Phytoplankton. The Biological Pump. Nutrient Cycling and the Marine Biological Pump. Phytoplankton and Zooplankton. CSU ATS Sco9 Denning 1
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1 Nutrient Cycling and the Marine Biological Pump Readings: SelecGons from Williams & Follows (2011) Sabine et al (2004): Ocean Sink for Anthropogenic CO 2 Phytoplankton Diameter: < 1 mm to over 100 mm ConcentraGon: 1000's to 1,000,000 per milliliter If you filled a Coke can with seawater sampled from a thick phytoplankton bloom, the can would contain about 100 million cells! Global Biomass: < 1% of the plant biomass on earth BUT responsible for nearly half the net photosynthesis of the biosphere! What does this imply for turnover Gme? Phytoplankton and Zooplankton Size can t be used to distinguish between autotrophs and heterotrophs (note log scale!) Smallest organisms don t sink well Some photosynthesize when they can, eat DOC when they can t The Biological Pump Ducklow et al (2001) Primary producgon limited by availability of nutrients and light Loss of nutrients from light by sinking must equal delivery of nutrients by upwelling Primary producgon generates communiges of phytoplankton from DIC and nutrients in the presence of light Zooplankton graze on phytoplankton Bacterial decomposigon and heterotrophic respiragon recycle DIC and nutrients to the water column Detrital pargcles from dead phytoplankton and zooplankton waste coagulate into progressively larger pargcles Larger pargcles sink faster than turbulence can resuspend them, so fall below euphogc zone CSU ATS Sco9 Denning 1
2 Marine Ecosystem Model Predict PZND, plus Chlorophyll Assume a single limiting currency (e.g., N) in each pool No interactions among individuals, just pools More recent models (red) include limitation by multiple nutrients, N-fixation, and size classes of organisms Diatoms (Bacillariophyceae) SiO 2 tests Important for forming large particles that sink efficiently Carbonate and Coccoliths Alteration of carbonate equilibria (pco 2 ) near surface Export of CaCO 3 to deep ocean dissolution below lysocline Williams_Fig CSU ATS Sco9 Denning 2
3 Primary ProducGon and the efficiency of the biological pump New ProducGon: driven by NO 3 delivered by vergcal mogon (upwelling, turbulence) Recycled producgon: ~ 90% of NPP, driven by nutrients recycled from organic ma9er decomposed in euphogc zone Export producgon: delivery of POM to deep water at base of euphogc zone In steady-state, NP = EP f-rago is defined as NP/NPP e-rago defined as EP/NPP Provides a link between remote sensing (of NPP) and carbon flux to the deep ocean Nutrient LimitaGon Primary producgon (photosynthesis) requires: Light Macro nutrients: C, N, P Trace nutrients: Fe, other metals Leibig s Law of the Minimum nutrient available in the smallest quan1ty rela1ve to the requirement of a plant limits produc1vity Redfield RaGo 106 CO NO HPO H H + photosynthesis (hv) decomposition (CH 2 O) 106 (NH 3 ) 16 (H 3 PO 4 ) O 2 Bio-Carbon Cycle Near Ocean Surface steady-state perturbed by wind Redfield, Ketchum, and Richards (1963) RKR Carbon dioxide and nitrate are reduced only phosphate remains at the same oxidagon state P:N:C:O - 1:16:117:170 colimitagon by N & P not carbon limited! data at constant density surfaces CSU ATS Sco9 Denning 3
4 Biology & CirculaGon Deep Sea DIC DIC is strongly depleted near surface, enhanced at depth Progressive enhancement of DIC at depth with time as deep water flows through thermohaline circulation Strong mixing in Southern Ocean ( mixmaster ) Further enhancement as water moves North through Pacific Williams_Fig Preformed and Regenerated Nutrients Deep Sea PO 4 PO 4 is strongly depleted near surface, enhanced at depth Progressive enhancement of PO 4 at depth with time as deep water flows through thermohaline circulation Strong mixing in Southern Ocean ( mixmaster ) Further enhancement as water moves North through Pacific Williams_Fig CSU ATS Sco9 Denning 4
5 Components of Deep Sea DIC Components of Deep Sea DIC ATLANTIC PACIFIC Williams_Fig Williams_Fig Remote Sensing of Ocean Color Sea-viewing wide-field spectrometer (SeaWiFS) launched August 1, bands in visible and near-ir provide imagery of plant pigments 705 km circular, noon, sun-synchronous orbit ~ daily global coverage with < 5 km pixels CSU ATS Sco9 Denning 5
