Fragen

Re = (a * v * ρ) / ɳ -> ratio of inert to viscous forces

a = length (m)

v = velocity (m/s)

ρ = density of liquid (kg/m3)

ɳ = dynamic viscosity (kg/m/s)

Inertial force (newtons second law)

Viscous force F = (ɳ * S * v) / a -> S = area

R>>1 : dominance of inertia -> turbulence flow

R<<1 : domiannce of viscosity and drag, laminar or no flow, diffusive boundary layer

water viscosity > air viscosity -> less mixing, more stratification -> less temporal variability, steeper gradients

Stokes law:

-> radius*density difference*shape

-> sinking speed increases with size, relative to weight, and density, form drag is a bit less important

• light intensity

• wave length

temperature

light

Circumpolar Antarctic Divergence

Balance between Coreolis and pressure gradient forces (sea level, temperature, salinity), direction of flow is orthogonal to balance between pressure gradient and coreolis force

• Wind stress on sea surface (wind-driven circulation): wind stress is transmitted downwards due to turbulence, internal friction in upper ocean -> increasing current angle with increasing depth (Ekman spiral) -> net transport to the right of the wind in NH -> gyre circulation

• Solar radiation on earths surface is unequal -> on rootating earth, wind follows contours of constant pressure (geostraphic balance), pressure distribution changes -> three wind circulation cells in each hemisphere -> trade winds -> Ekman transport causes upwelling events

• Buoyancy flux between ocean and atmosphere (thermohaline circulation): Saline cold water is denser than warm fresh water -> sinks down -> relatively saline water fills the void, cools, increases density -> northward warm water transport southward deepwater transport

Because the concentrations of Na and Cl are conservative as they are not used up by any biological processes; Ca and HCO3 are used up during the process of calcification

The time it takes for the entire reservoir of this element to be replaced once (mean conc*ocean volume/inflow per year)

a) Na>Ca>Si>Fe

b) Cl>SO4>NO3>HCO3

• Conservative: slight decrease in subsurface, otherwise stable concentration throughout water column

• Bioactive/-limiting: depletion in surface, highest concentration at ~1000m

• Particle-reactive: stable concentration throughout water column in Atlantic, increase towards depth in Pacific

-> great ocean conveyor belt: deep water formation in North Atlantic, transport through Pacific, next contact with atmosphere (gas exchange) in North Pacific, transport back to North Atlantic as warm surface water

-> Pacific deep water is older and had more time to accumulate bioactive or particle reactive elements sinking down from above

Ca: bio-limiting

Ba: particle-reactive

N: bio-limiting

P: bio-limiting

Mg: conservative

biological pump: CO2 uptake by phytoplankton, CO2 release by respiration, transport of organic C to depth via sinking, release in deep waters by respiration

physical pump: driven by thermohaline circulation, CO2 uptake at higher latitudes due to deeper water formation, CO2 release in tropics due to higher ventilation/ equatorial upwelling

regenerated production: recycling of released nutrients in euphotic zone by phytoplankton uptake, NH4+

new production: supported by external nutrient input e.g. N-fixation by diazotrophs, NO3-

• biological carbon pump is driven by PP, i.e. uptake of CO2 and formation of OM (CH2O)

• carbonate counter pump is driven by CaCO3-formation, i.e. Ca + HCO3 —> CaCO3 + H2O + CO2

• difference is CO2 uptake/release and export products CaCO3 and CH2O

a) Nitrogen fixation by diazotrophs -> supported by high Fe input and strong P depletion

b) Loss of N during remineralization (denitrification, respiration) when primary production is high

A bloom of coccos would lead to a reduction of ppCO2 as they are using CO2 for shell formation while a bloom of diatoms would result in high respiration rates

• energy (ATP)

• carbon (CO2, CH4 or C)

• liquid water

• various elements (N, P, S, Na)

Process by which prokaryotes synthesize C3 (glycerate), compounds from C1 (CH4) compounds using chemical energy

1) ammoinia is converted to intermediate hydroxylamine, the nitrite by nitrosomonas and nitrosococcus

2) nitrite is converted to nitrate by nitrobacter

ammonia oxidation under anaerobic conditions: NH4 + NO2 —> N2 + 2H2O

low O2 conditions (anoxic, microoxic conditions), have to avoid being incrusted into Fe3+ minerals

Carboxylating enzymes links inorganic CO2 or HCO3 with organic acceptor molecule which needs to be regenerated in the following steps

Microorganisms are the most abundant organisms on earth but only 1% can be cultivated

autotrophy: synthesis of nutrients by uptake of sunlight and chemical energy

heterotrophy: uptake of nutrients through organic matter (primary producers)

