Azolla

Taxobox
name = Mosquito fern

"Azolla" (mosquito fern, duckweed fern, fairy moss, water fern) is a genus of seven species of aquatic ferns, the only genus in the family Azollaceae. They are extremely reduced in form and specialized, looking nothing like conventional ferns but more resembling duckweed or some mosses.

Ecology

"Azolla" floats on the surface of water by means of numerous, small, closely-overlapping scale-like leaves, with their roots hanging in the water. They form a symbiotic relationship with the blue-green alga "Anabaena azollae", which fixes atmospheric nitrogen, giving the plant access to the essential nutrient. This has led to the plant being dubbed a "super-plant", as it can readily colonise areas of freshwater, and grow at great speed - doubling its biomass every two to three days. The only known limiting factor on its growth is phosphorus, another essential mineral. An abundance of phosphorus, due for example to eutrophication or chemical runoff, often leads to "Azolla" blooms.

Their nitrogen-fixing capability of "Azolla" has led to it being widely used as a biofertiliser, especially in parts of southeast Asia. Indeed, the plant has been used to bolster agricultural productivity in China for over a thousand years. When rice paddies are flooded in the spring, they can be inoculated with "Azolla", which then quickly multiplies to cover the water, suppressing weeds. The rotting plant material releases nitrogen to the rice plants, providing up to nine tonnes of protein per hectare per year. [cite web|url=http://www.fao.org/ag/AGA/AGAP/FRG/afris/DATA/558.htm|title=FAO figures]

"Azolla" are also serious weeds in many parts of the world, entirely covering some bodies of water. The myth that no mosquito can penetrate the coating of fern to lay its eggs in the water gives the plant its common name "mosquito fern". [cite web |url=http://www.americaswetlandresources.com/wildlife_ecology/plants_animals_ecology/plants/mosquitofern.html |title=Mosquito Fern |accessdate=2007-11-10 |work=America's Wetland Resource Center |publisher=Loyola University, New Orleans]

Most of the species can produce large amounts of deoxyanthocyanins in response to various stresses, [Wagner, G.M., 1997. "Azolla": a review of its biology and utilization. Bot. Rev. 63, 1–26,] including bright sunlight and extremes of temperature, [Moore, A. W. 1969. "Azolla": Biology and agronomic significance. Bot. Rev. (Lancaster) 35: 17-35] [Zimmerman, 1985. Biomass and Pigment Production in Three Isolates of "Azolla" II. Response to Light and Temperature Stress. Ann. Bot. 56: 701-709.] , causing the water surface to appear to be covered with an intensely red carpet. Herbivore feeding induces accumulation of deoxyanthocyanins and leads to a reduction in the proportion of polyunsaturated fatty acids in the fronds, thus lowering their palatability and nutritive value. [Cohen, M.F., Meziane, T., Tsuchiya, M., Yamasaki, H. 2002. Feeding deterrence of "Azolla" in relation to deoxyanthocyanin and fatty acid composition. Aquatic Bot. 74:181–187]

"Azolla" cannot survive winters with prolonged freezing, so is often grown as an ornamental plant at high latitudes where it cannot establish itself firmly enough to become a weed. It is not tolerant to salinity; normal plants can't survive in greater than 1-1.6‰, and even conditioned organisms die in over 5.5‰ salinity.cite journal
author = Brinkhuis, H.
coauthors = Schouten, S.; Collinson, M.E.; Sluijs, A.; Sinninghe Damsté, J.S.; Dickens, G.R.; Huber, M.; Cronin, T.M.; Onodera, J.; Takahashi, K. "et al".
year = 2006
title = Episodic fresh surface waters in the Eocene Arctic Ocean
journal = Nature
volume = 441
pages = 7093
url = http://www.nature.com/search/executeSearch?sp-q-9=NATURE&sp-q=Episodic+fresh+surface+waters+in+the+Eocene+Arctic+Ocean&sp-c=10&sp-x-9=cat&sp-s=date&submit=go&sp-a=sp1001702d&sp-sfvl-field=subject%7Cujournal&sp-x-1=ujournal&sp-p-1=phrase&sp-p=all
accessdate = 2007-10-17
doi = 10.1038/nature04692]

Reproduction

"Azolla" reproduces sexually, and asexually by splitting.

