Spirodela polyrhiza
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''Spirodela polyrhiza'' ( ''S. polyrrhiza'') is a species of
duckweed Lemnoideae is a subfamily of flowering aquatic plants, known as duckweeds, water lentils, or water lenses. They float on or just beneath the surface of still or slow-moving bodies of fresh water and wetlands. Also known as bayroot, they arose f ...
known by the common names common duckmeat, greater duckweed, great duckmeat, common duckweed, and duckmeat. It can be found nearly worldwide in many types of
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does incl ...
habitat.


Description

''Spirodela polyrhiza'' is a
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also widel ...
aquatic plant usually growing in dense colonies, forming a mat on the water surface. Each plant is a smooth, round, flat disc 0.5 to 1.0 cm wide. Its upper surface is mostly green, sometimes red, while the lower surface is dark red. It produces several minute roots and a pouch containing male and female flowers. The top part dies in the fall and the plant often overwinters as a turion. The turion sinks to the bottom of the water body and stays in a dormant phase, until water temperature reaches 15 °C. The turions then germinate on the bottom of the water body and start a new life cycle. As this species lives in ponds and slow-moving water bodies, differs developmentally from terrestrial plants in morphology and
physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemic ...
. It undergoes mainly vegetative growth in spring and summer, forming new fronds. ''Spirodela polyrhiza'' rarely flowers. In fall and winter it switches into a dormant phase represented by the turions due to nutrition starvation and freezing temperatures. Because of its fast growth, direct contact with media and small genome size (~150 Mb), ''S. polyrhiza'' is an ideal system for biofuels, bioremediation, and carbon cycling. A comprehensive genomic study of ''S. polyrhiza'' was published in February 2014. The results provide insights into how this organism is adapted to rapid growth and an aquatic lifestyle.


Turion induction by abscisic acid

Turions were induced by the plant hormone
abscisic acid Abscisic acid (ABA) is a plant hormone. ABA functions in many plant developmental processes, including seed and bud dormancy, the control of organ size and stomatal closure. It is especially important for plants in the response to environmental s ...
(ABA) in the lab. Researchers reported that turions were rich in
anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical com ...
pigmentation and had a density that submerged them in liquid media.
Transmission electron microscopy Transmission electron microscopy (TEM) is a microscopy technique in which a beam of electrons is transmitted through a specimen to form an image. The specimen is most often an ultrathin section less than 100 nm thick or a suspension on a ...
of turions showed in comparison to fronds shrunken
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic m ...
s, smaller intercellular space, and abundant
starch granules Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets, ...
surrounded by
thylakoid Thylakoids are membrane-bound compartments inside chloroplasts and cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a thylakoid membrane surrounding a thylakoid lumen. Chloroplast thyl ...
membranes. Turions accumulated more than 60% starch in dry mass after two weeks of ABA treatment.


Distribution

''Spirodela polyrhiza'' is found worldwide, namely in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, more rarely in
Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known a ...
and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
, but also in
Central Europe Central Europe is an area of Europe between Western Europe and Eastern Europe, based on a common historical, social and cultural identity. The Thirty Years' War (1618–1648) between Catholicism and Protestantism significantly shaped the a ...
. It grows in
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
climates. It is not prevalent in
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
and only rarely in
Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
.


Cultivation

Large scale cultivation is done in outdoor water tanks, mostly in connection with
wastewater Wastewater is water generated after the use of freshwater, raw water, drinking water or saline water in a variety of deliberate applications or processes. Another definition of wastewater is "Used water from any combination of domestic, industrial ...
treatment. Tanks are fed with wastewater and the floating duckweed is harvested from the surface. It is then further used as a
biofuel Biofuel is a fuel that is produced over a short time span from biomass, rather than by the very slow natural processes involved in the formation of fossil fuels, such as oil. According to the United States Energy Information Administration ...
from industrial wastewater or as animal feed from agricultural wastewater treatment facilities.


Use

''Spirodela polyrhiza'' can be used for bioremediation, removing
toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a sub ...
substances from aquatic environment as well as cleaning
eutrophic Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytoplan ...
waters, especially in
wastewater Wastewater is water generated after the use of freshwater, raw water, drinking water or saline water in a variety of deliberate applications or processes. Another definition of wastewater is "Used water from any combination of domestic, industrial ...
treatment plants. Its uses as
biofuel Biofuel is a fuel that is produced over a short time span from biomass, rather than by the very slow natural processes involved in the formation of fossil fuels, such as oil. According to the United States Energy Information Administration ...
and animal feed are also gaining importance. It is hardly used for human nutrition.


Bioremediation

Because of its capability to hyperaccumulate
heavy metals upright=1.2, Crystals of osmium, a heavy metal nearly twice as dense as lead">lead.html" ;"title="osmium, a heavy metal nearly twice as dense as lead">osmium, a heavy metal nearly twice as dense as lead Heavy metals are generally defined as ...
and its high uptake of nutrients from the water, ''S. polyrhiza'' is used for bioremediation. The main pollutants it can be used to remediate are
arsenic Arsenic is a chemical element with the symbol As and atomic number 33. Arsenic occurs in many minerals, usually in combination with sulfur and metals, but also as a pure elemental crystal. Arsenic is a metalloid. It has various allotropes, b ...
(As) and mercury (Hg) and common wastewater nutrients, like sulphate (SO42-),
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
(PO43-) and
nitrate Nitrate is a polyatomic ion with the chemical formula . Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are soluble in water. An example of an insolu ...
(NO3).


