Bacterioplankton

From Wikipedia, the free encyclopedia

Jump to navigationJump to search

Image from an epifluorescence microscope of seawater stained with a green dye to reveal
bacterial cells and smaller viral particles

Part of a series on

Plankton

hide

Trophic groups

 Phytoplankton
 Zooplankton
 Mixoplankton
 Mycoplankton
 Bacterioplankton
 Virioplankton
 show

Size groups

show

Taxonomic groups

show

Other groups

show

Blooms

show

Other

 v
 t
 e

Bacterioplankton refers to the bacterial component of the plankton that drifts in

the water column. The name comes from the Ancient
Greek word πλανκτος (planktos), meaning "wanderer" or "drifter", and bacterium,
a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They
are found in both seawater and freshwater.
Bacterioplankton occupy a range of ecological niches in marine and aquatic
ecosystems. They are both primary producers and primary consumers in these
ecosystems and drive global biogeochemical cycling of elements essential for life
(e.g., carbon and nitrogen). Many bacterioplankton species are autotrophic, and
derive energy from either photosynthesis or chemosynthesis. Photosynthetic
bacterioplankton are often categorized as picophytoplankton, and include
major cyanobacterial groups such as Prochlorococcus and Synechococcus.
Other heterotrophic bacterioplankton are saprotrophic, and obtain energy by
consuming organic material produced by other organisms. This material may
be dissolved in the medium and taken directly from there, or bacteria may live
and grow in association with particulate material such as marine snow.
Bacterioplankton play critical roles in global nitrogen
fixation, nitrification, denitrification, remineralisation and methanogenesis.
Bacterioplankton abundance depends on environmental variables like
temperature, nutrient availability and predation. Like other small plankton, the
bacterioplankton are preyed upon by zooplankton (usually protozoans), and their
numbers are also controlled through infection by bacteriophages.

Contents
 1Major groups
o 1.1Photosynthetic Bacterioplankton
o 1.2Heterotrophic Bacterioplankton
 2Biogeochemical cycling
o 2.1Carbon
o 2.2Nitrogen
o 2.3Dissolved Organic Matter
 3Trophic interactions
 4Ecological significance
 5See also
 6References
 7External links

Major groups[edit]
Photosynthetic Bacterioplankton[edit]
Photosynthetic bacterioplankton are responsible for a large proportion of the
total primary production of aquatic food webs, supplying organic compounds to
higher trophic levels. These bacteria undergo oxygenic and anoxygenic
photosynthesis. Differences between these processes can be seen in the
byproducts produced, the primary electron donor, and the light harvesting
pigments used for energy capture.

Transmission electron micrograph showing the cyanobacteria Synechococcus elongatus

Cyanobacteria are a large group of photosynthetic bacterioplankton, often

growing as cells or in filamentous colonies.[1] These organisms are the dominant
group of bacterioplankton using oxygenic photosynthesis in aquatic ecosystems.
Cyanobacteria, along with photosynthetic eukaryotes, are responsible for
approximately half of the total global primary production [2] making them key
players in the food web. They use photosynthesis to generate energy in the form
of organic compounds and produce oxygen as a byproduct. [3] Major light
harvesting pigments
include chlorophylls, phycoerytherin, phycocyanin and carotenoids.[4] The majority
of cyanobacteria found in marine environments are represented by the
genera Synechococcus and Prochlorococcus. Synechococcus is cosmopolitan,
having been reported across temperate and tropical waters. [5] Prochlorococcus is
a very small in size and is found mainly in the euphotic zone of tropical waters. [6]
[7]
 Factors including light, nutrients, and temperature can cause cyanobacteria to
proliferate and form harmful blooms.[8] Cyanobacteria blooms can cause hypoxia
and produce high levels of toxins, impacting other aquatic organisms as well as
causing illnesses in humans.
Some Cyanobacteria are capable of nitrogen fixation. The genus Anabaena uses
specialized cells called heterocysts to physically separate nitrogen fixation and
photosynthesis.[9] Trichodesmium is an example of cyanobacteria that is capable
of fixing nitrogen through an alternative photosynthetic pathway. [10]
Other photosynthetic bacterioplankton, including purple and green bacteria,
undergo anoxygenic photosynthesis in anaerobic conditions. The pigments
synthesized in these organisms are sensitive to oxygen. In purple bacteria the
major pigments include bacteriochlorophyll a and b and carotenoids. Green
bacteria have different light harvesting pigments consisting of bacteriochlorophyll
c, d and e.[1] These organisms do not produce oxygen through photosynthesis or
use water as a reducing agent. Many of these organisms use sulfur, hydrogen or
other compounds as an energy source to drive photosynthesis. Most of these
bacterioplankton are found in anoxic waters, including stagnant and hypersaline
environments.[11]
Heterotrophic Bacterioplankton[edit]
Heterotrophic bacterioplankton rely on the available concentration of dissolved
organic matter in the water column. Usually these organisms are saprophytic,
absorbing nutrients from their surroundings. These heterotrophs also play a key
role in the microbial loop and the remineralization of organic compounds like
carbon and nitrogen. Pelagibacterales, also known as members of an
alphaproteobacteria clade, are the most abundant bacterioplankton in the
oceans. Members of this group are found in waters with low nutrient availability
and are preyed on by protists.[12][13]

