zonation on rocky shores

A

Intertidal zonation refers to the frequently observed pattern by which species replace one another along a gradient from the low to high tide lines along many of the world’s coastlines.

variation in the distribution of organisms caused by differences in both biotic and abiotic conditions along an environmental gradient. Organisms living on the rocky shore have different adaptations to these factors and therefore will be able to survive at different heights on the shore accordingly.

The intertidal zone or littoral zone is the shoreward fringe of the seabed between the highest and lowest limit of the tides. The upper limit is often controlled by physiological limits on species tolerance of temperature and drying. The lower limit is often determined by the presence of predators or competing species.

The rocky intertidal ecosystem can be divided into four zones: the splash zone, high intertidal, middle intertidal, and low intertidal

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4
Q

What is the Coriolis effect (force)?

A

Apparent deflection of a moving object when viewed from a rotating frame of reference.
Freely moving objects on the surface of the Earth experience the ‘Coriolis force’.
On a non-rotating planet, ocean currents (and winds) would flow directly from areas of high pressure to low pressure.
Because Earth rotates, currents (and winds) flow to the right of this direction north of the equator, and to the left of this direction south of the equator.

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5
Q

What is Ekman Transport?

A

Friction between wind and water surface causes water to move in direction of wind.
The Coriolis effect deflects this current.
The surface layer drags the layer beneath, which is also deflected.
The net movement of the ocean’s surface layer is perpendicular to the right of the wind in the Northern Hemisphere and to the left in the Southern Hemisphere.

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6
Q

Deep Ocean Circulation and Thermohaline Circulation

A

Thermohaline circulation is due to differences in the density that arise from variations of temperature and salinity.
thermo = heat
haline = salt (halide ions)
Solar radiation warms waters in the tropics, causing them to expand (become less dense) and float.
The ocean loses heat to the atmosphere at high latitudes (air colder than water), causing the surface waters to cool and contract (become denser) and sink.

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7
Q

What affects the density of sea water?

A

Density = mass/volume
Units kg m-3
Density increases as temperature decreases
Density increases as salinity increases
Density increases during ice formation because salt is excluded from the ice
Density decreases following rain due to dilution of salt

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8
Q

Mechanisms of Deep Water Formation

A

The mechanisms of deep-water formation are different in North & South Atlantic.
In the North Atlantic, high salinity water (brought north by the Gulf Stream) is cooled in winter leading to deep convective mixing.
On the Antarctic continental shelf, ice formation increases salinity and upon further cooling, the dense water flows off the shelf and down the continental slope.

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9
Q

Thermohaline Conveyor Belt

A

North Atlantic Deep Water flows south along the western side of the N. Atlantic.
Antarctic Bottom Water is the densest water and flows north along the western side of the S Atlantic.
These two deep currents meet in the S. Atlantic and flow eastward into the Indian and S. Pacific Oceans

The deep water flows from the Atlantic to the Pacific, are balanced by a return flow of warm surface waters from the Pacific to the Indian and back to the Atlantic Ocean.
The combination of these slow deep and surface water flows is referred to as a conveyor belt.

The thermohaline circulation is sluggish compared with the wind-driven circulation.
The entire circulation and replacement of the deep waters takes about 1000 years
750 years for the Atlantic
1500 years for the Pacific

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10
Q

What is an ion?

A

an ion is a charged atom or charged molecule
ions form by adding or removing one or more electrons from an atom or molecule
a cation is a positively charged ion
e.g., sodium ion: Na+
an anion is a negatively charged ion
e.g., chloride: Cl-
sulfate: SO42-
ionic bonds hold crystals together
e.g., sodium chloride (= table salt): NaCl

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11
Q

Why is water sometimes described as a universal solvent?

A

Water (H2O) can dissolve more things than any other natural substance.
It is a polar molecule, that can form hydrogen bonds.
Water is good at dissolving salts
which consist of positively (+) and negatively (-) charged ions
NaCl → Na+ + Cl-

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12
Q

What are the sources of the ions in seawater?

A
Runoff from the continents (weathering of rocks)
Na+ , K+, 
Ca2+ , Mg2+
Volcanic activity (hydrothermal vents)
HS- , Cl-
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13
Q

How does the rule of constant proportions make measuring salinity easier?

A

The rule of constant proportions states that the relative amounts of the various ions in seawater are always the same (e.g., independent of salinity)
e.g.
Chloride = 55.03% of salinity everywhere in the sea
Sodium = 30.59% of salinity everywhere in the sea
the rule holds for other major ions
This allows chloride, which is easy to measure, to be used to calculate salinity
Chlorinity = mass of chloride in a kg of seawater

Conductivity is now commonly used to measure salinity in practical salinity units (psu)
1 psu = 1 ‰ or 1 ppt

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14
Q

Limiting Nutrients

A

The two main limiting nutrient elements for biological production in sea are nitrogen (N) and phosphorus (P).
These are present as dissolved inorganic ions
phosphate: PO43-
nitrate : NO3-
Concentrations of these ions are often very low in surface waters.
Productivity of the oceans often depends on the regeneration (recycling) of inorganic N and P from organic matter.

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15
Q

Vertical profiles of dissolved O2

A

Feature: oxygen minimum zone
located in thermocline
high respiration rate
limited exchange of water

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16
Q

What are Biogeochemical cycles?

A

Pathways by which a chemical element moves through different compartments (called reservoirs).

Examples
Carbon cycle
Nitrogen cycle
Sulphur cycle
Phosphorus cycle
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17
Q

The Carbon cycle

A

CO2 moves between ocean and atmosphere due to physical-chemical processes (Solubility pump).

C is exchanged between the ocean and the biota via:

Photosynthesis removes CO2 from the atmosphere and ocean.
CO2 + H2O + light → CH2O + O2

Respiration releases CO2 to the atmosphere and ocean:
CH2O + O2 → CO2 + H2O

Carbon enters the foodweb via grazing

Carbon is returned to the ocean as CO2 via respiration

Some primary produces will sink when they die, sequestering (storing) carbon in sediments

Dead stuff becomes detritus/POM (Particulate organic matter) which can be:

Decomposed by bacteria, producing CO2, POM, and DOM (Dissolved organic matter)

Feed on by animals, returning carbon to the foodweb

detritus/POM that is not decomposed sinks, sequestering carbon in sediments

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18
Q

The biological pump

A

Phytoplankton and macro-algae (seaweed) fix carbon

Incorporating C into the food chain (grazing) - or releasing it as DOC/POC

Feacal pellets, marine snow, and dead marine organisms sink to the deeper layers

Where they decay consuming dissolved oxygen and giving off CO2.