• oxygen sensitivity

• temperature tolerance

• requirements for metals (Fe, Cu, Ni, Mo)

• used CO2 species

• need for coenzymes

• availability of reducing compounds

The CCB cycle has an upper limit of 70-75°C and functions under oxic conditions, while the WL cycle does not have a temperature restriction (even hyperthermophyles) requires anoxia and high abundance of metals

RubisCO and Phosphoribokinase

Acetyl CoA + Ferredoxin(red) —> pyruvate + Ferredoxin(ox)

Calvin Benson Bassham cycle, reductive CoA-pathway

• diel (24h day and night)

• diurnal (during the day, cloud cover)

• seasonal (summer and winter)

• lattudinal (equator and arctic)

phytoplankton biomass varies throughout year and regions: continuous in tropics, higher in spring in north temperate regions, peak in summer in north pllar regions

Wavelength: PAR 300-720

attenuation coefficients of water, plankton, particles (turbidity)

-> climate change increases terrestrial DOC input (brownification of lakes and sea water) due to permafrost melting, more rain, increase of plant debris at higher temperatures

• Chlorophyll: green coloured, absorb blue and red light, fluorescence maximum at 680nm, light harvesting

• Carotenoids: red-yellow pigmented, absorb blue-green light, no fluorescence, light-harvesting and photoprotective

• Phycobilin: red-pink coloured, absorb green-yellow light, fluorescence at 570nm, light harvesting

• glacial melting

• rain

• release of plant debris

• weathering

• Shows whether cell growth is light limited/saturated

• used for quantifying and modelling of marine phyttoplankton productivity

A -> light limitation zone

B -> light saturation zone

C -> photoinhibition zone

D -> light compensation point

E -> lightt at max. photosynthetic rate

Y>0: net production

Y<0: respiration

• Reduction of antenna size: lower efficiency at lower irradiance due to reduced absorption but also less increase at high irradiation due to reduced photoinhibition

• Improved RubisCO function: same efficiency at los irradience, higher efficiency at higher irradience due to increase in catalytic rate and decreased photoinhibition

• Improved pigment composition: higher efficiency at lower irradiences due to utilisation of alternative wave lengths

Photoacclimation: phenotypic response, changes in pigment composition/concentration and enzyme activity.

-> maintains constant photosynthetic efficiency under varying light conditions

Photoadaption: genetic change (mutation/species selection), species have different pigments/enzymes/biogeochemistry at identical light conditions

Net rate of organic matter produced = gross primary production via uptake of inorganic carbon - respiration

-> incubation in light and dark bottle with 14C and O2 incubation measurements

• Carbon allocation inside the phytoplankton cell (storage in mitochondria)

• Extracellular release of OM

• The survivalist: high N:P ratio -> can sustain growth with limited resources via resource acquisition (proteines, pigments, enzymes)

• The bloomer: low N:P ratio -> adapted for exponential growth, contains growth machinery (rRNA)

• N:P near Redfield ratio -> balances growth and acquisition machinery

• coccolithophores

• pteropoda

• foraminifera

• diatoms

• radiolaria

• sponges

• Lag phase -> no diatoms

• exponential phase -> growth

• stationary phase -> maximum

• declining phase

-> 21 days

Yield of plant mass is proportional to the amount of limiting nutrient (nutrient that runs out first)

Sources:

• river

• dust

• hydrothermal vents

• weathering

Sinks:

• sediments

• air-sea interface

• Phaeophyta (brown algae)

• rhodophyta (red algae)

• chlorophyta (green algae -> plants, sea grasses -> plants)

Intensity of dark respiration: low if all cells are photosynthetically active (thin thalli e.g. crustose algae)

Intensity of photorespiration: low if CO2 uptake is efficient (particularly in green macroalgae)

Ulva can directly transport HCO3- into their cytoplasm with carrier enzymes. This allows C accumulation additionally to to transformation of CO2 to HCO3- by carbonic anhydrase

• thick epidermis

• cell walls with high tannin concentration

• thermal dissipation

• specialized amino acids

• light exposure

• air exposure

• Wave energy

• substrate solidarity

• salinity

• nutrient availability

• pollution

• eutrophication

• Greene algae -> eutrophication, shallow water -> Baltic Sea

• Brown algae -> cool/sunny/nutrient rich -> kelp

• Red macroalgae -> warm/shady/oligotrophic -> crustose algae

• Leathery/filiform/calcerous -> sensitive to herbivory/high productivity/early successional species

• crustose -> insensitive to herbivory/low productivity/late successional species