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8 Net Primary Production June, July, August of 1998 Net Primary Production December, January, February of 97 and 98 Note the stimulation of NPP from tropical upwelling in eastern Pacific Behrenfeld et al. [2001] Behrenfeld et al. [2001] Minimalist Marine Biology (OCMIP-1) Biogeochemical Fixation Diazotrophs Production/consumption just sufficient to restore model to observed sea-surface PO 4 3- (Redfield ratio) Implemented in many ocean GCMs Can reproduce many features of Chlorophyll distribution CSU ATS Sco9 Denning 8
9 Model can t capture Chl minima in HNLC regions without introducing Fe-limitation and prescribing Fedeposition Iron Fertilization Some areas have high N but low chlorophyll (HNLC) These regions are far from land (Eq Pac, circumpolar) Fe-limitation due to reduced atmospheric deposition of dust from continents Estimated Iron Deposition Dust model predicts lofting of terrestrial dust particles by the wind, transport, and deposition (composition) Desert dust is important, but grain size and mineral composition control transport and Fe content Doney et al (2001) Simulated Nutrient Limitations ML Ecosystem Model Evaluation Ecosystem model forced with observed T, S, ppt, etc Most of the marine biosphere is N-limited Large areas of iron limitation far (downwind) from arid continents Simulation Observations Compared to Bermuda-Atlantic timeseries station (BATS) data Seasonality, vertical profiles pretty well simulated CSU ATS Sco9 Denning 9
10 Simulated Preindustrial DIC Ecosystem model coupled to physical GCM Doney et al (2001) Advection of C, N, P, Fe, Si, Ca Chemical equilibria Agreement is pretty good! Simulated Global Biological Pump Doney et al (2001) Importance of coastal zones BioPump ~ 2/3 of total carbon delivery to deep ocean Solubility pump accounts for the remaining ~ 1/3 Is there a Gulf Stream? Sea-Surface Temperature Note baroclinic eddies and rings in Gulf Stream Slow currents, small Rossby radius Much of the meridional heat transport in the oceans is accomplished by these motions Effects on vertical motion and biology? Trajectories of derelict (drifting) ships (19 th century) CSU ATS Sco9 Denning 10
11 Mesoscale Physics -> Ecology Mesoscale Dynamics Vertical Motion, & the Biological Pump Growth of a baroclinic disturbance in an eddyresolving ocean model Top panels show maps, bottom panels show x- sections Big impact of vertical motions on NPP, shift from small to larger organisms, sinking POC Physics not resolved by most ocean models! Doney et al (2001) Atlantic Biogeochemicals carbon PO 4 Pacific pco 2 Analyses (after Takahashi) Advection/diffusion interpolation from point measurements Combines data from over 30 years of measurements, with corrections for d[pco 2 ]/dt and SST Note tropical upwelling, downwelling in gyres, highlatitude sinks SiO 4 (Feeley et al, 2001) CSU ATS Sco9 Denning 11
12 Spatial Patterns of Gas Exchange (Feeley et al, 2001) F = k ex S pco 2 Annual Flux Result Dependence of transfer velocity on wind speed convolved with wind and pco 2 patterns Little difference in subtropics (weak pco 2 ) More in tropics (lots of pco 2, weak winds) Most differences in higher latitudes (big pco 2, high winds) Anthropogenic CO 2 These are data from JGOFS stations Anthropogenic component estimated by correcting total DIC for anthropogenic component using stoichiometry Anthropogenic DIC is not accompanied by biological N and P Column Integrated Anthropogenic DIC Sabine et al (2004) Sabine et al (2004) CSU ATS Sco9 Denning 12
13 Distribution of Preindustrial DIC Surface water strongly depleted (~10%) due to biological removal Convection and deep water formation in North Atlantic Deep DIC: Pacific > Indian > Atlantic Anthropogenic DIC Estimated from total observed DIC using stoichiometry Most anthropogenic DIC confined to top few 100 m Shoaling in tropics, convection at higher latitudes Some contamination of bottom water in Atlantic (both hemispheres) (Feeley et al, 2001) CSU ATS Sco9 Denning 13
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