Upwelling returns this CO2 to the epipelagic.

The biological pump requires the input of nutrients (N, P) to sustain plankton blooms.

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19
Q

Microbes and the carbon cycle

A

Play a critical role in all nutrient cycles

Fixing carbon
Link to the food chain
Also a source of CO2

Microbial action can make carbon inaccessible recalcitrant – RDOM

This sinks and can be stored for 1000s of years

Microbes decompose POM and DOM, producing dissolved CO2

Microbes degrade the most assessable carbon first.

Carbon that is harder to degrade, called recalcitrant carbon (RDOM), sinks before is can be degraded and is stored in sediments.

The balance between how much carbon sinks and how much is releases is critical in global carbon budgets.

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20
Q

The Nitrogen cycle

A

Nitrogen is fixed by bacteria and cyanobacteria

Key players: Trichodesmium spp.
Atelocyanobacterium spp.
(cyanobacteria)

Bacteria and archaea cycle nitrogen between ammonia, nitrites, and nitrates via nitrification (aerobic)

Fixed nitrogen and nitrates are taken into the biota and cycle through the food web

This nitrogen can be excreted as DON, or sink when taxa die (POM).

Bacteria and archaea recycle POM and DON in decomposition (ammonification)

Nitrogen is returned to the atmosphere via bacterial Denitrification (anaerobic)

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21
Q

The phosphorus cycle

A

Not found as a gas (Contrast to Carbon and Nitrogen)

Normally as part of a phosphate ion: PO43-

Found as salts in ocean sediments or in rocks.

Uplift brings ocean sediments to land.

Phosphate becomes available by chemical weathering.

Input to oceans is via rivers.

Enters the foodweb through uptake by plants, algae and bacteria.

Dissolved phosphate is precipitated and sinks into sediments

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22
Q

More on Limiting nutrients

A

Nutrients, particularly N and P, limit the fertility of many undisturbed ecosystems.

P is typically the main limiting nutrient in freshwaters, followed by N.

N is usually most limiting in marine systems, followed by P.

Although CO2 may limit photosynthesis in terrestrial plants, inorganic C is rarely limiting in aquatic systems.

The micronutrient Fe, has recently been found to be the main limiting nutrient over about 1/3 of the ocean surface – Iron fertilisation hypothesis.

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23
Q

Jawless Fish (Agnatha)

A
 cylindrical, elongated body
 cartilaginous skull
 lack vertebrae
 lack jaws
 lack scales
 feed by suction using a round muscular
mouth and sharp teeth
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24
Q

Hagfishes

A
Marine
 Jawless elongate fish
 Lack fins
 No vertebra (sort of)
 Burrow in muddy bottoms at
moderate depths in cold waters
 Feed mainly on dead or dying fish
 Produce slime!
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25
Q

Lampreys

A

Mainly freshwater or costal
- Some migrate to the ocean to mature and bread
in freshwater
 Have unpaired fins
 Redemptory vertebra
 Most species are not parasitic!
- sea and river lamprey attach to fish, rasp a hole
into the victim, and feed on fluids and tissues

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26
Q

Cartilaginous Fishes

Class Chondrichthyes

A

Cartilage skeleton - lighter and more
flexible than bone
 Placoid scales - tough sandpaper like skin
 Moveable hinged jaws
 Ventral mouth with well developed teeth
 Maintain buoyancy through a oil-rich
liver
 Can detect weak electric field to locate
prey - ampullae of Lorenzini

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27
Q

Sharks

A

Fusiform shape (tapered at both ends) - aids
movement through water
 Strong caudal (tail) fin for propulsion
 Two dorsal fins and paired pectoral fins
 5-7 gill slits on each side of body behind the
head
 Most are carnivorous
 Largest sharks are filter feeders (whale
shark - Rhincodon typus)

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28
Q

Ampullae of Lorenzin

A

Sensing organ that uses electroreceptors to
detect the weak electrical currents emitted by
living organisms.
• For example fish hiding under sand.
• Consists of pores connected to gel filled
tubes.
• Also found in other chondrichthyes (e.g. rays)

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29
Q

Bony Fishes

Class Osteichtyes

A
Dominant vertebrates in the sea
 More taxonomically diverse than
rays & sharks
 Wide range of shape, size,
color, swimming ability, feeding
habits, reproduction and
behavior

Skeleton made of bone
 Anterior, midline mouth
 Swim bladder to provide buoyancy
 Scales made of bone, covered by skin and mucus
 Gills protected by a flap of bony plates
(operculum)

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30
Q

Swim bladder

A
Swim bladder (5% of animal’s volume).
 Two gas filled sacs – fish achieve neutral buoyancy by
changing the volume/pressure of the bladder.

Only in bony fish - so what about sharks?
 Altered salt content of cellular fluids (low salt) relative
to seawater.
 High lipid content
- liver may occupy up to 25% of volume of a shark
 Also “flying” using fins for lift

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31
Q

Lateral Line System

A

Gelatin-filled tunnel

  • runs beneath skin along each side from tail to head
  • contains clusters of nerve cells for detecting vibration
  • detects reflections of bow waves off nearby objects, including other fishes

Allows rapid movement near solid surfaces and tight schooling

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32
Q

Body form in relation to feeding and habitat in bony fishes

A

Rover predators (tuna, marlin, blue shark) long torpedo
shaped = efficient swimmers
 Lie-in-wait (ambush) predators (barracuda) – Body
does not taper. Acceleration from strong powerful tail,
(caudle peduncle is wide).
 Flatfish (flounder and plaice - bottom dwelling
 Deep-bodied (butterfly fish) – lots of contact with water
= manoeuvrable, tight turns
 Eel-like (moray eel) fishes are adapted for moving
through crevices