Biogeochemical cycling[edit]
Carbon[edit]
Atmospheric carbon is sequestered into the ocean by three main pumps which
have been known for 30 years: the solubility pump, the carbonate pump, and
the biological carbon pump (BCP).[14] The biological carbon pump is a vertical
transmission pump driven mainly by the sinking of organic rich particles. Bacterial
phytoplankton near the surface incorporate atmospheric CO 2 and other nutrients
into their biomass during photosynthesis. At the time of their death these
phytoplankton, along with their incorporated carbon, sink to the bottom of the
ocean where the carbon remains for thousands of years. [15] The other biologically
mediated sequestration of carbon in the ocean occurs through the microbial
pump. The microbial pump is responsible for the production of old recalcitrant
dissolved organic carbon (DOC) which is >100 years old. [14] Plankton in the ocean
are incapable of breaking down this recalcitrant DOC and thus it remains in the
oceans for 1000s years without being respired. The two pumps work
simultaneously, and the balance between them is believed to vary based on the
availability of nutrients.[16] Overall, the oceans act as a sink for atmospheric
CO2 but also release some carbon back into the atmosphere. [17] This occurs when
bacterioplankton and other organisms in the ocean consume organic matter and
respire CO2, and as a result of the solubility equilibrium between the ocean and
the atmosphere.
Nitrogen[edit]

A schematic showing the cycling of nitrogen within the ocean.

The nitrogen cycle in the oceans is mediated by microorganisms, many of which

are bacteria, performing multiple conversions such as: nitrogen
fixation, denitrification, assimilation, and anaerobic ammonia oxidation
(anammox). There are many different nitrogen metabolism strategies employed
by bacterioplankton. Starting with molecular nitrogen in the atmosphere (N 2),
which is fixed by diazatrophs such as trichodesmium into usable forms like
ammonia (NH4).[18] This ammonia can then be assimilated into organic matter like
amino and nucleic acids, by both photoautrophic and heterotrophic plankton, it
can also be nitrified to NO3 for energy production by nitrifying bacteria. Finally the
use of NO3 or NO2 as terminal electron acceptors reduces the nitrogen back into
N2, which is then released back into the atmosphere thus closing the cycle.
[19]
 Another important process involved in the regeneration of atmospheric N 2 is
anammox.[19][20] Anammox, a process in which ammonia is combined with nitrite in
order to produce diatomic nitrogen and water, could account for 30-50% of
production of N2 in the ocean.[20]
Dissolved Organic Matter[edit]
Dissolved organic matter (DOM) is available in many forms in the ocean, and is
responsible for supporting the growth of bacteria and microorganisms in the
ocean. The two main sources of this dissolved organic matter are; decomposition
of higher trophic level organisms like plants and fish, and secondly DOM in
runoffs that pass through soil with high levels of organic material. It is important
to note that the age and quality of the DOM is important for its usability by
microbes.[21] The majority of the DOM in the oceans is refractory or semi-labile
and is not available for biodegradation.[22] As mentioned above the microbial pump
is responsible for the production of refractory DOM which is unavailable for
biodegradation and remains dissolved in the oceans for thousands of years.
[14]
 The turnover of labile DOM organic material is quite high due to scarcity, this is
important for the support of multiple trophic levels in the microbial community.
[23]
 The uptake and respiration of DOM by heterotrophs closes the cycle by
producing CO2.
Trophic interactions[edit]

Schematic of the food chain in the freshwaters of Lake Ontario.