Like all ferns, sexual reproduction leads to spore formation, but "Azolla" sets itself apart from other members of its group by producing two kinds. During the summer months, numerous spherical structures called sporocarps form on the undersides of the branches. The male sporocarp is greenish or reddish and looks like the egg mass of an insect or spider. It is two millimeters in diameter, and inside are numerous male sporangia. Male spores (microspores) are extremely small and are produced inside each microsporangium. Curiously, microspores tend to adhere in clumps called massulae.Fact|date=October 2007

Female sporocarps are much smaller, containing one sporangium and one functional spore. Since an individual female spore is considerably larger than a male spore, it is termed a megaspore.

"Azolla" has microscopic male and female gametophytes that develop inside the male and female spores. The female gametophyte protrudes from the megaspore and bears a small number of archegonia, each containing a single egg. The microspore forms a male gametophyte with a single antheridium which produces eight swimming sperm. [AN EVOLUTIONARY SURVEY OF THE PLANTKINGDOM, by Robert F. Scagel, Robert J. Bandoni, Glenn E. Rouse, W. B. Schofield,Janet R. Stein, and T. M. Taylor, 1965: Belmont, California, Wadsworth Publishing Co.,Inc., 658 pp.] The barbed glochidia on the male spore clusters are assumed to cause them to cling to the female megaspores, thus facilitating fertilization.

Use as Food

In addition to its traditional cultivation as a bio-fertilizer for wetland paddy, Azolla is finding increasing use for sustainable production of livestock feed. [cite web |url=http://www.leisa.info/index.php?url=show-blob-html.tpl&p%5Bo_id%5D=76755&p%5Ba_id%5D=211&p%5Ba_seq%5D=1 |title=Azolla – A sustainable feed substitute for livestock|accessdate=2008-01-14 |author=P. Kamalasanana Pillai, S. Premalatha and S. Rajamony|publisher=LEISA India] Azolla is rich in proteins, essential amino acids, vitamins and minerals. Studies describe feeding azolla to dairy cattle, pigs, ducks, and chickens, with reported increases in milk production, weight of broiler chickens and egg production of layers, as compared to conventional feed. One FAO study describes how azolla integrates into a tropical biomass agricultural system, reducing the need for inputs. [cite web |url=http://www.fao.org/ag/AGAInfo/resources/documents/WAR/war/U7600B/u7600b04.htm |title=Sustainable intensive livestock systems for the humid tropics|accessdate=2008-01-14 |author=T.R. Preston and E. Murgueitio|publisher=FAO]

Climatic paleontology

A study of Arctic climatology reported that azolla may have had a significant role in reversing a greenhouse effect that occurred 55 million years ago that caused the region around the north pole to turn into a hot tropical environment. This research conducted by the Institute of Environmental Biology at Utrecht University claims that large dense patches of "Azolla" growing around freshwater lakes formed by the climate change eventually consumed enough carbon dioxide for the greenhouse effect to reverse.

See also

* Aquatic plant

References

*Scagel, R. F., Bandoni, R. J., Rouse, G. E., Schofield, W. B., Stein, J. R., & Taylor, T. M. C. (1966). "An Evolutionary Survey of the Plant Kingdom". Wadsworth.

External links

* [http://aquaplant.tamu.edu/database/floating_plants/mosquito_fern_mgmt.htm aquaplant.tamu.edu]
* [http://www.ars-grin.gov/cgi-bin/npgs/html/splist.pl?1209 Germplasm Resources Information Network: "Azolla"]
* [http://www.efloras.org/florataxon.aspx?flora_id=3&taxon_id=103297 Flora of China: "Azolla" species list]
* [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=103297 Flora of North America: "Azolla"]
* [http://waynesword.palomar.edu/plnov98.htm Marriage Between A Fern & Cyanobacterium]
* [http://plants.usda.gov/java/profile?symbol=AZPI plants.usda.gov]
* [http://www.royalbcmuseum.bc.ca/end_species/species/mfern.html royalbcmuseum.bc.ca]