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33
Q

Anadromous and catadromous fish

A

Anadromous: Feed and grow at sea - Reproduce in freshwaters
• Salmon

Catadromous: Feed and grow in freshwaters
 Reproduce at sea
 European and American Eels

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34
Q

The Epipelagic

A
Topmost zone, light is abundant
▪ This is where photosynthesis takes place =
lots of food
▪ You need to see your lunch before it sees
you – or you become lunch!
▪ Speed is king – escape or catch
Adaptations:
- Large eyes
- Strong muscles, fast swimming
- Counter shading
- Streamlined
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35
Q

Mesopelagic environment

A
Sufficient sunlight for vision, but not
photosynthesis
▪ Energy comes from sinking organic
material
- ≈20% of primary production sinks out of the epipelagic
- <5% of primary production reaches 1000 m
Depends on the surface layers for O2
Natural thermocline

Short on food: Most adaptations are
concerned with finding food, or conserving
energy
They also have to stay afloat, within these
constraints
Vision is still a key sense: Seeing in the dark,
and not being seen is key.

Despite the lack of food, as much as 95% of
fish biomass may be in the mesopelagic!

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36
Q

Mesopelagic Fishes

Adaptions to lack of food

A

Common adaptations:

▪ Small size – don’t waste energy
Make the most of every meal
▪ Large mouths
▪ Hinged extendible jaws
▪ Needle-like teeth
▪ Unspecialised diets
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37
Q

Mesopelagic Fishes

Adaptions to lack of food

A

Common adaptations:

▪ Small size – don’t waste energy
Make the most of every meal
▪ Large mouths
▪ Hinged extendible jaws
▪ Needle-like teeth
▪ Unspecialised diets

Carnivores have:
▪ large mouth
▪ extendible jaws & needle-like teeth (e.g.
viperfish & rattrap fish)

Zooplanktivores have:
▪ smaller mouths to rapidly ingest small prey
▪ upward facing eyes to spot zooplankton in
the downwelling light (e.g. lanten fish)

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38
Q

The two strategies of

mesopelagic fish

A

Stay put, sit and wait
2. Go find food
Many mesopelagic fish undergo a diel vertical migration,
moving up to the surface at night (when they are safer from
predators) to find food.
This is the biggest migration on earth.

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39
Q

Vertical migration

A

Following food source (e.g. zooplankton)
‘Safer’ from predators at night
Vertical migrators possess
- well developed muscles
- swim bladder (often filled with lipid rather than gas)
Migrators contribute to the deep-scattering layer
Migrators bring surface production down with them
- enriching food in the mesopelagic
- non-migratory predators feed on them

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40
Q

Characteristics of non migrators

A
non-migrators cope with the reduced food
supply (relative to migrators) by reducing
energy demand
▪ near neutral buoyancy
▪ lack swim bladders
▪ less muscle
▪ watery flesh
▪ soft weak bones
Sit and wait predators - lurk in the dark and wait
for a meal (use of lures)
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41
Q

Mesopelagic Fishes

Adaptions to low light

A
Look up
▪ Large, often tubular, eyes
▪ Upward orientation
▪ Main retina for upward field
▪ Secondary retina for lateral vision

Vision is still important in the mesopelagic, but
counter shading is less effective with less light.
Most fish are looking up at you so common
adaptations are:
• Transparency
• Lateral compression of body to reduce
silhouette

Counter illumination
Counter illumination -
- photophores (light emitting organs) on underside
(ventral)
- bioluminescence - blue light to match spectrum of
downwelling light
Countershading - using coloration (silver
sides and black backs) to blend in.

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42
Q

Bathy & abyssopelagic

Adaptations to Very Low food Supply

A

We see many adaptations that we saw in the
mesopelagic to deal with lack of food:
▪ Week flabby muscles (don’t waste energy)
▪ Large mouth/teeth
▪ Generally slow metabolisms
▪ Reduced or no swim bladder
▪ Bioluminescence

Adaptations seen in mesopelagic fish to low food are generally
exaggerated.
Don’t miss a meal:
▪ large mouth
▪ hinged extendible jaws
▪ long sharp teeth
▪ bioluminescent lure
Don’t waste energy:
▪ sluggish
▪ flabby watery muscles
▪ weak skeletons
▪ poorly developed respiratory and circulatory systems
▪ poorly developed nervous system
▪ lack swim bladders
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43
Q

Sex in the deep sea

A

Hard to find a mate in a sparsely populated
sea. Especially when you don’t move much.
▪ Some species are simultaneous
hermaphrodites
▪ Some use pheromones to find each other
▪ Angler fish have dwarf males that attach to
females (can’t waste an opportunity to mate)

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44
Q

Deep sea benthic fish

A

No natural light here and very little food.
Very little food reaches the bottom
- Food falls (whale carcasses etc)
- Marine snow and fecal pellets (POM)
Covered in a fine muddy sediment.
- bacteria and meiofaunna degrade POM/DOM and make
it available to the food chain
- Fish also scavenge larger food fall directly

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45
Q

Characteristics of deep sea benthic fish

A

Scavengers with strong muscles to move to
food falls
▪ Relatively large and slow growing
▪ Often elongated (eel-like)
▪ Strong muscles (will swim a long way to find
food).
▪ Small or no eyes (few are bioluminescent)
▪ Black/brown in color

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46
Q

phytoplankton: Diatoms

A

unicellular protist
about 40-50% of marine primary productivity
silica cell wall called a frustule
single cells or long chains

capable of rapid growth
tolerant of low light
temperate and polar regions
bloom forming (spring bloom, upwelling)
adequate light
high nutrient concentrations
subject to intense grazing pressure from zooplankton
cysts (dormant stage)
survive for years (e.g. in sediments)
seed for new population growth
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47
Q

Coccolithophores

A

Coccosphere
Cell wall composed of calcium carbonate (CaCO3) plates
Complex, and still poorly understood, life cycle
Often important in tropical waters
Capable of forming large blooms, usually following the diatom bloom

Major source of CaCO3 to the sea floor.
Responsible for extensive CaCO3 rock formations (e.g. White Cliffs of Dover).
Upon dying, coccolithophores produce large amounts of dimethyl sulfide (DMS), a gas that escapes to the atmosphere.
DMS may aid cloud formation and allows sulfur to be transported from the ocean to the land.