Variations in bacterioplankton abundance are usually a result of

temperature, zooplankton grazing, and availability of substrate. [24] Bacterial
abundance and productivity are consistently related to algal abundance and
productivity as well as organic carbon. Additionally, phosphorus directly
influences both algal and bacterial abundance and in turn, algae and bacteria
directly influence each other's abundance[24] In extremely oligotrophic
environments, both bacterial and algal growth is limited by phosphorus, but
because bacteria are better competitors they obtain a larger portion of the
inorganic substrate and increase in abundance more rapidly than algae.
In marine pelagic environments, heterotrophic nano-flagellates are the most
probable consumers of bacterial cell production. [25] Cultured flagellates in
laboratory experiments demonstrate that they are adapted to predation on
bacteria-sized particles and occur in concentrations to control bacterial biomass.
[26]
 Tight fluctuations in numbers of bacteria and flagellates have been found in
a eutrophic estuary, particularly in the summer.[25][27] The amplitude of these
fluctuations increases in response to artificial eutrophication with inorganic
nutrients and decreases in response to predation. Losses of bacterioplankton by
grazing is indirectly related to carbon balances and directly related
to prokaryotic inhibitors.[28] A surplus of substrate would cause increased flagellate
biomass, increased grazing on bacterioplankton and therefore decreased
bacterial biomass overall. Predation of ciliates is analogous to predation by
flagellates on bacteria as well.
With using prokaryotic inhibitors seasonally, there is a positive relationship
between bacterial abundance and heterotrophic nanoplankton grazing rates and
only 40-45 % of bacterioplankton production was observed to be consumed by
phagotrophic Protozoa.[29] Additionally, eukaryotic inhibitory experiments show
that protozoan grazing has a positive effect on bacterioplankton production
suggesting that nitrogen regeneration by Protozoa could be highly important for
bacterial growth. Eukaryotic inhibitors did not prove to be useful to determine
protozoan grazing rates on bacterioplankton, however they may help understand
control mechanisms in the microbial food web. [29]
Ecological significance[edit]

A large harmful bloom of cyanobacteria, more commonly known as blue-green algae, spread across
the lake in green filaments and strands that are clearly visible in this simulated-natural-colour image.

Bacterioplankton such as cyanobacteria are able to have toxic blooms in

eutrophic lakes which can lead to the death of many organisms such as fish,
birds, cattle, pets and humans.[30] A few examples of these harmful blooms is
the Microcystis bloom in the year 2000 in Swan River estuary, Australia, [31] and
the Oostvaarderplassen in the Netherlands in 2003. [32] The detrimental effects of
these blooms can range from heart malformation in fish [33] to constraining
copepod reproduction.[34]
High temperatures caused by seasonality increases stratification and preventing
vertical turbulent mixing which increases competition for light that favours
buoyant cyanobacteria.[35][36] Higher temperatures also reduce the viscosity of
water which allows faster movement which also favors buoyant species of
cyanobacteria.[30] These species are also very competitive with the ability to create
a surface cover preventing light to reach deeper species of plankton. [35][37][36]
Climate studies are also indicating that with increasing hot waves the likelihood
of detrimental cyanobacterial blooms will become more of a threat to eutrophic
freshwater systems.[38][39][40] Other implications of the increasing average air
temperature due to climate change is that there might be an expansion of the
cyanobacterial bloom season, extending from earlier in the spring to later in the
fall.[41]
Estimates of bacterioplankton abundance and density can be derived with a
variety of methods including direct counts, flow cytometry, and conclusions
drawn from metabolic measures.
Further, as discussed in the biogeochemical cycling section, plankton are
responsible for the recycling and movement of essential nutrients (i.e.
nitrogen/carbon/DOM) which are essential building blocks for many of the
organisms co-existing with bacterioplankton in these ecosystems. These
recycled nutrients can be reused by primary producers, thus increasing the
efficiency of the biological food web and minimizing energy waste.

See also[edit]
 Cyanobacteria
 Pelagibacter
 Polynucleobacter
 Limnohabitans
 Phytoplankton
 Plankton
 Zooplankton
 Marine bacteria
 Marine bacteriophage

