Azolla
| Mosquito fern | |
|---|---|
| |
| Azolla caroliniana | |
| Scientific classification | |
| Kingdom: | |
| Division: | |
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| Genus: | Azolla |
| Type species | |
| A. filiculoides[1] | |
| Species | |
|
See list below | |
Azolla (mosquito fern, duckweed fern, fairy moss, water fern) is a genus of seven species of aquatic ferns in the family Salviniaceae. They are extremely reduced in form and specialized, looking nothing like conventional ferns but more resembling duckweed or some mosses.
Selected species
Azolla caroliniana Willd.
[ Poss. synon. of A. filiculoides ]
Azolla circinata Oltz & Hall
Azolla filiculoides Lam.
Azolla japonica Franch. & Sav.
[ Poss. synon. of A. filiculoides ]
Azolla mexicana C.Presl
Azolla microphylla Kaulf.
Azolla nilotica Decne. ex Mett.
Azolla pinnata R.Br.
Azolla rubra R.Br.
[ Poss. synon. of A. filiculoides ]
At least six extinct species are known from the fossil record:
- Azolla intertrappea Sahni & H.S. Rao, 1934 (Eocene, India)[5]
- Azolla berryi Brown, 1934 (Eocene, Green River Formation, Wyoming)[5]
- Azolla prisca Chandler & Reid, 1926 (Oligocene, London Clay, Isle of Wight)[5]
- Azolla tertiaria Berry, 1927 (Pliocene, Esmeralda Formation, Nevada)[5]
- Azolla primaeva (Penhallow) Arnold, 1955 (Eocene, Allenby Formation, British Columbia)[5]
- Azolla boliviensis Vajda & McLoughlin, 2005 (Maastrichtian - Paleocene, Eslaboacuten Formation& Flora Formation Bolvia)[6]
Ecology
'Description Azolla filiculoides (Red Azolla) Azolla filiculoides (Red Azolla) is the only member of this genus and of the family Azollaceae in Tasmania. It is a very common native aquatic plant in Tasmania. It is particularly common on farm dams and other still water bodies. The plants are small (usually only a few cm across) and float, but can be very abundant and form large mats. The plants are typically red, and have very small water repellent leaves.'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 cyanobacterium 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.
The nitrogen-fixing capability of Azolla has led to Azolla 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.[7]
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".[8]
Most of the species can produce large amounts of deoxyanthocyanins in response to various stresses,[9] including bright sunlight and extremes of temperature,[10][11] 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.[12]
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.[13]
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.[5]
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.[14] The barbed glochidia on the male spore clusters are assumed to cause them to cling to the female megaspores, thus facilitating fertilization.
Human Use
Food
In addition to its traditional cultivation as a bio-fertilizer for wetland paddy (due to its ability to fix nitrogen), azolla is finding increasing use for sustainable production of livestock feed.[15] 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.[16]
Companion plant
Azolla has been used, for at least one thousand years [1], in rice paddies as a companion plant, because of its ability to both fix nitrogen, and block out light to prevent any competition from other plants, aside from the rice, which is planted when tall enough to poke out of the water through the azolla layer.
Larvicide
As an additional benefit to its role as a paddy biofertilizer, Azolla spp. have been used to control mosquito larvae in rice fields. The plant grows in a thick mat on the surface of the water, reducing the rate at which oxygen dissolves into the water, effectively choking the larvae.[17]
Climatic paleontology
A study of Arctic climatology reported that azolla may have had a significant role in reversing an increase in 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
References
- ^ a b In: Encyclopédie Méthodique, Botanique 1(1): 343. 1783. "Name - Azolla Lam". Tropicos. Saint Louis, Missouri: Missouri Botanical Garden. Retrieved February 19, 2010.
Annotation: a sp. nov. reference for Azolla filiculoides
Type Specimens HT: Azolla filiculoides - ^ "Name - Azolla Lam. subordinate taxa". Tropicos. Saint Louis, Missouri: Missouri Botanical Garden. Retrieved February 19, 2010.
- ^ "Query Results for Genus Azolla". IPNI. Retrieved February 19, 2010.
- ^ Hussner, A. (2006). "NOBANIS -- Invasive Alien Species Fact Sheet -- Azolla filiculoides" (PDF). Online Database of the North European and Baltic Network on Invasive Alien Species. Heinrich Heine Universität, Düsseldorf. Retrieved February 19, 2010.
- ^ a b c d e f Arnold, C.A. (1955). "A Tertiary Azolla from British Columbia" (PDF). Contributions from the Museum of. Paleontology, University of Michigan. 12 (4): 37–45.
- ^ Vajda, V (2005). "A new Maastrichtian-Paleocene Azolla species from of Bolivia, with a comparison of the global record of coeval Azolla microfossils". Alcheringa: an Australasian Journal of Palaeontology. 29 (2): 305–329. doi:10.1080/03115510508619308.
{{cite journal}}: Unknown parameter|coauthor=ignored (|author=suggested) (help) - ^ "FAO figures".
- ^ "Mosquito Fern". America's Wetland Resource Center. Loyola University, New Orleans. Retrieved 2007-11-10.
- ^ Wagner, G.M. (1997). "Azolla: a review of its biology and utilization". Bot. Rev. 63: 1–26. doi:10.1007/BF02857915.
- ^ Moore, A. W. (1969). "Azolla: Biology and agronomic significance". Bot. Rev. 35: 17–35. doi:10.1007/BF02859886.
- ^ Zimmerman (1985). "Biomass and Pigment Production in Three Isolates of Azolla II. Response to Light and Temperature Stress". Ann. Bot. 56: 701–709.
{{cite journal}}: Cite has empty unknown parameter:|author-name-separator=(help); Unknown parameter|author-separator=ignored (help) - ^ Cohen, M.F.; Meziane, T.; Tsuchiya, M.; Yamasaki, H. (2002). "Feeding deterrence of Azolla in relation to deoxyanthocyanin and fatty acid composition" (PDF). Aquatic Bot. 74: 181–187. doi:10.1016/S0304-3770(02)00077-3.
- ^ Brinkhuis, H. (2006). "Episodic fresh surface waters in the Eocene Arctic Ocean". Nature. 441 (7093): 7093. doi:10.1038/nature04692. PMID 16752440. Retrieved 2007-10-17.
{{cite journal}}: Unknown parameter|coauthors=ignored (|author=suggested) (help) - ^ AN EVOLUTIONARY SURVEY OF THE PLANT KINGDOM, 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.
- ^ P. Kamalasanana Pillai, S. Premalatha and S. Rajamony. "Azolla – A sustainable feed substitute for livestock". Farming Matters magazine. Retrieved 2008-01-14.
- ^ T.R. Preston and E. Murgueitio. "Sustainable intensive livestock systems for the humid tropics". FAO. Retrieved 2011-09-28.
- ^ Myer, Landon; Okech, Bernard A.; Mwobobia, Isaac K.; Kamau, Anthony; Muiruri, Samuel; Mutiso, Noah; Nyambura, Joyce; Mwatele, Cassian; Amano, Teruaki (2008). "Use of Integrated Malaria Management Reduces Malaria in Kenya". PLoS ONE. 3: e4050. doi:10.1371/journal.pone.0004050.
{{cite journal}}:|access-date=requires|url=(help)CS1 maint: unflagged free DOI (link)
- 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.
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