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48
Q

Dinoflagellates

A
Dinoflagellates
motile using two flagella 
longitudinal (trailing behind the cell)
transverse (around the cell's middle)
cell wall (theca) consists of cellulose 

Capable of vertical migration in stratified waters
Slow growing, requiring high light
Form cysts that can persist in sediments for years
seed for new population growth when environmental conditions are favourable
Red tides, not due to rapid growth, but rather due to
persistence and
accumulation of toxins associated with defences against grazers

Small contribution to marine primary productivity
None-the-less, ecologically and economically important
Includes about 100 toxic species, producing a variety of effects in man and fish.

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49
Q

Cyanobacteria

A

prokaryotic
blue-green to pink in colour
phycobiliproteins (phycocyanin, phycoerythrin) are major light-harvesting pigments
includes the smallest and some of the largest phytoplankton
Trichodesmium is capable nitrogen fixation - transforming N2 gas into ammonium
Synechococcus accounts for about 20-30% of total ocean primary productivity
Prochlorococcus is the smallest photosynthetic cell (<1 µm diameter)

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50
Q

What are the characteristics of the zooplankton?

A

heterotrophic protists and animals
rely directly, or indirectly, on phytoplankton for food
largest biomass in surface waters (epipelagic)

metazoan zooplankton
Crustacean zooplankton
Chaetognaths (arrow worms)
Pteropods (molluscs)
Gelantinuos zooplankton (coelenterates, ctenophores, salps)

protozooplankton
ciliates, flagellates, amoeboids

Single-celled animals that are grouped based on motility into 
flagellates
ciliates
amoeboids: 
foraminifera
radiolaria

consume bacteria, phytoplankton and other protozooplankton

fed upon by larger zooplankton

mixotrophic protists are both photosynthetic and phagotrophic

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51
Q

Protozooplankton

A

Radiolaria
silicate test
feed using long, thin, retractable pseudopodia
planktonic species may contain symbiotic algae (zooxanthellae)
many benthic species

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52
Q

BACTERIO- and VirioPlankton

A

The term bacterioplankton refers only to heterotrophic prokaryotic organisms in the plankton.
excludes cyanobacteria, which are autotrophic.
includes archaea, which are not bacteria.
Bacterioplankton consume organic matter consuming O2 and releasing CO2.
Bacterioplankton also play important roles in the nitrogen cycle.
Virioplankton are viruses that infect bacteria, phytoplankton, protozoa and other organisms.

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53
Q

Plankton DEFINITIONS

A
Meroplankton = zooplankton that spend part of their life in the plankton and the remainder in the benthos
Holoplankton = planktonic organisms that spend their entire life in the plankton
Ichtyoplankton = planktonic developmental stages of fish
Neuston = organisms associated with the sea surface
Pleuston = organisms that protrude into the air (Portuguese man-of-war)
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54
Q

Primary production

A

Autotroph = an organism that manufactures its own organic carbon using energy from the sun or other sources.
photosynthesis uses light energy (photoautotrophy)
chemosynthesis uses chemical energy (chemoautotrophy)
Primary production = the conversion of inorganic carbon into organic compounds by autotrophs.

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55
Q

What affects Primary production?

A

availability and uptake of nutrients
temperature
affects the rate of metabolic reactions
specific temperature optima and tolerance ranges
intensity and quality of light
net primary production is confined to relatively shallow water depths due to
the high attenuation of light in water
the relationship between photosynthesis and light intensity

Limiting Factors: Nutrients
Nitrogen is the principal limiting nutrient for many pelagic marine ecosystems.
Iron is a limiting factor in the Southern Ocean and other ‘high nitrate/low chlorophyll’ (HNLC) regions.
Diatoms require silicate.
Phosphorus can limit growth of N2-fixing cyanobacteria.

56
Q

What are the requirements for a phytoplankton bloom to take place?

A

nutrients in high concentrations near the sea surface
sufficient light throughout the surface wind-mixed layer to allow net phytoplankton growth, and
the growing phytoplankton population must also escape from the high mortality imposed by zooplankton grazers

57
Q

What are the adaptations and conditions that allow diatoms to persist through winter?

A

diatoms tolerate low light,
mixing keeps these non-motile cells in suspension,
silica cell wall and large cell size provides protection from many protozoan grazers

58
Q

Start of Spring Phytoplankton Bloom

A

phytoplankton growth rate increases due to
the increased availability of solar radiation, and
shallower mixed layer depth

diatoms are poised to take advantage of these favourable conditions because of their

ability to grow rapidly, and
to avoid being eaten by protozoa

59
Q

End of Spring Phytoplankton Bloom: Grazing

A

diatoms provide an excellent food source for crustacean zooplankton

calanoid copepods are the principal grazers of diatoms

adult female copepods may lay up to 50 eggs a day

comb jellies, tunicates and arrow worms become abundant, grazing on the copepods

zooplanktivorous fish become abundant in the declining phase of the spring bloom

60
Q

protozoan zooplankton in summer

A

protozoan zooplankton become increasingly important as the water column remains stratified
protozoa regenerate nutrients
in summer most of the primary productivity depends on

recycling of nutrients between protozooplankton,
bacterioplankton and phytoplankton

with little export in sinking particles

61
Q

Dissolved nutrient concentrations are typically very low in summer.
What adaptations contribute to success during summer?

A

picophytoplankton have high surface area-to-volume ratios and high nutrient affinities
N2-fixing cyanobacteria can tap the atmospheric N2 source
vertically migrating dinoflagellates, because they can migrate to nutrient rich deep waters.

62
Q

Latitudinal variability in primary production

A

Regulated by the same environmental factors and ecological interactions as the seasonal cycle.
Typically nutrient-limited in tropical seas.
Little seasonal variability.
Typically light-limited in polar seas.
Summer bloom.

63
Q

Predation Pressure in the Pelagic

A

One of the major differences between the pelagic and most other environments is that there is virtually no place to hide in the open water.
Exceptional, are clumps of the floating plant Sargassum and other floating debris.
These ‘islands’ often attract animals.

64
Q

How do organisms protect themselves from high predation pressure

A
Body armour (Chitin, CaCO3, SiO2, Cellulose)
Chemical warfare (toxic dinoflagellates)
Camouflage
transparency
colour/shading
counter illumination
Diel vertical migration
65
Q

Vertical Migration

A

There are two broad classes of vertical migration.

