Trichodesmium

plex microenvironments.

1 Species

Trichodesmium erythraeum described by Ehrenberg in

1830.[3] T. erythraeum is the species responsible for dis-
coloring the Red Sea during blooms. This is the only se-
quenced genome in the genus thus far and is the focus of
most laboratory studies (Trichodesmium IMS 101).
Trichodesmium thiebautii Described by Gomont in
1892.[4]
Trichodesmium hildebrantii Described by Gomont in
1892.[4]
Trichodesmium contortum Described by Wille in
1904.[5]
Trichodesmium tenue Described by Wille in 1904.[5]
Trichodesmium radians Described by Wille in 1904.[5]

Trichodesmium bloom o the Great Barrier Reef

2 Cell Structure
Trichodesmium, also called sea sawdust, is a genus of
lamentous cyanobacteria. They are found in nutrient
poor tropical and subtropical ocean waters (particularly Like most cyanobacteria, Trichodesmium has a gram
around Australia and in the Red Sea, where they were negative cell wall. However, unlike other aerobic
rst described by Captain Cook). Trichodesmium is a diazotrophs, heterocysts (structures found in cyanobac-
diazotroph; that is, it xes atmospheric nitrogen into teria which protect nitrogenase from oxygenation) are
ammonium, a nutrient used by other organisms. Tri- lacking in Trichodesmium. This is a unique character-
chodesmium is the only known diazotroph able to x ni- istic among aerobic diazotrophs which x nitrogen in
trogen in daylight under aerobic conditions without the daylight. Photosynthesis occurs using phycoerythrin -
use of heterocysts.[1] light-harvesting phycobiliprotein which is normally found
Trichodesmium can live as individual laments, with tens within heterocysts in other diazotrophs.
to hundreds of cells strung together, or in colonies consist- Instead of having localized stacks of thylakoids, Tri-
ing of tens to hundreds of laments clustered together.[2] chodesmium has unstacked thylakoids found throughout
These colonies are visible to the naked eye and some- the cell. Trichodesmium is highly vacuolated and the
times form blooms, which can be extensive on surface content and size of the vacuoles shows diurnal variation.
waters. These large blooms led to widespread recognition Large gas vesicles (either along the periphery as seen
as sea sawdust/straw"; in fact, the Red Sea gets most of in T. erythaeum or found distributed throughout the cell
its eponymous colouration from the corresponding pig- as seen in T. thiebautii) allow Trichodesmium to regu-
ment in Trichodesmium erythraeum. Colonies of Tri- late buoyancy in the water column. These gas vesicles
chodesmium provide a pseudobenthic substrate for many can withstand high pressure, presumably those up to 100
small oceanic organisms including bacteria, diatoms, 200 m in the water column, allowing Trichodesmium
dinoagellates, protozoa, and copepods (which are its pri- to move vertically through the water column harvesting
mary predator); in this way, the genus can support com- nutrients.[6]

