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Calcification - growth of the reef
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In ocean, mostly find 3 forms of CaC03 Calcite
Mostly of mineral origin Aragonite Fibrous, crystalline form, mostly from corals Magnesian calcite Smaller crystals, mostly plant origin
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Calcification Calcite Aragonite Magnesian calcite (Mg carbonate)
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Examples: organism CaCO3 Molluscs calcite & aragonite Corals
just aragonite Some green algae Red algae magnesian calcite Sponges aragonite (with silica) Some bryozoans all 3
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remove Ca++ & CO3-- from seawater Combines them to CaCO3
Corals remove Ca++ & CO3-- from seawater Combines them to CaCO3 transports them to base of polyp Calcicoblastic epidermis minute crystals secreted from base of polyp Energy expensive Energy from metabolism of algal PS products
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Calcification
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CO2 and seawater What forms of C are available to the coral ?
Organic and inorganic forms DIC - dissolved inorganic carbon CO2 (aq) HCO3- CO3--
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DIC comes from: Weathering dissolution of oceanic rock
Run-off from land Animal respiration Atmosphere etc.
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DIC in ocean constant over long periods
Can change suddenly on local scale E.g. environmental change, pollution Average seawater DIC = mmol/Kg Average seawater pH = pH affects nature of DIC
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Carbon and Seawater normal seawater - more HCO3- than CO3--
when atmospheric CO2 dissolves in water only 1% stays as CO2 rest dissociates to give HCO3- and CO3--
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H2O + CO2 (aq) H2CO3 HCO3- + H+ (1)
HCO CO H+ (2) equilibrium will depend heavily on [H+] = pH relative amounts of different ions will depend on pH
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dissolved carbonate removed by corals to make aragonite
Ca CO CaCO (3) pulls equilibrium (2) over, more HCO3- dissociates to CO3-- HCO CO H+ (2) removes HCO3-, pulls equilibrium in eq (1) to the right H2O + CO2 (aq) H2CO HCO H (1) more CO2 reacts with water to replace HCO3-, thus more CO2 has to dissolve in the seawater
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Can re-write this carbon relationship: 2 HCO3- CO2 + CO3-- + H2O
used to be thought that symbiotic zooxanthellae remove CO2 for PS pulls equation to right makes more CO3-- available for CaCO3 production by polyp No
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demonstrated by experiments with DCMU
stops PS electron transport, not CO2 uptake removed stimulatory effect of light on polyp CaCO3 deposition therefore, CO2 removal was not playing a role also, in deep water stony corals if more food provided, more CaCO3 was deposited more energy available for carbonate uptake & CaCO3 deposition
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Now clear that algae provide ATP (via CHO) to
allow polyp to secrete the CaCO3 and its organic fibrous matrix Calcification occurs 14 times faster in open than in shaded corals Cloudy days: calcification rate is 50% of rate on sunny days There is a background, non-algal-dependent rate
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Environmental Effects of Calcification
When atmospheric [CO2] increases, what happens to calcification rate ? goes down more CO2 should help calcification ? No
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H2O + CO2 (aq) H2CO3 HCO3- + H+ (1) HCO3- CO3-- + H+ (2)
Add CO2 to water quickly converted to carbonic acid dissociates to bicarbonate: H2O + CO2 (aq) H2CO HCO H+ (1) HCO CO H+ (2) Looks useful - OK if polyp in control, removing CO3--
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H2O + CO2 (aq) H2CO3 HCO3- + H+ (1) HCO3- CO3-- + H+ (2)
Add CO2 to water quickly converted to carbonic acid dissociates to bicarbonate: H2O + CO2 (aq) H2CO HCO H+ (1) HCO CO H+ (2) Looks useful - OK if polyp in control, removing CO3-- BUT, if CO2 increases, pushes eq (1) far to right
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H2O + CO2 (aq) H2CO3 HCO3- + H+ (1) HCO3- CO3-- + H+ (2)
Add CO2 to water quickly converted to carbonic acid dissociates to bicarbonate: H2O + CO2 (aq) H2CO HCO H+ (1) HCO CO H+ (2) Looks useful - OK if polyp in control, removing CO3-- BUT, if CO2 increases, pushes eq (1) far to right [H+] increases, carbonate converted to bicarbonate
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So, as more CO2 dissolves, more protons are released acidifies the water the carbonate combines with the protons produces bicarbonate decreases carbonate concentration
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Also, increase in [CO2] H2O + CO2 + CaCO3 2HCO3- + Ca++
leads to a less stable reef structure the dissolving of calcium carbonate H2O + CO2 + CaCO HCO Ca++
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addition of CO2 pushes equilibrium to right
Also, increase in [CO2] leads to a less stable reef structure the dissolving of calcium carbonate H2O + CO2 + CaCO HCO Ca++ addition of CO2 pushes equilibrium to right increases the dissolution of CaCO3
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Increases UV - DNA, PS pigments etc.
anything we do to increase atmospheric [CO2] leads to various deleterious effects on the reef: Increases solubility of CaCO3 Decreases [CO3--] decreasing calcification Increases temperature, leads to increased bleaching Increases UV - DNA, PS pigments etc.
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Reef Photosynthesis
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principally through photosynthesis
Productivity the production of organic compounds from inorganic atmospheric or aquatic carbon sources – mostly CO2 principally through photosynthesis chemosynthesis much less important. All life on earth is directly or indirectly dependant on primary production.