Ontogenetic - associated with different developmental stages in an organism’s life history (seasonal).

Diel - a behavioural response that involves a trade-off between:
maximising feeding and
avoiding being eaten.

Diel Vertical Migrations (DVM) often involve upward migration at or after dusk (following isolumes) and downward migration prior to dawn (endogenous biorhythm)

66
Q

Langmuir Circulation

A

Accumulation of buoyant cells in sinking regions of Langmuir circulation, or
accumulation of sinking cells in rising regions of Langmuir circulation.

67
Q

Cetacea

A

Order Cetacea

Odontoceti: Toothed whales: dolphins & porpoises
Mysticeti: Baleen whales: the great whales

streamlined, elongate body to reduce drag
forelimbs reduced to stabilising paddles
hind limbs completely reduced (vestigial bones)
highly muscular posterior ending in a horizontal tail for propulsion - fluke
nearly hairless body
thick subdermal fat layer (blubber)
insulation
streamlining
energy reserve

blow-hole (nostrils) located on back of head
expel CO2 upon reaching the surface
Mysteceti – two
Odontoceti – one 
storage of O2
haemoglobin in blood
myoglobin in muscle  
hearing is the most important sense
sound travels faster and further in water than in air
light is attenuated rapidly
some river dolphins are blind
68
Q

Toothed whales and porpoisesOdontoceti

A
sperm & killer whales, dolphins & porpoises
active hunters of large prey
sonic and subsonic clicking signals
echolocation
communication
social animals

killer whales travel in small pods
dominant female, several females and subordinate males
subordinate males remain with mother’s pod but mate with females from other pods

69
Q

Echolocation

A

Clicks are generated by moving air between internal cavities (air sacs) and are focused using a fatty structure (melon)
Incoming sound is perceived by lower jaw, and transmitted to the inner ear

70
Q

Baleen whales

A

filter feeders having baleen plates instead of teeth

right whales swim slowly = current = zooplankton (<3 mm length) strained on baleen plates

rorqual whales intermittent suspension feeders
-expandable pleats of elastic tissue in throat and thorax
-enclose swarms of zooplankton in mouths and use tongue to force water through baleen
plates to strain prey

71
Q

Migrations

A

most toothed whales do not migrate, but do travel to search for food

many baleen whales

  • feed in polar regions on abundant plankton in summer
  • migrate to warmer regions to breed in winter

still uncertain how whales navigate

  • visual clues when close to shore
  • earth’s magnetic field
72
Q

Pinnipeds (seals)

A

large eyes for vision in low light
hearing most important sense
whiskers are tactile organs used in detecting prey
moult annually
feed on fish, cephalopods, krill, other seals
some bottom feeders (crabs and shellfish)
prey to sharks, killer whales

migrate between feeding ground and haul out sites
mate and rear young on rocky shore areas
suitable beaches are limited in extent, hence large colonies and competition for space
large males arrive first and establish territories, and maintain them by display and combat
dominant males maintain harems
subordinate males at the fringes of the territories of the dominant males

73
Q

Adaptations for diving in animals

A

Adaptations to prevent the bends
-formation of N2 bubbles in blood and joints during rising from depth

collapse lungs before diving
-limits the amount of N2 that can dissolve in blood under high pressure experienced at depth

Adaptations to reduce O2 demand
Heart rate slows (in northern elephant seal, for example, from 85 to 12 beats per minute)
Blood supply to vital organs (brain, heart) is maintained
Blood supply to extremities and intestine is reduced

74
Q

Flow of Energy in Marine Systems

A
Trophic interactions (i.e. predation) lead to flow of energy
Not all consumers eat all prey
Not all consumers eat producers
Energy flow through steps – “Food-Chain”
Each step = Trophic level
Many chains in every web!
75
Q

Tides are one of the dominant forces affecting near-shore life

A

tides alternatively submerge and expose intertidal plants and animals affecting

  • physical stress (heat and water stress)
  • food supply
  • susceptibility to predation

tides drive water circulation in bays

tides trigger spawning in some species

76
Q

What causes the tides?

A

Two forces cause the tides

  • gravitational forces exerted by moon and sun, and
  • centrifugal force due to rotating earth.

Result is the equilibrium tide
-two bulges on opposite sides of the earth.

77
Q

Spring & Neap Tides

A

Spring tides = greatest tidal range (highest high and lowest low) occur during new and full moon

Neap tides have the lowest tidal range (lower high and higher low) occur when sun and moon are at 90o.

78
Q

Micro and Macro tidal

A

The tidal range in the ocean varies from 0 (Caribbean) to 15 m (Bay of Fundy, Canada)

Macrotidal = large tidal range
> 4 m range
British Isles
tidal currents are important in sediment transport

Microtidal = small tidal range
<2 m
Mediterranean sea
waves are important to sedimentary processes

79
Q

Wave formation

A

Waves begin to form as soon as the wind starts to blow
Size of wave depends on
speed and duration of wind
fetch = span of water over which the wind blows
Waves break when their orbital motion feels the friction of the sea bed

80
Q

Vertical zonation in Rocky Intertidal Zone

A

Vertical zonation

  • dominant species occur in distinct horizontal bands
  • along a gradient from fully terrestrial to fully marine.

Zones are compressed in protected areas with low tidal range.

On open coasts, the zones are extended by wave splash.