• Roseobacteria and Flavobactteria break down DMSP produced by phytoplankton to DMS and MeSH

• bacteria are highly motile and have a positive sense for degraded phytoplankton/DOM

• concentrated food input to deep seawhich is. usually nutrient deprived

• one carcass = carbon sinking of 100-200 yrs per hectar at abyssal sea floor

• complete degradateion is a matter of many years and includes all kinds all kinds of organisms from carnivorous fish and crabs to microbes and worms

1. mobile scavengers remove soft tissue (hag fish/sharks/rat tail fish)

2. enrichment opportunist stages: colonization by dense assemblages of worms and crustaceans

3. sulphophilic stage: chemoatotrophic and heterotrophic microbal assemblages

corals and algae produce mucus (DOM) -> uptake by sponges -> release of POM by sponges upon death -> uptake of sponge. POM by detrivores

-> 1/3 of sponge biomass is microbial (cyanobacteria, choanocytes, pinacocysts, archaeocytes)

• Proteobacterium

• 25% of all microbial plankton

• carbon oxidizing bacteria (DOM)

• Aerobic

• free living

• chemoautotrophic

• small genome (1-3Mb)

archeae have metabolism of bacteria and replication machinery of eukaryotes

• P. furiosus: Hyperthermophiles -> fast growth at 80-103°C, optimum at 100°C)

• N. archaeum: symbiosis/parasistism with bacteria, tiny cell, tiny genome (400Mb)

• S. solfactorius: growth optimum at 80°C and pH=2-3, grows on sulfur, extremely stable enzymes

it is a relatively conservative sequence that can be used to identifiy taxa independently from cultivation

Bacterial clade with no cultivated representatives that have been discovered by metagenomic sequencing

ppO2 increases during day time due to 2x photosynthesis (hyperoxia), decreases during night time due to respiration (hypoxia/anoxia)

muscle isn’t dependent on oxygen as it is only used for propulsion to make quick movemements BUT anaerobic mettabolism produces H+ alongside ATP and endproducts -> muscle gets sour

• bivalves (A. islandica & A. borealis) due to their exceptionally high tolerance to anoxia

• massive glycogen reserves

• alternative anaerobic endproducts decrease metabolic heat production by 99%

-> 100x extended survival time by metabolic reduction, 3x extended by alternative endproducts = 300x longer survival time with same amount of glycogen reserves

• Exhaustion of fermentable substrate

• ATP and pH fall

• vital cellular processes fail

• membrane rupture

• cell death

Respiratory coefficient = CO2 formed/ O2 consumed

proteins: (+) more oxygen efficient than fats

(cephalopods) (-) can be less dense packed than fats

fatty acids: (+) less mass to carry, lower body volume, reduced friction -> energy expenditure

(turtles) (-) metabolic rate is not sustainable for a long time

high water solubility, low production costs compared to urea -> excretion via gills ion transporters, gas channels

sensing of food particles -> engulfment -> food vacuole (phagosome) -> intracellular digestion via fusion with digestive enzymes (phagolysosome) -> exocytosis of undigested material

Simplified system of postulates, data, mathematical descriptions of an entity or state of affairs

Test hypotheses, concepts, ideas: causes of ice ages, mass extinctions, deoxygenation

Isolated observations into coherent context: reanalysis products, process studies (mesocosms)

Scenario simulations/predictions: consequences of human action

Michaelis-Menten model provides a relatively small curvature because it assumes photosynthesis = growth; other models take into account that unused photons are lost and cannot be recycled whereas unused nutrients can be recycled/ stay in the system -> Michaelis-Menten model represents growth limitation by nutrients while growth limitation by light has a steeper initial linear slope and more sudden saturation

Ocean is not uniform but homogenous, concentrations are affected by advection/diffusion/gas exchange/redox chemistry

Prokaryotes…viruses…

10^-8 – 10-6m (phages: 10-7 – 10-8m) so on the boundary between DOM and POM (0.2 µm)

Tiny agents made up of DNA or RNA that require a specific host cell to replicate

lytic: phage attaches to host cell, injects DNA -> phage DNA circularizes & enters lytic or lysogenic cycle -> new phage DNA and proteins are synthesized and assembled into virions -> cell lyses, releases phage virions

lysogenic: circularized phage DNA integrates within bacterial chromosome by recombination (prophage) -> lysogenic bacterium reproduces normally (cell division) -> daughter cell has virus genes -> occasional excision of prophage from bacterial DNA & initiation of lytic cycle