References[edit]
1. ^ Jump up to:a b Mann, Nicholas H; Carr, Noel G, eds. (1992).  Photosynthetic
Prokaryotes | SpringerLink.  doi:10.1007/978-1-4757-1332-9.  ISBN  978-1-4757-1334-
3. S2CID  6924271.
2. ^ Field, Christopher B.; Behrenfeld, Michael J.; Randerson, James T.; Falkowski,
Paul (1998-07-10). "Primary Production of the Biosphere: Integrating Terrestrial and Oceanic
Components".  Science.  281  (5374): 237–
240.  Bibcode:1998Sci...281..237F. doi:10.1126/science.281.5374.237. ISSN 0036-8075.  P
MID  9657713.  S2CID 45140824.
3. ^ Peschek, Günter A.; Bernroitner, Margit; Sari, Samira; Pairer, Martin; Obinger,
Christian (2011). Bioenergetic Processes of Cyanobacteria. Springer, Dordrecht. pp. 3–
70.  doi:10.1007/978-94-007-0388-9_1. ISBN 9789400703520.
4. ^ Colyer, Christa L.; Kinkade, Christopher S.; Viskari, Pertti J.; Landers, James P.
(2005-06-01). "Analysis of cyanobacterial pigments and proteins by electrophoretic and
chromatographic methods". Analytical and Bioanalytical Chemistry. 382 (3): 559–
569.  doi:10.1007/s00216-004-3020-4. ISSN 1618-2642.  PMID  15714301. S2CID  3538803
0.
5. ^ Johnson, Paul W.; Sieburth, John McN. (1979-09-01). "Chroococcoid
cyanobacteria in the sea: A ubiquitous and diverse phototrophic biomass1". Limnology and
Oceanography. 24(5): 928–
935.  Bibcode:1979LimOc..24..928J.  doi:10.4319/lo.1979.24.5.0928.  ISSN  1939-5590.
6. ^ Chisholm, Sallie W.; Frankel, Sheila L.; Goericke, Ralf; Olson, Robert J.; Palenik,
Brian; Waterbury, John B.; West-Johnsrud, Lisa; Zettler, Erik R. (1992-02-01).
"Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote
containing divinyl chlorophyll a and b". Archives of Microbiology.  157  (3): 297–
300.  doi:10.1007/bf00245165. ISSN 0302-8933.  S2CID 32682912.
7. ^ Chisholm, Sallie W.; Olson, Robert J.; Zettler, Erik R.; Goericke, Ralf; Waterbury,
John B.; Welschmeyer, Nicholas A. (July 1988). "A novel free-living prochlorophyte abundant
in the oceanic euphotic zone". Nature. 334 (6180): 340–
343.  Bibcode:1988Natur.334..340C. doi:10.1038/334340a0. ISSN 1476-4687.  S2CID 43731
02.
8. ^ Reynolds, C. S.; Walsby, A. E. (1975-11-01). "Water-Blooms". Biological
Reviews. 50 (4): 437–481. doi:10.1111/j.1469-185x.1975.tb01060.x.  ISSN  1469-185X.
9. ^ Agnihotri, Vijai K (2014). "Anabaena flos-aquae".  Critical Reviews in Environmental
Science and Technology.  44  (18): 1995–
2037.  doi:10.1080/10643389.2013.803797.  S2CID 84472933.
10. ^ Bergman, Birgitta; Sandh, Gustaf; Lin, Senjie; Larsson, John; Carpenter, Edward J.
(2013-05-01).  "Trichodesmium– a widespread marine cyanobacterium with unusual nitrogen
fixation properties".  FEMS Microbiology Reviews.  37  (3): 286–302.  doi:10.1111/j.1574-
6976.2012.00352.x.  ISSN  0168-6445. PMC  3655545.  PMID  22928644.
11. ^ Kopylov, Alexander I.; Kosolapov, Dmitriy B.; Degermendzhy, Nadezhda N.; Zotina,
Tatiana A.; Romanenko, Anna V. (2002-04-01). "Phytoplankton, bacterial production and
protozoan bacterivory in stratified, brackish-water Lake Shira (Khakasia, Siberia)".  Aquatic
 Ecology. 36 (2): 205–218. doi:10.1023/a:1015611023296.  ISSN  1386-
2588.  S2CID 20520535.
12. ^ Morris, Robert M.; Rappé, Michael S.; Connon, Stephanie A.; Vergin, Kevin L.;
Siebold, William A.; Carlson, Craig A.; Giovannoni, Stephen J. (December 2002). "SAR11
clade dominates ocean surface bacterioplankton communities". Nature. 420 (6917): 806–
810.  Bibcode:2002Natur.420..806M.  doi:10.1038/nature01240.  ISSN  1476-4687. PMID 124
90947. S2CID  4360530.
13. ^ Cole, JJ; Findlay, S; Pace, ML (1988).  "Bacterial production in fresh and saltwater
ecosystems: a cross-system overview".  Marine Ecology Progress Series.  43: 1–
10.  Bibcode:1988MEPS...43....1C. doi:10.3354/meps043001.
14. ^ Jump up to:a b c Legendre, Louis; Rivkin, Richard B.; Weinbauer, Markus G.; Guidi,
Lionel; Uitz, Julia (2015-05-01). "The microbial carbon pump concept: Potential
biogeochemical significance in the globally changing ocean". Progress in
Oceanography. 134: 432–
450.  Bibcode:2015PrOce.134..432L. doi:10.1016/j.pocean.2015.01.008.  ISSN  0079-6611.
15. ^ De La Rocha, C.L.; Passow, U. (2014). Treatise on Geochemistry. pp. 93–
122.  doi:10.1016/b978-0-08-095975-7.00604-5.  ISBN  9780080983004.
16. ^ Polimene, Luca; Sailley, Sevrine; Clark, Darren; Mitra, Aditee; Allen, J Icarus
(2017-03-01).  "Biological or microbial carbon pump? The role of phytoplankton stoichiometry
in ocean carbon sequestration".  Journal of Plankton
Research.  39  (2).  doi:10.1093/plankt/fbw091. ISSN 0142-7873.
17. ^ The ocean carbon cycle and climate. Follows, Mick., Oguz, Temel., North Atlantic
Treaty Organization. Scientific Affairs Division. Dordrecht: Kluwer Academic Publishers.
2004.  ISBN  9781402020872.  OCLC 54974524.