81
Q

Upper Intertidal Zone

A

Lichens (Verrucaria)
-black blotches

cyanobacteria (Calothrix)
-dark green mats

Tufts of algae

  • algae (Ulothrix)
  • brown (Pelvetia)
  • red (Porphyra)

Periwinkles (Littorina) and Limpets (Patella) graze on algae

Top predators include terrestrial animals (e.g., birds, rats) and marine predatory snails

82
Q

Middle Intertidal Zone

A

Acorn barnacles form bands at upper end (Chthamalus, Balanus & Semibalanus)

Lower in the mid-intertidal are

  • mussels (Mytilus)
  • gooseneck barnacles (Pollicipes)
  • rockweed (Fucus & Pelvetia)
83
Q

Lower Intertidal Zone

A

Mussels & barnacles are rare due to high predation pressure

Zone is dominated by seaweeds

Sea urchins are common grazers

84
Q

Rocky Intertidal Zone: Causes of Zonation

A

Four major factors involved in zonation are
larval settlement (dispersal & colonisation)
physiological tolerance of stress
Predation (e.g. by seastars, birds etc)
competition for space

85
Q

Physical Stress - Wave Action

A
Epifauna = live on the surface
Some animals avoid being dislodged by seeking shelter in cervices
behavioural response
Adaptations to hold on
holdfasts in seaweeds
mussels glue themselves 
on using byssal threads
limpets and chitons (right) cling 
on using a muscular foot
86
Q

Physical Stress - Temperature

A

extremes of temperature during periods of emergence
sea has more stable temperature than air due to high heat capacity of water
adaptations include:
ability to tolerate high temperature range
use of light colours to reflect sunlight
sheltering in moist places to allow evaporative cooling (under rocks… like the below)

87
Q

4 classifications of estuaries

A

Drowned river valley (= coastal plain estaury)
formed when sea level rose at end of last ice age
Examples include
Thames Estuary
Delaware Bay
Chesapeake Bay

Fjords are estuaries in deep valleys cut by glaciers
formed when sea level rose at end of last ice age
high latitudes including Norway, Southwestern Chile

Bar built estuaries: sediments build up to form barrier sand bars & islands
North Carolina Bays behind protection given by Cape Hatteras
North Sea coast of the Netherlands & Germany

Tectonic estuaries are formed by subsidence

88
Q

Estuarine circulation

A

Freshwater flows out near the surface
Saline water flows in near the bottom
Salt wedge = a layer of denser, saltier seawater that flows along the bottom of the estuary
There is mixing at the interface between salt and fresh waters due to wind, tides and river flow

Tidal prism = the volume of water that is moved in and out of an estuary with the change in the tides

89
Q

Salinity and biodiversity

A

Mid-salinity zones of estuaries are often regions of low taxonomic diversity

Estuarine species must be able to tolerate salinity fluctuations

Stenohaline species tolerate low salinity range

Euryhaline species tolerate a high salinity range

Osmoconformers allow the salinity of their body fluids to change with external salinity

Osmoregulators keep the salt content of their body fluids constant

90
Q

Primary producers of muddy shores

A

Diatoms (Bacillariophyceace)

Cyanobacteria (Cyanophyceace)

  1. Euglenoids e.g. Euglena
  2. Macroalgae e.g. Ulva, Enteromorpha
  3. Angiophyceace e.g. Zostera (eel grass)
  4. Chemoautotrophs e.g. Sulphur oxidising bacteria
91
Q

Salt marshes

A

Salt marshes are intertidal habitats dominated by cord grasses (Spartina spp.)

Spartina typically spreads by vegetative growth of rhizomes (root networks within sediments)

Often a marsh consists of one or a few clones

Roots and rhizomes trap & stabilise sediment, leading to build-up of the marsh and infilling of estuaries

Spartina blades contain siliceous deposits that deter grazing

Typically, <10% of blade growth is consumed by grazers; the remainder fuels detritus food chains

Salt marsh creeks are often productive nurseries for marine fish species.

92
Q

Spartina anglica

A

Spartina anglica originated in southern England in about 1870 when the European cordgrass S. maritima hybridized with the introduced American S. alterniflora

more vigorous than either native species
-rapid rate of growth, high fecundity and aggressive colonisation

planted for coastal erosion control

  • rhizome systems bind coastal mud
  • stems increase silt deposition, thereby assisting in land reclamation from the sea
93
Q

Mangroves

A

mangroves are salt tolerant tropical and subtropical trees

salt is secreted from leaves by salt glands

mangrove forests (mangal) support detritus-based food webs

root systems are adapted to anoxic, waterlogged conditions

oxygen is transported to below ground tissue by upward facing pneumatophores allowing aerobic metabolism in an anoxic sediment

fine roots take up nutrients

prop roots support the body of the tree

94
Q

Seagrass Meadows

A

Like cord grass (Spartina), these grasses have extensive rhizome systems

Reduced turbulence traps fine sediments

Roots extract nutrients from sediments

Shoots obtain CO2 from the water column

Beds expand largely by vegetative growth of shoots from the rhizomes

95
Q

Kelp

A

Kelp are macroalgae (brown seaweeds)
a holdfast attaches the kelp to hard bottoms
the stipe provides support allowing the blades to reach the sea surface
buoyancy is provided by pneumatocysts

96
Q

Macrocystis Life Cycle

A

Alternation between large sporophyte (diploid, spore-producing)

and microscopic gametophyte (haploid, gamete producing) stages

Spores settle on bottom, develop into male or female gametophytes

Sperm produced by male gametophyte fertilize eggs produced by female gametophytes to produce diploid sporophytes

97
Q

Climate Variability and Kelp Ecology

A

Kelp growth off California depends on upwelling of cold nutrient rich waters

Upwelling, in turn, varies with large scale patterns in the winds and ocean circulation

Coastal upwelling is associated with along shore winds that blow towards the equator on the west coast of continents

98
Q

Coastal Upwelling

A

occurs as a result of Ekman transport of surface waters

the Coriolis force causes surface water movement to be deflected to the right of the direction of the wind in the N. hemisphere (to the left in the S. hemisphere)

thus, when winds blow from the N to S off California (or S to N off Peru), the surface layer flows off shore

99
Q

What is El Ni~no ?

A

El Ni~no Southern Oscillation (ENSO)

The term El Ni~no was coined in 1891 to describe an event that devastated the Peruvian fishery

  • warm surface current flowed over the coastal waters off South America near Christmas 1891
  • thus associated with Christmas (hence El Ni~no, the (Christ) child)

It is a short-term weather disturbance that influences wind patterns, sea surface height and precipitation, and affects marine and terrestrial plant and animal populations

100
Q

What is the Southern Oscillation?

A

a change in atmospheric pressure distributions that accompany El Ni~no

normally, atmospheric pressure is

  • high over the eastern Pacific and
  • low over the Indian Ocean
  • prevailing winds blow from east to west
101
Q

What happens during a La Ni~na Event?