Kill bacteria -> release carbon (10 billion tons/day)

change host metabolism/gene expression (control)

gene transfer

dsDNA

rRNA as universal, conserved, variable regions that are phylogenetically informative and can be used for molecular analysis; previously: 2-kingdom hypothesis (1900s) with only animals and plants, 5-kingdom hypothesis with monera, protists, plants, fungi, animals

gene trees don’t always reflect species trees due to horizontal gene transfer, gene duplications result in paralogous genes, composition changes among genes, mutations

Archaeal (Lokiarchaeota as gap-filling between prokaryotes and eukaryotes) -> from hydrothermal vents

Cyanobacteria:

chlorophyll, some heterocysts for N2-fixation

Heterocontophyta: 

2 heterokont flagella, diatoms (epitheca/hypotheca Si-cell wall, Pseudonitzschia), silicoflagellates (internal Si-skeleton, only marine)

Prymnesiophyta:   

2 flagella, 1 haptonema, coccolithophorales (CaCO3, Emiliana), prymnesiales (Wellenschaum)

Chryptophytes:                  

2 hairy flagella, no cell wall, barely dominant in ocean, small, Teleaulax

Dinophytes:           

flagellates, “naked” (Gymnodinium) and armoured (Dinophysis, thick cellulose plates)

Chlorophytes:                    

green algae, equal flagella, ancestors of higher plants

Phytoplankton, Zooplankton, Bacterioplankton, Mixotrophs

Pico: <2µm

Nano: <20µm

Micro: 200µm

Meso: <2mm

Macro: <20mm

Mega: >2cm

laminar flow around particle, diffusive boundary layer, water sticks to cell, viscous forces dominate, sinking velocity follows Stoke’s law (Re < 1)

More molecules can encounter cell surface in relation to volume, cell can host more nutrient transporters in relation to volume -> small cells have higher affinity and competitive advantage in low nutrient conditions

proto-: mouth opening of gastrula stays mouth, secondary anal opening, spiral cleavage of cells

deutero-: mouth opening becomes anus, secondary mouth opening, radial cleavage of cells -> chordata

bilateria: bilateral symmetrie, triploblastic (ecto-, endo-, mesoderm) -> spiralia, ecdysozoa, deuterostomia

spiralia: animals that evolved with spiral cleavage of cells around axis -> annelida, mollusca, plathelmintha

ecdysozoa: process ecdysis (moulding) -> nematoda, arthropoda, tardigrada

roles that organisms fulfil in different habitats and community compositions, based on traits that are essential for ecosystem stability

suspension feeders, predation, filter feeding, grazing

percent of particles that are retained by filter apparatus during one passage

bivalves have 100% retention efficiency at 5µm, sponges at <1µm

bivalves have cilia in lamellae that are specialized on catchment and transport, sponges have filtration chambers with choanoflagellates

cilia, cnidocysts/cleptocnides, radula, choanoflagellate chambers, tentacles

not a common ancestor but similar evolution of several monophyletic groups:

Myxini (hagfish, secondary reduction of vertebra to chorda dorsalis), Petromyzonta (lampreys/agnatha, +vertebra), Chondrichthyes (Gnathostomata, sharks/rays/chimeras, +jaws), Actinopterygii & Crossopterygii & Dipnoi (Osteichthyes, swim bladder, skeleton, most diverse vertebrate taxon)

cartilaginous fishes don’t have a swim bladder for reduction of specific weight (pelagic zone), enrichment of fat and oils in liver to ensure buoyancy, constant swimming with large pectoral fins as lift producing wings

bony fishes: large number of eggs

sharks/rays: vivipary (“placenta”), ovovivipary (eggs with embryo develop within female) & ovipary (eggs)

Consequences: (ovo)viviparous fishes have late maturity age and only few offspring -> vulnerable populations

Independent evolution of similar trait. E.g. streamline body shape

Predator confusion, detection dilution (less spots where predators can find prey  levy-flight of predators)

1500x larger (25x larger area). Adaptations: bioluminescence (prey attraction/location), photophores on ventral side (invisibility against water surface), vertical swimming (decreased surface area from below), extendable stomach and jaws (food shortage), dwarf males (mating partners scarce -> stay attached)

gigantism in deep sea, chromatophores for colour changing (signalling/camouflage), arms & tentacles for prey/detritus catchment; generation time only 1yr -> one reproductive investment, then death (semelparous) -> mass spawning and death events -> huge amounts of biomass sinking fast to the seafloor during patchy events

Historically: direct exploitation because they were easy catch on long sea journeys; Now: bycatch

Penguins, Albatrosses, Pelicans, Herons/storks, Shore/wading birds/skuas/auks -> inaccessible breeding sites

streamer (tori) lines -> scare birds away, line weighting -> more rapid sinking of lines

decrease of S:V ratio by exceeding 1m in length + 50kg mass; insulation via body fat; high metabolic rate