Arsenic

Greater duckweed showed accumulation of arsenic in laboratory tests. Arsenic uptake was found to be negatively correlated with phosphate and positively correlated with
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in ...
uptake. This indicates that phosphate and arsenic compete for uptake by ''S. polyrhiza'', while arsenic's absorption is facilitated by iron oxides, because it shows an affinity to the root surface of ''S. polyrhiza'', where it is taken up. Greater duckweed is thought to detoxify the arsenic by reducing As (V) to the less toxic As (III). Difficulties arise with the management of the plants with high As contents. One possible use of the biomass containing As is production of charcoal and gas as a byproduct, which can be used as a fuel. The problems with this approach are low charcoal quality and high investments. Direct burning or burning of the coal is thought to release arsenic into the air, which would
pollute Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, the ...
the environment. Other options for
fuel A fuel is any material that can be made to react with other substances so that it releases energy as thermal energy or to be used for work. The concept was originally applied solely to those materials capable of releasing chemical energy b ...
production would be
hydrolysis Hydrolysis (; ) is any chemical reaction in which a molecule of water breaks one or more chemical bonds. The term is used broadly for substitution, elimination, and solvation reactions in which water is the nucleophile. Biological hydrolysi ...
and
fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food p ...
, which are economically not feasible. The
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...
would have to be treated with strong acids and heat, which are both capital intensive. Briquetting is considered one of the best options, where the plants are dried and pressed into pellets of briquets. This raises the question of whether the arsenic is released back into the environment during the burning process. The production of biogas is also considered, but again, the redistribution of the As has to be avoided .


Mercury

''Spirodela polyrhiza'' was found to be an efficient bio accumulator of mercuric chloride (HgCl2) in laboratory settings. Its plant biomass showed a 1000 times higher mercuric chloride concentration than its aquatic environment. ''Spirodela polyrhiza'' showed the highest accumulation factor compared to ''
Lemna gibba ''Lemna gibba'', the gibbous duckweed, swollen duckweed, or fat duckweed, is a species of ''Lemna'' (duckweed). It has a simple plant body, known as a thallus, which floats on the surface of the water and measures in diameter. A single root hang ...
'' and '' L. minor'', which were also investigated.


Urban wastewater treatment

''d''Greater duckweed has been used to remove common pollutants from wastewater. In a laboratory setting, ''S. polyrhiza'' showed a maximum of 90% removal efficiency of nitrate, 99.6% of phosphate and 69.8% of sulphate. The efficiency for all three pollutants combined was 85.6%, which makes it an environmentally and economically viable bioremediatory for wastewater treatment.


Biofuel

Due to space-efficient starch production and good growth in animal wastewater, ''S. polyrhiza'' has great potential in
bioethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a hyd ...
production. Despite environmental problems associated with production and competition from human and animal feed, corn is the main raw material for bioethanol. ''Spirodela polyrhiza'' could produce up to 50% more bioethanol on the same area. At the same time the production of bioethanol from ''S. polyrhiza'' is not in competition with human food. The production of bioethanol from ''S. polyrhiza'' is still in the development phase.


Animal feed

In small-scale agriculture ''S. polyrhiza'' is used as fish or poultry feed. Due to its fast growth and high protein content, it is an interesting feedstuff. Because of sanitary problems and the risk of heavy metal accumulation, it is not yet used for feeding in larger animal husbandry systems. For
rainbow trout The rainbow trout (''Oncorhynchus mykiss'') is a species of trout native to cold-water tributaries of the Pacific Ocean in Asia and North America. The steelhead (sometimes called "steelhead trout") is an anadromous (sea-run) form of the coast ...
, poorer growth rates were found when ''S. polyrhiza'' was added to the feed. For tilapia ( ''Oreochromis niloticus'' L.), greater weight gains were found when 30% of the
fish meal Fish meal is a commercial product made from whole wild-caught fish, bycatch and fish by-products to feed farm animals, e.g., pigs, poultry, and farmed fish.R. D. Miles and F. A. Chapman.FA122: The Benefits of Fish Meal in Aquaculture DietsFisher ...
in the feed was replaced with ''S. polyrhiza''. A review has also shown that duckweed can be used in cattle, pig and poultry diets. However, the problems of heavy metals and pathogen contamination occur.


Human nutrition

Although other duckweed species, such as ''
Wolffia arrhiza ''Wolffia arrhiza'' is a species of flowering plant known by the common names spotless watermeal and rootless duckweed, belonging to the Araceae, a family rich in water-loving species, such as ''Arum'' and ''Pistia''. It is the smallest vascula ...
'', are consumed by people in rural areas, ''S.polyrhiza'' is not cultivated for human consumption. This is because of high concerns about heavy metal accumulation and possible
contamination Contamination is the presence of a constituent, impurity, or some other undesirable element that spoils, corrupts, infects, makes unfit, or makes inferior a material, physical body, natural environment, workplace, etc. Types of contamination ...
with ''
Escherichia coli ''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Esc ...
'' or '' Clostridium botulinum''. In contrast to '' W. arrhiza'', ''S. polyrhiza'' contains, like most duckweed species,
calcium oxalate crystals Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrat ...
which are known to cause kidney stones.


References


External links


Spirodela polyrhiza genomicsJepson Manual TreatmentFlora of North AmericaWashington Burke Museum
{{Taxonbar, from=Q154500 Lemnoideae Plant models Freshwater plants