Apr 1, 2013: Properties of seawater that influence conditions experienced by biota High density fluid Charged molecule (with high density = viscous) Solubility: of ions; of gases – oxygen and carbon dioxide High light extinction coefficient High heat capacity Intertidally, desiccation and wave force Tides occur due to joint gravitational force of sun and moon. The moon’s orbit around the earth is 25 days, therefore patterns in the tidal cycle recur at this frequency. Tides progress each day by 24.8 hours. Spring tides occur every 2 weeks because they are driven by alignment of sun and moon on either the same or opposite sides of the earth. Neap tides occur when sun and moon do not align. slack Amplitude ebb flood Mean range includes all highs and lows Mean semidiurnal range includes all higher highs and lower lows 1 semidiurnal tidal cycle of 25 hr Chart datum =Mean lower low water calculated over many years Apr 3, 2013 – Habitats Intertidal (eulittoral) zones (Kozloff 1983) 1: supralittoral fringe affected only by higher tides; upper portion wet only from spray or rain (>7-9’ MLLW in Puget Sd) 2: Upper midlittoral zone (4-7’ MLLW in Puget Sd) = above mean tide level; rockweed on hard substratum 3: Lower midlittoral zone (0-4’ MLLW in Puget Sound) = below mean tide level; mussels on hard substratum 4: Infralittoral fringe below MLLW to the most extreme low tides; algae in particular “frustratingly rich for a beginner” Dethier habitat types: Marine = salinity 33 o Intertidal (rock, boulders, hardpan, cobble, mixed-coarse, gravel, sand, mixed-fine, mud, organic, artificial, reef) o Subtidal (bedrock and boulders, cobble, mixed coarse, gravel, mixed-fine, mud and mixed-fine, organic, artificial, reef) = below extreme lower low water Estuarine = salinities 0.5 to 30 o Intertidal (bedrock, hardpan, mixed-coarse, gravel, sand, mixed-fine, mixed-fine and mud, mud, organic, artificial, reef) o Subtidal (bedrock-boulder, cobble, mixed-coarse, sand, mixed-fines, mud, sand and mud, organic, artificial, reef) Environmental factors determining habitat type: Freshwater input: mid-estuary organisms do not necessarily need to tolerate moderate salinity, but rather variable salinity. Why is this challenging? Are there estuarine-adapted species or not? Water motion, wave action, morphodynamic state: Rapidly-moving water carries larger particles; in slow-moving water, larger particles drop out because gravity exceeds resuspension. p. 229 Boundary layers Free stream Logarithmic layer Laminar sublayer (or Not if turbulent) No slip conditions = water does not move Wetness/dryness, Emersion time: Paradigm is that upper limit of organisms is determined by physiological tolerance to heat and desiccation, and lower limit is determined by species interactions. April 8, 2013 – Primary production 1. Basic photosynthesis and respiration relationships 6CO2 + 6H20 + energy <-> C6H12O6 + 6O2 2. Primary producers in the ocean Bacteria Some flagellates Incl. coccolithophores Incl. foraminifera Dinoflag., Ciliates Diatoms, Brown algae Some amoebae Animals Fungi Rhodophyta Green algae+ land plants 3. Light in water (another habitat axis) a. Extinction coefficient = exponential rate at which light energy declines with depth, influenced by… organisms, DOM, bubbles, turbidity b. Light quality = red wavelengths are absorbed, whereas blue wavelengths penetrate (exception – CDOM from rivers reflects red light) 4. Photosynthesis dynamics – a. CHECK p. 65-66 for variable names; Michaelis-Menten equation for gross photosyntehsis: 𝑉 = 𝑉𝑚𝑎𝑥×[𝑆] [𝑆]+𝐾𝑚 1 𝐾𝑚 1 1 and as inverses 𝑉 = 𝑉𝑚𝑎𝑥 × 𝑆 + 𝑉𝑚𝑎𝑥. b. Plants respire, therefore negative net photosynthesis in low light or darkness c. PS rates plateau with high inputs (of light or CO2), likely as RUBISCO saturates 5. Definitions of productivity terms a. Instantaneous individual-level variables: i. Compensation point is light level at which net photosynthesis = 0 b. Integrated (day-night) individual-level variables: i. Compensation depth is the bottom of the (eu)photic zone, where carbon assimilation by phytoplankton equals respiratory losses (instantaneously) ii. Critical depth is where integrated daily photosynthetic carbon assimilation equals integrated daily respiratory carbon losses. (This depth increases from March to May in Box 2.5 as daylength increases C assimilation) iii. If mixing depth > critical depth, phytoplankton blooms are unlikely c. Whole-ecosystem variables: i. ii. iii. iv. v. Photosynthesis = P (gross production) Algal respiration = Ra Whole community respiration = Rc Net photosynthesis = NPP = P-Ra Net community production or Net ecosystem production = P-Rc (net autotrophic if PS dominates, and net heterotrophic if respiration dominates) What’s different about primary production in the ocean: diversity of organisms phylogenetically; light variability that generates niches. April 10, 2013 – energy flow and nutrient cycles 1. General principles: a. Inefficient energy transfer across trophic levels – i. Growth yield (efficiency) – growth per food intake ranges 10-30% ii. Trophic yield (efficiency) – production at trophic level relative to next lower trophic level ranges 5-20% iii. Inefficient energy transfer results in remineralized carbon and nutrients available to primary producers iv. Inefficient energy transfer limits biomass of large-bodied organisms b. Top-down control – biomass limited by consumption or disease (particularly viral mortality in marine microorganisms?). i. Trophic cascades = special case of indirect effects; top-down control extends for at least two sequential trophic interactions. c. Bottom-up control – i. of abundance or biomass (Liebig’s law of the minimum); ii. of population growth or metabolic rate (Blackman limitation) 2. Marine-specific concepts: a. Nitrogen cycling – f-ratio is the ratio of new to total (new and recycled) nitrogen taken up during primary production. i. Recycled nitrogen = ammonium, nitrite, urea, amino acids ii. New nitrogen = nitrate; can derive from atmosphere (precipitation, dry deposition), rivers (?), upwelling, or N-fixation. iii. F-ratio typically 0.3-0.5 but as low as 0.1 in deep or low-upwelling areas iv. Inverse of f-ratio tells how many times N moves through biotic compartment per year b. Carbon – distribution of respiration and CO2 by depth Epipelagic in light Mesopelagic 150-1000m bulk of respiration Bathypelagic 1000-4000m c. Microbial loop – Fig. 3.12 G PS C G G C R G C G R R G G DOM POM Abyssopelagic 4000-6000m Hadal 6000-10000m C C R R R All compartments have arrows recycling to DOM and POM d. Distribution of metabolism by body size is dominated by smallest organisms in ocean: i) combined biomass is highest in smallest-sized organisms; ii) specific metabolic activity – surface to volume ratio is highest, which determines rates of processes; iii) yield<100%