Baleen whales: positive allometry in filter-feeding adaptations, energy efficiency increases with weight

Toothed whales: investment in biosonar (high costs), energy efficiency decreases with whales, sperm & beaked whales are large to push physiological limits for dive duration and depth

Whale falls are stepping stones for gene flow, demographic exchange and spatial distribution of sulphophilic organisms that normally live at hydrothermal vents

• symbiosis is ubiquitous across the tree of life -> presence of microbes

• symbionts can be transmitted between generations

  -> vertical (microbes in egg) + horizontal

• microbial symbionts affect holobiont fitness

• variation in the symbiont community aids in holobiont acclimation to environmental changes

-> corals: microbes in “skin”, allow corals to adapt to changing environmental conditions more rapidly than via mutation and selection

largest tidal flat system in the world, sediments supplied from sea, high productivity, many endemic species, important for migratory birds, fish nursery

lower: predation, competition         upper: physical factors (dessication, wave shock)

-> removal/ addition/ replacement series, predator removal, direct observation of overgrowth competition

Ecological: more competition by opportunists, bioinvasions, increased grazing rates

Physical: de-/hypersalination, storms + increased wave action, warming

-> poleward migration of intertidal species

fringing reef: grows seaward directly from shore

barrier reef: grow seaward separated from land (lagoon)

atoll: in open ocean, open lagoon in center, patch reef

-> symbiosis with unicellular algae for photosynthesis, ability to fix calcium carbonate (build skeleton)

salt stress: salt excretion via leaves, salt exclusion to prevent it from entering the vascular system

anoxic sediments: pneumatophores, prop/ kneed/ plank roots to gain alternative access to air

habitat in the mesophotic zone (30-150m), inhabited by light dependent organisms BUT extreme light-limited conditions (adaptations), continuation of neighbouring shallow habitats (same species, different appearance)

algae, corals, sponges

mesophotic populations can provide viable reproductive material for shallow reef areas following disturbances -> only tested on limited taxonomic, geographic, bathymetric, climatic basis

potential of connectivity varies by location/taxa; enhanced/inhibited by larval supply, behavioural movement, phenotypic plasticity, local hydrodynamic patterns

Important because they could serve es reservoir to replenish biodiversity lost in shallow waters, but they also have their own unique biological communities that should be protected

Photic zone in pelagic ocean, biodiversity reservoir, feeding grounds for endangered mammals, carbon sink

Bioluminescence -> prey detection/attraction, counterillumination, signalling -> Lantern-/ anglerfishes, cephalopods

Abyssal plain; food: terrestrial vegetation, migrating deep-sea fish, large food falls; interest: Mn nodules

Position: increased S:V ratio, flat shape, constant swimming, increased buoyancy

Sense: well developed eyes, lateral line, echolocation, colouration, transparency, silhouette reduction

Feeding: diel vertical migration, predation, cannibalism, extendable stomach, catch sinking detritus

Mimicry: sargassum seaweed inhabited by a number of organisms that mimic its appearance

low solar irradiance at high latitudes, negative net radiation budget, extreme seasonality in radiation budget (light availability), high albedo due to ice and snow, both poles are glaciated, short summers

Transpolar drift: current from new Siberian islands across North pole towards Greenland

Beaufort gyre: circulation towards Bering strait; little water exchange with Pacific because Bering strait is so narrow -> Inflow of warm water via Eastern Fram strait, outflow of cold water via Western Fram strait, increasing inflow of Atlantic water masses

stable surface water layer due to summer warming & freshwater inflow -> nutrient limitation; vertical thermohaline convection in Greenland basin in winter -> oxygen transport to deep Arctic ocean

Winds: east wind drift over pole, west wind drift around pole

Currents: Antarctic divergence below east wind drift, Antarctic convergence (polar front) below west wind drift (ACC)

isolated surface currents, weak vertical stability (downwelling at margin, constant upwelling at divergence)

the one with the lowest nutrient requirement

F

T

T

T

at eastern boundaries and equator; high primary productivity and food web activity, high fish production, high carbon fluxes to midwater depths, especially old water masses at midwater depths, high oxygen demand & low supply (OMZ)

Long-term adaptations: increasing gill/lung size, reduction of diffusion distance, enhanced oxygen uptake capacity of respiratory proteins, counter-flow principle, small body size, low metabolic activity, enhanced capacity for anaerobic metabolism

Short-term: increasing respiratory protein concentrations, faster breathing, low activity, metabolic suppression, moving away, only temporarily use the low oxygen habitat