1
2 4 ECOLOGY

3 Nitrogen-xation
N2 is the most abundant element within the atmosphere.
However, nitrogen packed in this form is not usable for
most biological processes. Nitrogen xation is the pro-
cess of converting inert atmospheric N2 into biologically
usable forms of nitrogen such as ammonium and nitrogen
oxides. This process requires a substantial amount of en-
ergy (in the form of ATP) in order to break the triple bond
between the nitrogen atoms.[7]
Trichodesmium is the major diazotroph in marine pelagic
systems [6] and is an important source of new nitrogen
in the nutrient poor waters it inhabits. Nitrogen xation Trichodesmium erythraeum bloom, between Vanuatu and New
in Trichodesmium is unique among diazotrophs because Caledonia, SW Pacic Ocean.
the process occurs concurrently with oxygen production
(via photosynthesis[1] ). In other cyanobacteria, N2 and released into dissolved pools, or get exported to the deep
CO2 reduction are separated either in space (using hete- sea.[7]
rocysts to protect the sensitive nitrogenase enzyme from
oxygen) or time. However, Trichodesmium lacks hete- Compared to eukaryotic phytoplankton, Trichodesmium
rocysts and nitrogen xation peaks during daylight hours has a slow growth rate, which has been hypothesized to
(following a diel ux initiated in the morning, reaching be an adaptation to survival in high energy but low nutri-
a maximum xation rate midday, and ceasing activity at ent conditions of oligotrophic waters. Growth rate is lim-
night).[6] Since the rst realization of this enigma, Tri- ited by iron and phosphate concentrations in the water. In
chodesmium has been the focus of many studies to try and order to obtain these limiting nutrients, Trichodesmium
discover how nitrogen xation is able to occur in the pres- is able to regulate buoyancy using its gas vacuole and
ence of oxygen production without any apparent structure move vertically throughout the water column, harvesting
separating the two processes. nutrients.[9]
Inhibitor studies even revealed that photosystem II ac-
tivity is essential for nitrogen xation in this organ- 4.1 Colonies
ism. All this may seem contradictory at rst glance,
because the enzyme responsible for nitrogen xation, Various species of Trichodesmium have been described
nitrogenase, is irreversibly inhibited by oxygen. How- based on morphology and structure of colonies formed.
ever, Trichodesmium utilises photosynthesis for nitrogen Colonies may consist of aggregates of several to several
xation by carrying out the Mehler reaction, during which hundred trichomes and form fusiform (called Tufts)
the oxygen produced by PSII is reduced again after PSI. colonies when aligned in parallel, or spherical (called
This regulation of photosynthesis for nitrogen xation in- Pus) colonies when aligned radially.[6]
volves rapidly reversible coupling of their light-harvesting
Trichodesmium colonies have been shown to have large
antenna, the phycobilisomes, with PSI and PSII.[8]
degree of associations with other organisms, includ-
ing bacteria, fungi, diatoms, copepods, tunicates, hy-
drozoans, and protozoans among other groups. These
4 Ecology colonies may provide a source of shelter, buoyancy, and
possibly food in the surface waters. Most of these associ-
Trichodesmium is found in oligotrophic waters, often ations appear to be commensal, with the Trichodesmium
when waters are calm and the mixed layer depth is shal- providing substrate and nutrition while deriving no ob-
low (around 100 m).[9] Trichodesmium is found primar- vious benet from the organisms dwelling within the
ily in water between 20 and 34 C and is frequently en- colonies.[11]
countered in tropical and sub-tropical oceans in western
boundary currents.[9] Its presence is more pronounced
4.2 Blooms
in nitrogen poor water and can easily be seen when
blooms form, trapping large Trichodesmium colonies at Trichodesmium forms large, visible blooms in the surface
the surface.[10] waters. Blooms have been described in the Baltic Sea,
As a diazotroph, Trichodesmium contributes a large por- the Red Sea, the Caribbean Sea, the Indian Ocean, the
tion of the marine ecosystems new nitrogen, estimated North and South Atlantic and the North Pacic, and o
to produce between 60 and 80 Tg of nitrogen per year.[8] the coast of Australia.[12] One of the earliest blooms was
Nitrogen xed by Trichodesmium can either be used di- described by E. Dupont in the Red Sea, noticed for turn-
rectly by the cell, enter the food chain through grazers, be ing the surface of the water a reddish color. This bloom

