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Lemnoideae is a subfamily of flowering
aquatic plant Aquatic plants are plants that have adapted to living in aquatic environments ( saltwater or freshwater). They are also referred to as hydrophytes or macrophytes to distinguish them from algae and other microphytes. A macrophyte is a plant that ...
s, 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 from within the arum or aroid family ( Araceae), so often are classified as the subfamily Lemnoideae within the family Araceae. Other classifications, particularly those created prior to the end of the twentieth century, place them as a separate family, Lemnaceae. These plants have a simple structure, lacking an obvious stem or leaves. The greater part of each plant is a small organized " thallus" or "
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
" structure only a few cells thick, often with air pockets ( aerenchyma) that allow it to float on or just under the water surface. Depending on the species, each plant may have no root or may have one or more simple rootlets. Reproduction is mostly by asexual
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
( vegetative reproduction), which occurs from a meristem enclosed at the base of the frond. Occasionally, three tiny "flowers" consisting of two stamens and a pistil are produced, by which
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
occurs. Some view this "flower" as a pseudanthium, or reduced
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed ...
, with three flowers that are distinctly either female or male and which are derived from the spadix in the Araceae. Evolution of the duckweed inflorescence remains ambiguous due to the considerable evolutionary reduction of these plants from their earlier relatives. The flower of the duckweed genus '' Wolffia'' is the smallest known, measuring merely 0.3 mm long. The fruit produced through this occasional reproduction is a utricle, and a
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
is produced in a bag containing air that facilitates flotation.


Duckweed in natural environments

One of the more important factors influencing the distribution of wetland plants, and aquatic plants in particular, is nutrient availability. Duckweeds tend to be associated with fertile, even eutrophic conditions. They can be spread by waterfowl and small mammals, transported inadvertently on their feet and bodies, as well as by moving water. In water bodies with constant currents or overflow, the plants are carried down the water channels and do not proliferate greatly. In some locations, a cyclical pattern driven by weather patterns exists in which the plants proliferate greatly during low water-flow periods, then are carried away as rainy periods ensue. Duckweed is an important high-
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respon ...
food source for waterfowl. The tiny plants provide cover for fry of many aquatic species. The plants are used as shelter by pond-water species such as bullfrogs and fish such as bluegills. They also provide shade and, although frequently confused with them, can reduce certain light-generated growths of photoautotrophic
algae Algae ( , ; : alga ) are any of a large and diverse group of photosynthetic, eukaryotic organisms. The name is an informal term for a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from ...
.


Use as human food crop

Duckweed is consumed as a poor person's food in some parts of Southeast Asia, namely Laos,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is b ...
, and
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John Wells explai ...
. It contains more protein than soybeans, so sometimes it is cited as a significant potential food source. Some initial investigations to what extent duckweed could be introduced in European markets show little consumer objection to the idea. NASA's Caves of Mars Project identified duckweed as a top candidate for growing food on Mars.


Invasive species

Despite some of these benefits, because duckweed thrives in high-nutrient wetland environments, the plants can be seen as a nuisance species when conditions favor excessive proliferation in environments that are traditionally low in nutrients or oligotrophic. One example of this problem occurs within the Everglades, a mostly oligotrophic environment, when excess chemicals (that include fertilizers) are carried by storm runoff, or surface runoff, into its waterways. Urban runoff and agricultural pollution then begin to introduce increased levels of nutrients into the surrounding wetlands and waterways, which can cause a disruption to native ecology. These conditions allow for the invasion of a fast growing species such as duckweed to establish themselves, spread, and displace other native species such as sawgrass, and over time, result in widespread changes to the ecology of native sawgrass and slough habitats within the Everglades.


Taxonomy

The duckweeds have long been a taxonomic mystery, and usually have been considered to be their own family, the Lemnaceae. They primarily reproduce asexually. Flowers, if present at all, are small. Roots are either very much reduced, or absent entirely. They were suspected of being related to the Araceae as long ago as 1876, but until the advent of molecular phylogeny, testing this hypothesis was difficult. Starting in 1995, studies began to confirm their placement in the Araceae and since then, most systematists consider them to be part of that family. Their position within their family has been slightly less clear; however, several twenty-first century studies place them in the position shown below. Although they are in the same family as '' Pistia'', another aquatic plant, they are not closely related. The genera of duckweeds are: '' Spirodela'', ''
Landoltia ''Spirodela punctata'' (or ''Landoltia punctata'', common name dotted duckmeat) is a species of duckweed (Lemnoideae). The species is morphologically intermediate between ''Lemna'' and other species of ''Spirodela ''Spirodela'' is a genus of ...
'', '' Lemna'', '' Wolffiella'', and '' Wolffia''. Duckweed
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
sizes have a ten-fold range (150~1,500 MB), potentially representing diploids to octaploids. The ancestral genus of ''Spirodela'' has the smallest genome size (150 MB, similar to '' Arabidopsis thaliana''), while the most derived genus, ''Wolffia'', contains plants with the largest genome size (1,500 MB). DNA sequencing has shown that ''Wolffiella'' and ''Wolffia'' are more closely related than the others. ''Spirodela'' is at the
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
position of the taxon, followed by ''Lemna'', ''Wolffiella'', and ''Wolffia'', which is the most derived. To identify different duckweed genomes, a DNA-based molecular identification system was developed based on seven plastid-markers proposed by the Consortium for the Barcode of Life. The ''atpF-atpH'' non-coding spacer was chosen as a universal DNA barcoding marker for species-level identification of duckweeds.