A

central Pacific trade winds increase in intensity
the South Equatorial current speeds up
upwelling of cold, nutrient rich water increases
surface waters off Peru & California cool down

102
Q

Soft Bottom benthic communities

A

Soft bottoms are those into which organisms can burrow
sand, silt, and/or clay depending on
how much water motion there is and
the source material for the sediment

Exposed places with strong waves have course sand or gravel bottoms
Calm, sheltered areas with little wave action often have muddy bottoms
Soft bottoms are unstable and shift in response to
waves
tides
currents

Sediments may be
vegetated (e.g., sea grass beds)
or unvegetated (e.g., sands, muds)

103
Q

Sediment Characteristics

A

Sediment is a mixture of
inorganic particles
decomposing organic mater
labile = readily degradable carbohydrates, lipids and proteins, with high extractable energy content (freshly produced)
refractory = what is left over after labile material has been extracted - decomposition products including cellulose, chitin
microorganisms
bacteria, algae, protozoa

104
Q

Sediment type affects physical properties

A

course sediments are well drained and can dry out

fine sediments retain water

poorly sorted sediments can become water logged

O2 (oxygen) content is determined by
-the microbial activity (respiration) in the sediment -which consumes O2
and by the extent of mixing which introduces O2

Anoxia develops where
-the rate of O2 consumption by respiration exceeds the supply of O2 by
percolation of water through the sediment
mixing of the sediment by waves or currents

Anaerobic bacteria break down organic matter using

  • sulphate (SO4) instead of O2,
  • produce hydrogen sulphide (H2S)
105
Q

Classification of soft bottom fauna

A
Meiofauna
live in the interstitial spaces between sediment grains
Macrofauna
Where they live:
Epifauna live on the sediment surface.
Infauna burrow within the sediment
How they feed:
Predators & scavengers
Deposit feeders:
Surface feeders
Head-down (in sediment) feeders
Filter feeders
106
Q

The main source of organic matter to the bottom is via a Rain of Detritus

A

faecal pellets, chitinous exoskeletons, and marine snow are produced by plankton in the epipelagic
much is remineralized (broken down) along the way by bacteria and meso- and bathypelagic organisms
sediment traps (cylinders placed at different depths in the water column) are used to sample this material
there is a pulse following the spring bloom in temperate waters
it can take months for this material to reach the bottom

107
Q

Food Supply to the Deep Sea

A

Food falls of dead fish and sea mammals provide “hot spots” of activity on the deep-sea bed.
Deep-sea fishes and invertebrates can home in on these food falls.

108
Q

Hot Vent Communities

A

high biomass of clams, crabs, mussels, limpets and tube worms
most feed on mats and suspensions of free-living sulphide oxidising bacteria
dissolved H2S from the vents provides energy that supports these communities
chemosynthesis uses the energy from oxidation of H2S to fix CO2 into organic matter

109
Q

Zooxanthellae

A

(zooxanthellae) are essential for reef-building by hermatypic corals
corals that lack zooxanthellae are ahermatypic and generally do not form reefs
zooxanthellae contribute to success of hermatypic corals by
removing CO2 which promotes CaCO3 precipitation
providing organic matter to the host
zooxanthellae benefit from
stable environment
freedom from predators
nutrients supplied by host

110
Q

Factors Affecting Reef Growth

A

high calcification only occurs in warm water
zooxanthellae require light
rapid growth is limited to depths <50 m
tidal exchange and coastal currents bring nutrients and zooplankton to the reef
suspended solids (turbidity) reduce light and smother coral polyps
strong waves can damage corals
role of tropical storms
parrot fish and burrowing/boring invertebrates attack living corals creating rubble

111
Q

Types of Coral Reefs

A

Fringing reefs develop near shore, throughout the tropics, where there is a hard surface for larval settlement.
Barrier reefs are separated from the shore by a lagoon.
The lagoon is protected from the waves, and has a soft bottom.
An atoll is a ring of reef, sand bars and islands surrounding a central lagoon.

112
Q

Fringing Reef

A
susceptible to damage by sedimentation, pollution and freshwater runoff. 
reef flat is 
sand, mud and coral rubble.
living hard corals
seaweeds, seagrasses and soft corals.

reef crest is subject to intense wave action
reef slope is steep with dense coral cover

113
Q

Barrier Reef

A
Example:  Great Barrier Reef (Australia)
consists of
back-reef slope,
reef flat, 
reef crest and 
fore reef slope
separated from shore by a lagoon

fore reef slope is steep with dense coral cover

114
Q

Competition for Space amongst Corals

A

Fast-growing corals grow up and branch out to collect sunlight
also shade their neighbours.
Other species attack their neighbours with
mesenterial filaments (used for digeston and absorption of prey) or
sweeper tentacles that are loaded with nematocysts.
The most aggressive species are slow-growing massive corals.

115
Q

Food chain

A

the steps of energy transfer from primary producers through consumers.

Energy is lost at each step in a food chain.

The longer the food chain, the lower the biomass and productivity of the top carnivores.

116
Q

Open Ocean (Pelagic) Fisheries

A

low primary productivity
long food chains to the commercially important top carnivores (sharks, tuna, salmon)
Temperate and Equatorial open ocean regions
Southern Ocean

117
Q

Managing the impact of fishing

A

Managing effort
limiting catch per boat
limiting total catch (close fishery once limit is reached)

Restrictions on access
number of boats
length of season
size of fish that are allowed to be caught

Restrictions on gear
power of boats
mesh size
prohibition of some types of gear

118
Q

What is pollution?

A

The introduction by human activity of substances that decrease the quality of the marine environment

An oil spill is pollution, a natural oil seep is not.

Humans have affected the entire marine environment from beaches to the deepest oceans.

119
Q

Atmospheric deposition

A

Sources:

  • Nitrogen oxide from burning fossil fuel
  • Volatilisation of nitrogen from fertilizer and animal waste
  • Biomass burning

Input via wet (rain) and dry (dust) deposition.

This is the primary source of anthropogenic nitrogen into the open ocean

120
Q

Land runoff

A

Fertilizers washed off from agricultural land into rivers

High in nitrogen and phosphorus
Also toxicants such as pesticides and pesticides

Increase in extreme weather events means this is likely to get worse. (Christensen & Christensen, 2003)

The primary input into costal systems

121
Q

Sewage

A

Sewage is the wastewater generated by domestic and industrial activity.

