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Introduction to Oceanography Introduction to Oceanography Lecture 14: , Biological Productivity Memorial Day holiday Monday no lab meetings Go to any other lab section this week (and let the TA know!)

Bay of Fundy -- low , Photo by Dylan Kereluk, . Creative Commons A 2.0 Generic, Mudskipper (Periophthalmus modestus) at low tide, photo by OpenCage, Wikimedia Commons, Creative http://commons.wikimedia.org/wiki/File:Bay_of_Fundy_-_Tide_Out.jpg Commons A S-A 2.5, http://commons.wikimedia.org/wiki/File:Periophthalmus_modestus.jpg

Tides Earth-Moon-Sun System Planet-length waves Cyclic, repeating rise & fall of level • Earth-Sun Distance – Most regular phenomenon in the 150,000,000 km Daily tidal variation has great effects on life in & around the (Lab 8) • Earth-Moon Distance 385,000 km Caused by gravity and Much closer to Earth, but between Earth, Moon & much less massive Sun, their orbits around • Earth Obliquity = 23.5 each other, and the degrees Earth’s daily spin – Seasons

Photos by Samuel Wantman, Creative Commons A S-A 3.0, http://en.wikipedia.org/ wiki/File:Bay_of_Fundy_Low_Tide.jpg and Figure by Homonculus 2/Geologician, Wikimedia Commons, http://en.wikipedia.org/wiki/ Creative Commons A 3.0, http://en.wikipedia.org/wiki/ File:Bay_of_Fundy_High_Tide.jpg File:Lunar_perturbation.jpg

Tides are caused by the gravity of the Moon and Sun acting on Scaled image of Earth-Moon distance, Nickshanks, Wikimedia Commons, Creative Commons A 2.5 Earth and its ocean.

Pluto-Charon mutual orbit, Zhatt, Wikimedia Commons, Public Domain, To Sun http:// en.wikipedia.org/ wiki/File:Orbit2.gif Not to scale!

Newton’s cannon, Wikimedia Commons, Creative Commons A S-A 3.0, http://en.wikipedia.org/wiki/File:Newton_Cannon.svg

• Basic Orbital Mechanics • Earth-Moon Distance Brews Ohare, Wikimedia, Public Domain, http://en.wikipedia.org/wiki/File:Earth- • Planetary objects stay in orbit due to balance of Gravity – 384,000 km Moon.PNG and Centrifugal (at their center of mass) • Revolution period of the Moon • Like a spun on the end of string – 27.3 Days • Rotation period of the Moon also 27.3 Days • Synchronous Rotation: We always see the same side of the Moon

1 Phases of the Moon Phases of the Moon • New Moon • Waxing Crescent • 1/2 Moon: First

quarter 1st Qtr WAXING CRESCENT • Full Moon First • Etc. Quarter New – 7 days/quarter Full Full Moon

Third Quarter WANING CRESCENT rd Tom Ruen, Wikimedia 3 Qtr Commons, Public Domain, http://en.wikipedia.org/wiki/ Figure from NASA Starchild, Public Domain), File:Moon_phase_calendar_ http://starchild.gsfc.nasa.gov/Images/StarChild/icons/ May2005.png moon_above.gif

The Big Picture 3: Bulges The Moon and Sun both influence tides

Figures from U. Tennessee, http://csep10.phys.utk.edu/astr161/lect/time/tides.html

• Moon’s gravitational acting on the Earth tugs out a tidal bulge towards moon • Centrifugal force pushes a bulge away from moon on the far side of the Earth – TIDES TRY TO TRACK THE MOON

NOAA, Public Domain, http://oceanservice.noaa.gov/education/kits/tides/media/tide06a_450.gif The Sun’s gravity has a similar, Constructive interference: Sun and Moon tidal bulges oriented the but smaller effect (1/2 as strong). same way, resulting in strong tides – Spring Tide

Destructive interference: Sun and Moon tidal bulges partially Andrew Buck, Wikimedia Commons, Creative Commons A S-A 3.0, http:// commons.wikimedia.org/wiki/File:Tidal_braking.svg cancel each other, resulting in weak tides – Neap Tide

Effect of Sun & Moon Together Why is the moon’s effect on the tide • Spring Tides & Neap Tides greater than the sun’s? • Gravity balances Centrifugal at Earth’s center of mass • Elsewhere they don’t cancel – TIDE GENERATING FORCE:

GMMoon Ftides ∝ 3 REarth− Moon

• Tide generating force falls off faster with radius than gravity!