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gC/m2/d TropicalCoral Reef Tropical open ocean Mangrove 2.46 Tropical Rain Forest 5.5 Oak Forest 3.6
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no single major contributor to primary production on the reef
Productivity no single major contributor to primary production on the reef a mixture of photosynthetic organisms can be different at different locations
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net productivity values (varies with location):
gC/m2/d Calcareous reds 1 - 6 Halimeda 2 -3 Seagrass 1 - 7 N.S. kelp 5
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Overall productivity of the reef: 4.1 - 14.6 gC/m2/d from
epilithic algae, on rock, sand etc., few phytoplankton seagrasses Zooxanthellae (in coral etc.) Fleshy and calcareous macroalgae
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One obvious differences between different algae is their colour
Different colours due to the presence of different photosynthetic pigments
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Light Reflected Chloroplast Absorbed light Granum Transmitted
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Light and Photosynthesis
Air & water both absorb light a plant at sea level receives 20% less light than a plant on a mountain at 4,000m this reduction occurs faster in seawater depends a lot on location get 20% light reduction in 2m of tropical seawater get 20% light reduction in 20cm of Maritime seawater
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a very specific part of the EM spectrum
Photosynthetically Active Radiation nm
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Gamma rays X-rays UV Infrared Micro- waves Radio 10–5 nm 10–3 nm 1 nm
380 450 500 550 600 650 700 750 nm Visible light Shorter wavelength Higher energy Longer wavelength Lower energy
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Measure it as IRRADIANCE
moles of photons per unit area per unit time mol.m-2.s-1 E = Einstein = 1 mole of photons mE.m-2.s-1
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As light passes through seawater it gets ABSORBED & SCATTERED
= ATTENUATION (a reduction in irradiance) pure water attenuation lowest at 465nm increases towards UV and IR ends of spectrum TRANSMITTANCE is highest at 465nm not dealing with pure water Seawater has all kinds of dissolved salts, minerals, suspended material etc.:
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Attenuation is different in different locations - different light transmittance spectra:
To fully exploit a particular location, marine plants have a wide variety of PS pigments they can use.
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Mesophyll Chloroplast Outer membrane Thylakoid Intermembrane space
Inner Thylakoid Granum Stroma 1 µm
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An overview of photosynthesis
CO2 CALVIN CYCLE O2 [CH2O] (sugar) NADP ADP + P i H2O Light LIGHT REACTIONS ATP NADPH Chloroplast
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In the thylakoid membrane,
Light Reactions In the thylakoid membrane, chlorophyll molecules, other small molecules & proteins, are organized into photosystems photosystems composed of a reaction center surrounded by a number of light-harvesting complexes (LHC) LHC = pigment molecules bound to proteins
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LHC = pigment molecules bound to proteins
funnel energy of photons to the reaction center reaction-center chlorophyll absorbs energy One of its electrons gets bumped up to a primary electron acceptor electron transport ATP & NADPH production
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(INTERIOR OF THYLAKOID)
Photosystems Primary election acceptor Photon Thylakoid Light-harvesting complexes Reaction center Photosystem STROMA Thylakoid membrane Transfer of energy Special chlorophyll a molecules Pigment THYLAKOID SPACE (INTERIOR OF THYLAKOID) e–
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Light The visible light spectrum includes
the colors of light we can see the wavelengths that drive photosynthesis Photosymthetic pigments absorb light
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Gamma rays X-rays UV Infrared Micro- waves Radio 10–5 nm 10–3 nm 1 nm
380 450 500 550 600 650 700 750 nm Visible light Shorter wavelength Higher energy Longer wavelength Lower energy
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Light Reflected Chloroplast Absorbed light Granum Transmitted
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different pigments have different absorption spectra
combine in different amounts in different species to give each a unique absorption spectrum tells us which wavelengths of light are being absorbed (and thus it’s colour)
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Wavelength of light (nm)
Absorption spectra of pigments Absorption of light by chloroplast pigments Chlorophyll a Wavelength of light (nm) Chlorophyll b Carotenoids
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doesn’t tell us what the alga is doing with the light
For this you need to look at the ACTION SPECTRUM measures photosynthesis at different wavelengths
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The action spectrum of a pigment
show relative effectiveness of different wavelengths of radiation in driving photosynthesis Plots rate of photosynthesis versus wavelength
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Marine PS pigments 3 major groups of PS pigments in marine organisms
Chlorophylls Phycobiliproteins Carotenoids
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Chlorophyll a is essential
find it in all plants and algae the other pigments are accessory pigments in the antennae complexes funnel electrons to chlorophyll a in the reaction centres
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5 types of chlorophyll commonly found in marine organisms
all are tetrapyrrole rings with Mg++ in the middle chlorophyll a, b, c1, c2 & d a all green plants and algae b Chlorophyceae c1 & c2 Phaeophyceae d Rhodophyceae
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Chlorophyll a Chlorophyll b Is the main photosynthetic pigment
Is an accessory pigment C CH CH2 N H3C Mg H CH3 O CHO in chlorophyll a in chlorophyll b Porphyrin ring: Light-absorbing “head” of molecule note magnesium atom at center Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts: H atoms not shown Accessory pigments absorb different wavelengths of light and pass the energy to chlorophyll a
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Also a wide range of carotenoids
C40 TETRATERPENES very hydrophobic sit in membranes 2 types of carotenoids CAROTENES (hydrocarbons) XANTHOPHYLLS (have 1 or 2 oxygens)
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b-CAROTENE is the most common carotenoid in marine organisms
often see a mixture of b-CAROTENE & FUCOXANTHIN (another carotenoid) in the Phaeophyceae gives the brown colour
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PHYCOBILINS are linear tetrapyrroles attached to proteins
red pigments no ring, no chelation of a metal Only found in Rhodophyceae & Cyanophyceae and a few species of Cryptophyceae
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