Contains high amounts of organic nutrients (N and P) and chemical toxicants:
Drugs (pharmaceutical & illicit)
Cleaning products
Heavy metals
Pathogens

During heavy rain untreated sewage is directly discharged into rivers, and untimely into the marine environment.

122
Q

Ecological impacts of eutrophication?

A

Shift in the balance of an ecosystem from slow growing corals or seagrass to fast growing macro- or microalgae.

Algal blooms
Increased phytoplankton
Increased benthic and epiphytic algae
Oxygen depletion
Turbid water
Decreased biodiversity
123
Q

Impacts of cultural eutrophication

A

Harmful algal blooms
Toxic algae
Shellfish poisoning

Anoxic (no O2) or hypoxic conditions (low O2)
Fish kills (suffocation)
Loss of harvestable shellfish/fish

Odour & loss of cultural value
H2S is produced when SO42- replaces O2 in the decomposition of organic matter

124
Q

Hypoxic zones

A

In extreme cases eutrophication can lead to permanent or seasonal hypoxic zones.

This occurs when large amount of phytoplankton sink and are decomposed by bacteria, depleting oxygen

Animals either leave the area or die, undermining ecosystems and fisheries

Hypoxic zones are a natural phenomenon but are increasing in size and number

125
Q

Effects of microplastics on marine animals

A

Plastics break down (e.g. exposure to UV) and enter the food chain when consumed by filter feeders such as jellyfish and mussels.

Negatively impact fitness due to lack of energy pay off.
Toxic additives used in the manufacture of plastics can affect endocrine, reproductive and/or immune systems.
They also for agglomerations with other organic pollutants (Pesticides, PCBs) facilitating bioaccumulation in fatty tissues.

Plastic pollution has potential to impact biogeochemical cycles such as the biological pump

126
Q

Biomagnification

A

Organic pollutants such as:
chlorinated hydrocarbons (many pesticides)
polychlorinated biphenyls (PCBs) manufacture of plastics
dioxins and furans

As they can dissolve in fat they are very persistent once ingested, and are then accumulate up the food web.

Due to biomagnification can reach high concentrations in top predators.

127
Q

Biomagnification and bioaccumulation def

A

bioaccumulation is a build up of a given toxin over time due to PROLONGED exposure

Biomagnification increases in concentration UP the food web

BOTH are dangers to Human Health and Marine Life

128
Q

Effects of heavy metals

A

Direct mortality/toxicity (e.g. copper)
Effect sense organs in fish (smell), impacting:
-predator avoidance
-foraging
-schooling
-navigation (delayed migration in salmon)
-Breeding (finding mates/delayed spawning salmon)

Slow growth
Increased disease
Antibiotic resistance via co-selection

129
Q

Noise pollution

A

Propeller driven ships

Oil, & gas uses seismic surveys that create pulses of intense noise.

Anti-submarine measures and sonar generate very loud short pulses of sound that may damage the brain and other organs.

Offshore construction (e.g. wind farms) requires prolonged pile-driving and underwater use of explosives.

130
Q

Thermal pollution

A

Seawater is used as a coolant in power plants, oil refineries etc. – leading to elevated temperature around discharges.
Fish may be attracted to these areas
In tropics, most organisms live in waters that are close to their upper temperature tolerance limit.
Fish that should migrate don’t, but are venerable to cold shock if the hot water is disrupted
Reduces dissolved oxygen levels
Localized change in plant communities as macroalgae/seagrass are replaced with phytoplankton

131
Q

Radioactive waste

A

Low level radioactive wastes are discharged to environment, and high levels of radioactivity may be released during accidents.
Low-level radioactive waste has been discharged into the Irish Sea from Sellafield since 1952.
Greenpeace described the Irish Sea as the most radioactively contaminated sea in the world.

132
Q

Impacts of oil pollution: toxicity

A

Inhibit growth of phytoplankton/primary producers
kill invertebrates and fish
Increase susceptibility of fish to disease
Cause hypoxia and anoxia as oil is degraded (what is this similar too?)
Destruction of habitats
Bioaccumulation of toxic components from water, sediments, or food.

lose their insulating ability,
leading to death of mammal and birds due to exposure.

133
Q

Microbes matter!

A

Marine phytoplankton perform 50% of global photosynthetic CO2fixation
They also release CO2!
Nutrient re-cycling, underpinning food webs.
They are venerable to change:
Microorganisms have fast turnover rate, for example compared with trees (days versus decades).
how microbial communities have/will respond to change is uncertain

134
Q

Sink or Source

A

The balance between marine microbial autotrophy and heterotrophy will be of key importance with a changing climate - (Sarmento, Philos Trans R Soc B Biol Sci . 2010).
There is evidence that the balance may shift towards heterotrophic processes as the climate warms - (Vaqué, Environ Res Lett 2015; 10)
This could result in a feedback loop between CO2 production and microbial heterotrophic activity!

135
Q

Effects of ocean acidification on Fish

A

Lower pH of fish blood (acidosis). This costs energy to correct and reduces fitness.

Changes in behavior e.g. reduced predator avoidance and have trouble navigating. (See Moodle for the debate)
corrosion of placoid scales in sharks

136
Q

Changes in habitat due to acidification

A

Tropical seeps
dead reef substratum (yellow) is eroded
structural complexity of living corals declines
algae proliferate
sub-tropics
loss of hard coral/coralline algal cover
diverse macroalgal communities replaced by turf algae.
temperate coasts
loss of brown algal canopy cover.
coralline algae decline as turf algae proliferate

In each case, calcified invertebrates (sea urchins) become fewer and smaller

137
Q

Problems with the data used for fishery management

A

Difficult to assess stock size because of
patchiness in distribution
differences in efficiency of capture methods
intermixing of stocks on feeding grounds
Usually assessed by landings divided by effort (Catch per Unit Effort = CPUE) which reflects
population size
spatial distribution of the population
intensity of fishing effort
number of fishing boats, their size and fishing hours
However, fishing is not random
Effort and catch are self-reported by fishers
verified (sometimes) by inspectors