was said to extend about 256 nautical miles. Most blooms
are several kilometers long and last one to several months.
Blooms can form in coastal or oceanic waters, most fre-
quently when the water has been still for some time and
surface temperatures exceed 27 C.[13]
Trichodesmium blooms release carbon, nitrogen and
other nutrients into the environment. Some species of
Trichodesmium have been shown to release toxins which
cause mortalities in some copepods, sh, and oysters. [9] Capone D.G.; Zehr J.P.; Paerl H.W.; Bergman B.; Car-
Blooms have also been credited with releasing the toxin
which causes clupeotoxism in humans after ingesting
sh which have bioaccumulated the toxin during Tri- [10] Post, A.F. (2005). Huisman J.; Matthijs H.C.P.; Visser
chodesmium blooms. The larger impact of these blooms
is likely important to the oceanic ecosystem and is the
source of many studies.[8] Blooms are traced and tracked
using satellite imaging where the highly reective gas vac-
uole makes Trichodesmium blooms easily detectable.[14] [11] O'Neil J.M.; Roman M.R. (1991). Carpenter E.J.;
It is expected that blooms may increase due to anthro-
pogenic eects in the coming years. Phosphate loading
of the environment (through fertilizer pollution, waste
disposal, and mariculture) will reduce the growth con-
straints associated with limited phosphate and likely in- [12] Sellner K.G. (1991). Carpenter E.J.; Capone D.G.;
crease bloom occurrences.[10] Likewise, global warming
is projected to increase stratication and cause a shallow-
ing of the mixed layer depth. Both of these factors are as-
sociated with Trichodesmium blooms and may also cause
[8]
an increase in the occurrence of blooms in the future.
5 References
[1] Carpenter, E.J., Capone, D.G., Rueter, J.G. eds. (1991).
Marine Pelagic Cyanobacteria: Trichodesmium and other
diazothrophs. Dordrecht.: Kluwer Academic Publishers.
[2] Capone, Douglas G.; Zehr, Jonathan P.; Paerl, Hans
W.; Bergman, Birgitta; Carpenter, Edward J. (1997-05-
23). Trichodesmium, a Globally Signicant Marine
Cyanobacterium. Science. 276 (5316): 12211229.
doi:10.1126/science.276.5316.1221. ISSN 0036-8075.
[3] Ehrenberg,E.O. (1830). Neue Beobachtungen uber
blauartige. Erscheinungen in Aegypten und Siberien nebst
einer Ubersicht und Kritik der fruher bekannten.. Ann
Phys Chem. 18: 477514.
[4] Gomont, M. (1892). Monographie des oscillariees (Nos-
tocacees Homocystees) I and II.. Ann Sci Nat Bot Ser. 7
(15): 263368.
[5] Wille, N. (1904). Die Schizophyceen der plankton-
expedition.Ergebnisse der Plankton-Expedition der
Humbol-Stiftung.(Hensen,V., ed.)".
[6] Siddiqui P.J.A.; Carpenter E.J.; Bergman B. (1991).
Carpenter E.J.; Capone D.G.; Rueter J.G., eds. "Tri-
chodesmium: Ultrastructure and protein localization.
Marine Pelagic Cyanobacteria: Trichodesmium and other
diazotrophs. Dordrecht: Kluwer Academic Publishers.
3
[7] Zehr J.P. (2008). Kirchmad., ed. Molecular ecological
aspects of nitrogen xation in the marine environment.
Microbial Ecology of the Oceans. Wiley Blackwell.
[8] Bergman, B.; Sandh, G.; Lin, S.; Larsson, H. & Car-
penter, E.J. (2012). "Trichodesmium a widespread
marine cyanobacterium with unusual nitrogen xation
properties.. FEMS Microbiology Reviews: 117.
doi:10.1111/j.1574-6976.2012.00352.x.
penter E.J. (1997). "Trichodesmium, a globally signicant
marine cyanobacterium.. Science. 276: 12211229.
P.M., eds. Nutrient limitation of marine cyanobacte-
ria: Molecular ecology of nitrogen limitation in an olig-
otrophic sea. Harmful Cyanobacteria. Netherlands:
Springer: 87108.
Capone D.G.; Rueter J.G., eds. Grazers and associated
organisms of Trichodesmium". Marine Pelagic Cyanobac-
teria: Trichodesmium and other Diazotrophs. Dordrecht:
Kluwer Academic Publishers: 6174.
Rueter J.G., eds. Trophodynamics of marine cyanobac-
teria blooms. Marine Pelagic Cyanobacteria: Tri-
chodesmium and other Diazotrophs. Dordrecht: Kluwer
Academic Publishers: 7594.
[13] Carpenter E.J.; Capone D.G. (1991). Carpenter E.J.;
Capone D.G.; Rueter J.G., eds. Nitrogen xation in
Trichodesmium blooms. Marine Pelagic Cyanobacte-
ria: Trichodesmium and other Diazotrophs. Dordrecht:
Kluwer Academic Publishers: 7594.
[14] Zehr J.P.; Paerl H.W. (2008). Kirchman D.L., ed.
Molecular ecological aspects of nitrogen xation in the
marine environment. Microbial Ecology of the Oceans
2nd edition. Wiley-Blackwell.
6 Bibliography
Kana, T.M. (1993) Rapid oxygen cycling in Tri-
chodesmium thiebautii. Limnology and Oceanogra-
phy 38: 1824.
Berman-Frank, I., Lundgren, P., Chen, Y.-B., Kp-
per, H., Kolber, Z., Bergman, B., and Falkowski,
P. (2001) Segregation of nitrogen xation and oxy-
genic photosynthesis in the marine cyanobacterium
Trichodesmium. Science 294: 15341537.
Kpper, H., Ferimazova, N., etlk, I., and Berman-
Frank, I. (2004) Trac lights in Trichodesmium:
regulation of photosynthesis for nitrogen xation
studied by chlorophyll uorescence kinetic mi-
croscopy. Plant Physiology 135: 21202133.
Capone, D.G., Zehr, J., Paerl, H., Bergman, B., and
Carpenter, E.J. (1997) Trichodesmium: A globally
signicant marine cyanobacterium. Science 276:
12211229.
4 7 EXTERNAL LINKS

7 External links
Publications on Trichodesmium from a Marine Bio-
geochemistry laboratory at the University of South-
ern California

Charles Darwins description of sailing through a

Trichodesmium bloom

Trichodesmium in Florida 2004, Florida Fish

and Wildlife Conservation Commission Fish and
Wildlife Research Institute
 5

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