Adapted from figure by Nicky McLean, Wikimedia Commons, Public Domain, http://en.wikipedia.org/wiki/ € File:Tide.Bridgeport.30d.png

2 Equilibrium Theory of the Tides Tides in narrow, tapering bays • In narrow bays attached to the ocean, tides can slosh straight in and out Sun is much more massive! • Large tides can occur when the tidal frequency matches natural 7 » Msun ~ 3x10 Mmoon (resonant) oscillations BUT Sun is much further away! of the bay

» Rsun ~ 400 Rmoon

GMs F R3 M R3 1 S ≈ SE = s mE = 3×107 × = 0.47 GM 3 3 FL m Mm RSE 400 R3 Image from NOAA Online mE School for Weather, Public Domain, http:// www.srh.noaa.gov/ jetstream/ocean/ Solar tide 1/2 as big as lunar tide fundy_max.htm €

Bay of Fundy tides QUESTIONS?

Mont Saint-Michel and Tombelaine (tidal islands), France, Uwe Küchler, Wikimedia Commons, CC A S-A 3.0, http://commons.wikimedia.org/wiki/File:Mont_st_michel_aerial.jpg

Marine Life & Biological Productivity CLASSIFICATION SCHEMES

FOR MARINE ORGANISMS

1. Taxonomy: Based on genealogical relationships between organisms (ie, felines) 2. Mode of Nutrition 3. Habitat 4. Mobility

Estimated marine chlorophyll & terrestrial vegetation coverage map 1997-1998, SeaWiFS/NASA, Public Domain, http://en.wikipedia.org/wiki/File:Seawifs_global_biosphere.jpg

3 Genetic classification: Three Domains of Life Bacteria Cyanobacterial colonies, left: NASA, Public Domain, http://microbes.arc.nasa.gov/images/content/gallery/lightms/ 1. Bacteria: Simple single celled organisms, lack publication/lyngbya.jpg; right: Hamelin Pool -- Shark’s Bay, , photo by Happy Little Nomad, Wikimedia commons, CC A S-A 2.0, http://en.wikipedia.org/wiki/File:Stromatolites_in_Shark_Bay.jpg nucleus (E. coli) 2. Archaea: Outwardly similar to Bacteria; many live in extreme environments (hot springs, nuclear reactors, saline lakes, etc.) 3. Eucarya: Have a membrane-enclosed nucleus and other organelles; include protists, animals, fungi, plants

Whole genome tree of life, diagram by User_A1, based on Ciccarelli (2006) and Letunic (2007), Public Domain. http:// en.wikipedia.org/wiki/ File:CollapsedtreeLabels- simplified.svg

Archaea Eukaryota

Ostreococcus, a picoplankton (<1x10–6 m across!), Wenche Eikrem and Jahn Throndsen, University of Copepod, NOAA, Public OsloWikimedia Commons, CC A S-A Domain, http:// 2.5, http://en.wikipedia.org/wiki/ www.glerl.noaa.gov/pubs/ File:Ostreococcus_RCC143.jpg photogallery/Waterlife/ pages/0737.html

Thermococcus Gammatolerans, an Archaebacterium, Halobacteria (actually Archaea) and Eukarya (Dunaliella Angels Tapias, Wikimedia Commons, Creative salina), San Francisco Bay CA, dro!d, Wikimedia Commons A 3.0 Unported, http:// Commons, Creative Commons A S-A 2.0 http:// commons.wikimedia.org/wiki/ commons.wikimedia.org/wiki/ File:Thermococcus_gammatolerans.jpg File:Salt_ponds_SF_Bay_%28dro!d%29.jpg Public Domain

Questions? What does it eat? 1. Autotrophs: Make their own food; Cyanobacteria, NASA, Public are the base of the Domain, http:// microbes.arc.nasa.gov/images/ content/gallery/lightms/ a. Autotrophs are Primary Producers publication/lyngbya.jpg b. Photosynthesizing plants, algae, some bacteria (store solar energy) c. Chemosynthetic bacteria

