Lecture 17: Marine Microbes and their Significance in Biogeochemical Cycles Flashcards
What is Sulfur?
- Atomic no. 16
- ~1% of dry weight of organisms
- Two mineral forms:
a. Sulfide (S^2-)
b. Sulfate (S04^2-) - Used as:
a. fertiliser (plants = high demand)
b. Manufacturing insecticides, fungicides,
etc.
Why is sulfur important?
- Metabolism - fuel and respiratory
electron donor - Proteins - AAs; cysteine and methionine
- Disulphide bonds - protein stability +
structure - Cofactors in enzymes - e.g., FeS in
Aconitase (TCA cycle) - Antioxidant molecules glutathione and
thioredoxin
What are the different sulfur processes in microbes?
1. Assimilation - using sulfate to form organic compounds (E dependent) (animals use preformed S-compounds) 2. Dissimilatory sulfur metabolism - bacteria and archaea use sulfur in E yielding reactions (Ox or An) 3. Electron acceptors or donors (e.g., Sulfate reduction & oxidation) 4. Generating signalling molecules 5. Generating anti-stress molecules
What does assimilate mean?
Get something into a form that is useable
Explain assimilatory uptake of sulfate
- Sulfate abundant in marine environments
(~10^6 x more abundant than other S) - Some incorporated into polysaccharides
- Sulfate reduced to Sulfide in many
microbes (requires E) - Sulfide used to make AAs cysteine and
methionine
What are different types of organisms transforming sulfur
- Phytoplankton
- Seaweeds
- Few species from angiosperms
- Some corals
- Many heterotrophic bacteria
What is marine DMSP synthesis?
1. Reduce sulfate to Sulfide to make methionine (all req. E) 2. L-methionine is then turned into "Dimethylsulfoniopropionate" (DMSP) a. stable and soluble zwitterion ( can be + & - charged b. Prod. at v high [mM] e.g., 1/2M c. > 8 billion tonnes made per year 3. E.g., a. Some marine eukaryotes b. Marine bacteria (Alphaproteobac teria)
Why is DMSP produced?
- Suggested roles:
a. Osmoprotectant - OSMOLITES
(balancing the osmotic difference
between cell’s surroundings and
cytosol)
b. Cryoprotectant - CRYOLITES
(maintaining enzyme activities in
high and low temperatures)
c. Grazing deterrent - deters predators
due to their bad taste/toxicity
d. Oxidative stress protectant -
ANTIOXIDANTs DMSP and its
catabolites scavenge oxygen free
radicals generated by oxidative
stress
e. Storage molecule for carbon and
sulfur - lots of carbon and sulfur in
its structure so can be broken down
at a later date
Why only suggested roles?
none have been definitively established
Why is it that we do not know the single gene or enzyme involved in these DMSP pathways, yet we know the biochemical synthesis pathways?
1. Marine biology generally lacking in molecular biological approaches compared to terrestrial systems 2. Marine eukaryotes that make DMSP lack genetic manipulation tools
What is known about DMSP pathways?
- > 8 billion tonnes is made annually in
surface oceans - 3-10% of global marine primary carbon
production (GPP) - DMSP catabolism supplies 3-10% of
carbon requirements of heterotrophic
bacteria - DMSP supplies 50-100% of sulfur
demands for heterotrophic bacteria
(despite there being 1-10 million x more
sulfate) VIA CATABOLISM
What is DMSP catabolism
1. organisms eat DMSP and gain benefits such as E, carbon, sulfur 2. DMSP - major nutrient for heterotrophic bacteria (N & S) (& E) a. 3-10% of carbon supply b. 30-100% sulfur supply
What are the two pathways for DMSP catabolism
- Sulfur - demethylation pathway
a. dmd genes
b. 70% of DMSP in marine environment
eaten forms MeSH (methanethiol)
c. MeSH used as a source of sulfate
What is SAR11 bacteria?
- most abundant and ubiquitous clade of
heterotrophic marine bacteria (1/3 of
cells in photic zone) - Examples
a. Candidatus Pelagibacter ubique - 1st
cultured
b. P. ubique genomes small (1.3Mbp)
c. smallest no. genes in a living organism
d. pathways for all 20 AAs and most
cofactors - these bacteria have lost the genes allowing assimilation of sulfate into Sulfide
a. therefore need reduced sulfur sources
What is DMSP lysis?
- ddd genes
- approx. 30% of DMSP eaten generates
Dimethylsulfide (DMS)
DMSP -> DMS + Acrylate or 3-hydroxypropionate
What can SAR11 do?
DMSP to produce MeSH (using dmd genes) for sulfur, or produce DMS (using dddK genes)for carbon
What are other examples of bacteria heterotrophic bacteria that lyse DMSP?
- (alpha) Roseobacter
2. (gamma) Halomonas
What are Roseobacters?
- marine alpha-proteobacteria
- 10-25% bacteria in marine surface
waters, sediments and sea ice
(abundant) - Easily cultured (ADVANTAGE OVER
SAR11 FOR STUDIES) - Many genomes available
- Just as abundant SAR11 but more diverse
(more genera)
What is the problem with SAR11?
- VERY DIFFICULT TO CULTURE
a. cant grow on plates
b. cant isolate single colonies
c. can only grow in liquid culture
d. difficult to monitor growth and its rate
Why is roseobacter a good model organism?
- First studied: Ruegeria pomeroyi DSS-3
- Isolated from DMSP enrichments of coastal Georgia Seawater
- Genome sequenced in 2004
- Genetically tractable
- Much like SAR11:
a. can produce DMS (using dddP, dddQ,
dddW genes) lyse
b. MeSH (dmd genes) demethylate
What happens to DMS?
- 90% re-catabolised by marine bacteria as
carbon source (Methylophaga spp.) - Other 10% transferred from marine
system to atmosphere (~35 million
tonnes)
a. major biogenetic source of sulfur in
atmosphere
What happens to DMS in the atmosphere?
What is CNN?
- DMS oxidation produces products
- these products act as “Cloud
Condensation Nuclei” (CNN) - Climate cooling effect by reflecting
sunlight - AS WELL AS THIS, clouds then move, from marine to terrestrial systems, COMPLETEING GLOBAL SULFUR SYSTEMS
CNN is the ‘nucleus’ for which water droplets form around, leading to a cloud
Explain the CLAW hypothesis
1. Proposition that DMSP dependent DMS production as a negative mechanism by which phytoplankton regulate their environment in CLAW a. Phytoplankton prod. DMSP... b. If it gets too cold they make more DMSP = more DMS c. = more DMS in atmosphere d. increases cloud cover e. BACTERIA ESSENTIALLY REGULATING THE ENVIRONEMT FOR THEMSELF