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Presentation Here 2/4/11 Microbes: drivers of global biogeochemistry proportal.mit.edu 1/28/2011 Prochlorococcus marinus forces.si.edu Soil fungus Virginia Rich ‐ EEB Dennis Kunkel Diatom www.bioquest.org forces.si.edu Soil bacteria Termite prost www.nhm.ac.uk www.morning-earth.org/Graphic-E/ Biosphere/Bios-Microbe-Image/M- PCophryoscolex Rumen prost Ophryoscolex Coccolithophore With material from Drs. Sco Saleska (U of Az), Gene Tyson (U of Queensland, Australia) and Kostas Konstannidis (Georgia Tech) Outline I. Big Picture II. How do microbes make a living (and thus interact with biogeochemical cycles)? III. Microbes & the C cycle – the highlights A. Primary Producon B. Decomposion Soil respiraon, ocean respiraon IV. Menon of other cycles, & Summary 1 2/4/11 I. Big picture: microbes drive biogeochemical cycles • ~ Half planetary primary marine forests & production (C fixation): microorganisms other big (cyanobacteria & green stuff phytoplankton) • Organic matter degradation: Without microbial • Metabolic diversity: recycling, nutrients Microbes perform all would be locked up & major metabolic become unavailable. pathways, and courses.worldcampus.psu.edu periodically reveal entirely new ones • Biomass: ~109 microbial cells/ (e.g. proteophodopsin, gram surface soil and ~106 cells/ml anaerobic methane oxidation). http://newscenter.lbl.gov/ seawater. (You have more microbial cells in your body than human cells). If all multi-cellular life disappeared 50-90% of marine biomass is microbial tomorrow the major biogoechemical (Census of Marine Life). cycles would likely proceed with very little change... II. How do microbes make a living? • “Microbes” can mean several things!!! For this overview defined as single‐celled organisms: bacteria and archaea (together oen called the “prokaryotes”, also “microbes”) plus single‐celled eukaryotes (aka “protozoans”, and the majority of “prosts”) How are microbes involved in all these biogeochemical cycles? What do microbes – indeed all cells – need to make a living? • CARBON for bulk of biomass • NUTRIENTS (N,P, S) and micronutrients for proteins, nucleic acids, etc. • WATER as a solvent (and a reactant in biomass producon) • ENERGY to allow them to work against entropy • ELECTRONS to transfer energy via redox reacons, and perform chemical transformaons – so a source and a sink for electrons ‐ 2 2/4/11 Let’s take an example we’re more familiar with: us. In our special case, our foods (complex organic compounds) provide us with Carbon, with Bond energy, AND with electrons. How are organic compounds used for all these things? Recall the mitrochondria? (Which are, as you’ll recall from high school bio, descendents of free‐living bacteria that took up residence inside eukaryoc cells long ago). Mitochondrial electron transport chain aka Krebs cycle Wikipedia Source for C, energy + electrons = carbon compounds Mitochondrial electron transport chain Carbon compounds (= “food”) www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia 3 2/4/11 Source for C, energy + electrons = carbon compounds Mitochondrial electron transport chain Carbon compounds (= “food”) biosynthesis biosynthesis biosynthesis www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia Mitochondrial electron transport chain Carbon compounds (= “food”) ATP biosynthesis biosynthesis biosynthesis www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia 4 2/4/11 Mitochondrial electron transport chain Carbon compounds (= “food”) ATP biosynthesis biosynthesis biosynthesis www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia Mitochondrial electron transport chain Carbon compounds (= “food”) ATP biosynthesis biosynthesis biosynthesis www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia 5 2/4/11 Mitochondrial electron transport chain Carbon compounds (= “food”) ATP biosynthesis biosynthesis biosynthesis www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia Mitochondrial electron transport chain Carbon compounds (= “food”) ATP biosynthesis biosynthesis biosynthesis www.power2fitness.com/images/krebs_cycle-lrg.gif Wikipedia Sink for electrons = O2 (producing water) 6 2/4/11 Words we use to describe where organisms get their carbon, energy, and electrons 1. Carbon • Autotroph Greek autos = self, trophe = nutrion. So what is their C source? How do they get it? What are some examples? wikipedia proportal.mit.edu earthobservatory.nasa.gov • Heterotroph heteros = other, trophe = nutrion. So what is their C source? How do they get it? Examples? Associated Press www.morning-earth.org/ Graphic-E/Biosphere/Bios- Microbe-Image/M- PCophryoscolex.jpg protistuser.uni- frankfurt.de.~schauder.ter mites.proto7_bg 2. Energy • Phototroph photo = light Energy comes from photons • Chemotroph chemo = chemical Energy comes from converng energy stored in chemical bonds (via their electrons) In both cases, captured energy is stored as ATP, carbs, lipids or proteins. 