Sargassum natans (eukarya), James St. John, Creative Commons Attribution 2.0 Generic, https:// commons.wikimedia.org/wiki/ Comb jelly(?) (Eukaryota), Nick Hobgood, Wikimedia Commons, Creative Commons A S-A 3.0, File:Sargassum_natans_(brown_algae)_(San_Salvador_Is http://commons.wikimedia.org/wiki/File:Combjelly.jpg land,_Bahamas)_1_(15867880028).jpg

4 What does it eat? Photosynthesis Living systems require chemical energy 1. Heterotrophs: cannot make their own food; must Chlorophyll: a green pigment that captures photons and transfers their energy to eat other organisms or their remains electrons, an through a series of steps creates carbohydrate molecules (chemical energy) and . a. Herbivores: eat plants Chlorophyll looks green because it absorbs red and blue light, and reflects green light b. Carnivores: eat animals Sargassum algae, NOAA, Public Domain, http://oceanexplorer.noaa.gov/ Adapted c. Omnivores: eat plants & explorations/02sab/logs/aug09/media/ from figure lines.html by Aushulz, animals Wikimedia Commons, Creative d. Bacteria: many decompose Commons A dead organic matter (E. coli) Barracuda are heterotrophs, NOAA, Public S-A 3.0, Domain, http://www.photolib.noaa.gov/htmls/ http:// reef2567.htm commons.wi kimedia.org/ wiki/ File:Chlorop hyll_ab_spe So are yeast, ctra2.PNG Masur, Wikimedia Commons, Creative Commons A S-A 2.5, http://en.wikipedia.org/wiki/ File:S_cerevisiae_under_DIC_microscopy.jpg

Photosynthesis Reaction sunlight PRIMARY PRODUCTION 6CO + 6H O ——> C H O + 6O 2 2 6 12 6 2 • Amount of inorganic carbon (mainly Carbon CO2) “fixed” by autotrophic organisms + (yields) Glucose + Oxygen into organic compounds dioxide (a sugar) – Based on reactions harnessing solar or chemical energy

Typically, ~100 grams carbon/ year / meter2 is fixed to sugar in the open ocean

Respiration • : opposite reaction of photosynthesis Questions • Dis-assembly of carbohydrate (food) molecules in the presence of oxygen to release chemical energy

• The main byproducts of respiration are H2O and CO2. These are released to the environment • Both plants & animals use respiration • Some bacteria & archea also respire + ENERGY

C6H12O6 + 6O2 ——> 6CO2 + 6H2O Glucose + Carbon (a sugar) Oxygen (yields) + water dioxide Sargassum, photo from South Atlantic Fishery Management Council, http://www.safmc.net/Portals/6/weedline%202.jpg

5 How can we measure productivity? How can we measure productivity? Color-imetry! (yes, it means what you think) -Timed weighing of autotrophs - Chorophyll enables photosynthesis by absorbing blue and red Good for big land plants, possible for large seaweed. light. Green light is reflected or scattered. - Green ocean implies lots of chlorophyll However, measurements are only local. - Lots of chlorophyll implies lots of productivity! Hard to weigh microorganisms, particularly when they - Satellites like SEASTAR can measure color from space. This have a very short lifecycle. makes colorimetry ideal for global ocean surveys, if it works. This technique will be bad for long-lived plants (chlorophyll is -Timed “weighing” of inorganic carbon present even when big plants aren’t active, I.e. spruce trees) HYPOTHESIS: Green color in the ocean correlates 14 Add labeled inorganic carbon ( CO2) to ocean water with primary productivity. See how fast organisms convert it to organic molecules PREDICTION: Productivity estimated from color Works well for microorganisms, but still a local should be the same as productivity measured by measurement. “weighing” uptake of inorganic carbon.

Colorimetry compared with “weighing” Productivity from SeaWiFS M.J. Behrenfeld & P.G. Falkowski. 1997. Photosynthetic rates derived from satellite-based chlorophyll . Limnol. Oceanogr. 42:1-20

There is good agreement, but also a lot of scatter. Colorimetry may be a reasonable model, but we still Colorimetry

From Satellite Satellite From don’t know all the details!