3. Electron Source • Organotroph organic = C‐containing. Use carbon compounds as electron donors. This includes us! • Lithotroph lithos = rock Use inorganic compounds as electron donors 4. Electron Sink • Aerobic respiraon uses O2 as terminal electron acceptor. When it’s available, it gets used because of highly favorable energecs. • Anaerobic respiraon occurs in absence of O2, using alternate ‐ terminal electron acceptor. E.g. denitrificaon uses nitrate (NO3 ), sulfate 2‐ reducon uses sulfate (SO4 ), methanogenesis uses carbon (CO2 or acetate) 7 2/4/11 Examples • How would we be classified under this trophic nomenclature? – Get C from others – Get electrons from C compounds – Get energy from bond energy Therefore we are Chemo organo heterotrophs, as are all mulcellular carnivores, herbivores, and many http://www.popartuk.com/g/l/lgsb0015+da-vincis-vitruvian- man-homer-simpson-the-simpsons-art-print.jpg many microbes. • How would land plants be classified? – Fix CO2 – Use sun for energy – What is their electron source? Is it organic or inorganic http://www.missouriplants.com/ Ferns/Equisetum_hymenale_stems Photo litho autotrophs not so important to memorize terms as to understand that a diversity of lifestyles exist, & thus a diversity of interactions with biogeochem. cycles Many biogeochemical transformations are unique to Bacteria and Archaea, and not found in Eukaryotes, e.g. Nitrogen fixation N2 ⇒ NH3 - - Nitrification NH3 ⇒NO2 ⇒NO3 Anaerobic respiration Use of electron acceptors other than O2 Examples Methanogenesis CO2 (or CH3COOH) ⇒ CH4 Denitrification NO - ⇒ N What jumps out just of this 3 2 brief sampling? The N cycle is dominated by 2- microbial transformations. Sulfate reduction SO4 ⇒ H2S 8 2/4/11 III. Microbes & the Carbon Cycle Sco (and others) says: “Carbon is the currency of life”: photosynthesis H2O + CO2 CH2O + O2 respiration Carbon in Reduced Carbon atmospheric CO2 in organic matter (biomass & energy supply) A. Primary Producon aka autotrophy Field et al. 1998 Science Ocean NPP Land NPP earthobservatory.nasa.gov ≈ • Ocean NPP ~ land NPP. Much sparser biomass, so why? • The oceans cover a lot of territory (~71% of the earth’s surface) • Not a lot of mul‐cellular primary producers in the oceans – sea grasses & most seaweeds limited to coasts, etc. – so it’s all about the microbes 9 2/4/11 Who is doing this marine phototrophy? EUKARYOTIC 40-70% of ocean primary PHYTOPLANKTON production proportal.mit.edu Dennis Kunkel CYANOBACTERIA www.nhm.ac.uk Coccolithophore (e.g. Prochlorococcus marinus) Diatom 30-60% of ocean primary production earthobservatory.nasa.gov http://www.photolib.noaa.gov/ bigs/fish1880.JPG But that’s just photoautotrophy… what about chemoautrophy? • In the oceans, chemoautrophy occurs in: – high organic‐maer coastal zone waters, – oxygen minimum zones (low O2‐horizontal stretches of the oceans, the expansion of which is linked to climate change in several ways. “Dead zones” are an extreme anthropogenic type of oxygen minimum zone), – hydrothermal vent systems, – marine sediments, – marine basalts, – the water column of the open ocean, – …. NSF press release for DeLong et al 2006 Science Basically everywhere in the sea, to varying degrees 10 2/4/11 Example: Chemolithoautotrophs living on and below the sea floor on basalt. “Laser confocal photomicrograph of a microbial biofilm aached to the surface of basalt chips from a depth of 1500 meters. Green is reflected light from the basalt surface and red is from Nile red–stained bacterial cells.” Stevens & McKinley, 1995 Science. Santelli et al., 2008 Nature Subseafloor microbes = 10–30% of the total living biomass of Earth - Whitman et al. 1998 PNAS. How dynamic might these pools be? ANY data on SIZE of chemoautotrophic C fixaon? Very few. One elegant example in a deep‐sea, low‐carbon, open‐ocean habitat: Used carbon isotopes in microbial biomass to track autotrophy (“you are what you eat”). Ingalls et al., 2006 PNAS “Total [autotrophic] biomass producon by archaea in deep waters is ... 1% of annual marine primary producon... of a magnitude significant to the global carbon cycle and greater than... that buried in marine sediments.” (It’s on scale of 1/10th annual human C addion to atmosphere. What kind of chemoautotrophy? NH3 nitrification, by ammonia- oxidizing archaea (ammonia is electron DONOR) Walker et al., 2010 PNAS 11 2/4/11 Summary of microbial autotrophy • Marine ~ terrerstrial NPP • Marine NPP ~ microbial • Marine NPP = cyanos + earthobservatory.nasa.gov eukaryoc phytoplankton • But wait, there’s a lot of non‐ phoc habitat on the planet • Chemoautrophs are NSF for DeLong et al 2006 abundant and diverse • Unclear how much C they
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