14 From Measured uptake of inorganic C SeaWiFS/NASA/Rutgers University, Public Domain http://marine.rutgers.edu./opp/swf/Production/gif_files/PP_9809_9908.gif

Primary Production 1. Amount depends on: a. Driving Energy (Solar or chemical) b. Nutrients Figure from University of Michigan, http://www.globalchange.umich.edu/ 2. Regions of Highest Productivity globalchange1//lectures/ kling/energyflow/typeeco2.gif a. Continental margins: (Ekman pumping) and vertical mixing common along margins. Also close to rivers, dust sources b. Equatorial Divergences c. Antarctic Divergence d. Northern Pacific & Northern Atlantic i. Deepwater upwelling in Pacific; Divergence within subpolar Arctic/Atlantic gyres

6 Primary Production Primary Production 3. Regions of Lowest Productivity 4. Primary Producers (Autrotrophs) a. Interiors of subtropical gyres a. Eukaryotic Algae (Seaweeds & Single celled i. This is where ocean water is most stably photosynthesizers) stratified i. Benthic (coastal): minor component i. Seaweeds --- Strong, stable , little vertical ii. Pelagic phytoplankton: primary component mixing. Few nutrients are brought up to the i. Diatoms, dinoflagellates, , etc. surface. b. Cyanobacteria: blue-green algae --- These are the “deserts” of the ocean, most c. Picoplankton / Archea(?) nutrients lost as dead organisms sink into the deep ocean d. Chemosynthetic Bacteria: Use inorganic compounds to get energy

i. They oxidize compounds such as H2S (Hydrogen sulfide)

HABITAT Questions

Abalone, Oregon Aquarium, photo by Little Mountain 5, Wikimedia Commons, Creative Commons A S-A 3.0, http:// commons.wikimedia.org/wiki/ File:Abalone_OCA.jpg NOAA, Public Domain, http://oceanexplorer.noaa.gov/explorations/04deepscope/background/deeplight/media/diagram3_600.jpg

Habitat Ocean Habitats 1. Pelagic (oceanic): live in the water • Where do organisms live in the oceans? column – Biozones 2. Benthic: Live in or on ocean bottom • Benthic vs. Pelagic Whale shark, Georgia aquarium, Zac Wolf, Creative Commons A S-A 2.5, – Sea floor vs. Free-floating/free-swimming http://commons.wikimedia.org/wiki/File:Whale-shark-enhanced.jpg

– Light Zones • Photic, Dysphotic, Aphotic

Coral polyp, Nick Hobgood, Creative Commons A S-A 3.0, http:// commons.wikimedia.org/wiki/ File:Euphyllia_glabrescens_%28Hard_coral%29_with_polyps_exte nded.jpg

7 Ocean Habitats Relative Habitat Sizes

• Abyssal pelagic: 54% oceans by volume • Abyssal: 75% sea floor by area Figure by Chris_huh, Wikimedia Commons, Creative Commons A S-A 3.0, http://en.wikipedia.org/wiki/ File:Oceanic_divisions.svg • Yet most of the bioproduction occurs elsewhere, near the surface

•Neritic vs. Oceanic –Shelf vs. deep waters –Neritic/Sublittoral: reaches to sea floor

Questions Zonation by Lighting

• Photic Zone: lit by sunlight, ~ 100 - 500m deep – Euphotic Zone: autotrophs capture more energy than they use; net fixation of

carbon; net production of O2 – Dysphotic Zone: Not enough light for profitable photosynthesis

Spotted garden eel, photo by Nick Hobgood, • : Permanent darkness Wikimedia Commons, Creative Commons A S- A 3.0, http://commons.wikimedia.org/wiki/ File:Heteroconger_hassi_%28Spotted_garden_ eel%29.jpg

Coastal Ocean Open Ocean

NOAA, Public Domain, http://oceanexplorer.noaa.gov/explorations/04deepscope/background/deeplight/media/diagram3_600.jpg NOAA, Public Domain, http://oceanexplorer.noaa.gov/explorations/04deepscope/background/deeplight/media/diagram3_600.jpg

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