Fossil record

Extinct free-floating aquatic plants and
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
with affinities to the Lemnoideae first appear in the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
( Maastrichtian) as evidenced by floating leaves described as ''Aquaephyllum auriculatum'' from
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, t ...
, and the lemnoid pollen
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
''Pandaniidites''. Fossils of floating leaves with rootlets from the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
of southern
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,
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, that were originally described as ''Lemna'' (''Spirodela'') ''scutata'' by John William Dawson in 1885, have been redescribed as ''Limnobiophyllum''. In addition to western North America, ''Limnobiophyllum'' has been reported from the Paleocene of eastern Russia and the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
of the
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. Unusually complete specimens from the Paleocene of
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
, Canada, range from single leaves up to about 4 cm in diameter to rosettes of up to four leaves, some of which were connected to adjacent plants by stolons, and a few of which bear remains of
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
s with anthers that contain ''Pandaniidites'' pollen. Occurrences of lemnoid seeds described as ''Lemnospermum'' have also been reported.


Research and applications

Research and applications of duckweeds are promoted by two international organizations, The International Lemna Association and the International Steering Committee on Duckweed Research and Applications. In July 2008, the
U.S. Department of Energy The United States Department of Energy (DOE) is an executive department of the U.S. federal government that oversees U.S. national energy policy and manages the research and development of nuclear power and nuclear weapons in the United States. ...
(DOE) Joint Genome Institute announced that the Community Sequencing Program would fund sequencing of the genome of the giant duckweed, '' Spirodela polyrhiza''. This was a priority project for DOE in 2009. The research was intended to facilitate new
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 biom ...
and bioenergy programs. The results were published in February 2014. They provide insight into how this plant is adapted to rapid growth and an aquatic lifestyle.


Potential clean energy source

Duckweed is being studied by researchers around the world as a possible source of clean energy. In the U.S, in addition to being the subject of study by the DOE, both
Rutgers University Rutgers University (; RU), officially Rutgers, The State University of New Jersey, is a public land-grant research university consisting of four campuses in New Jersey. Chartered in 1766, Rutgers was originally called Queen's College, and wa ...
and North Carolina State University have ongoing projects to determine whether duckweed might be a source of cost-effective, clean, renewable energy. Duckweed is a good candidate as a biofuel because it grows rapidly, produces five to six times as much starch as corn per unit of area, and does not contribute to
global warming In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate variability and change, Climate change in a broader sense also includes ...
. Duckweed removes carbon dioxide from the atmosphere, and it may have value for climate change mitigation.


Filtration of contaminants and nutrients

The plants can provide nitrate removal, if cropped, and the duckweeds are important in the process of bioremediation because they grow rapidly, absorbing excess mineral nutrients, particularly nitrogen and phosphates. For these reasons, they are touted as water purifiers of untapped value. The Swiss Department of Water and Sanitation in Developing Countries, associated with the Swiss Federal Institute for Environmental Science and Technology, asserts that as well as the food and agricultural values, duckweed also may be used for wastewater treatment to capture toxins and for odor control, and that if a mat of duckweed is maintained during harvesting for removal of the toxins captured thereby, it prevents the development of algae and controls the breeding of mosquitoes. The same publication provides an extensive list of references for many duckweed-related topics. These plants also may play a role in conservation of water because a cover of duckweed will reduce evaporation of water when compared to the rate of a similarly sized water body with a clear surface. Duckweed also functions as a bioremediator by effectively filtering contaminants such as bacteria, nitrogen, phosphates, and other nutrients from naturally occurring bodies of water, constructed wetlands, and wastewater. File:LentejasDeAgua.jpg,
Common duckweed ''Lemna minor'', the common duckweed or lesser duckweed, is a species of aquatic freshwater plant in the subfamily Lemnoideae of the arum family Araceae. ''L. minor'' is used as animal fodder, bioremediator, for wastewater nutrient recovery, and ...
in
Galicia Galicia may refer to: Geographic regions * Galicia (Spain), a region and autonomous community of northwestern Spain ** Gallaecia, a Roman province ** The post-Roman Kingdom of the Suebi, also called the Kingdom of Gallaecia ** The medieval King ...
, Spain File:Lemna minor1.jpg, '' Lemna minor'' File:Lemna trisulca0.jpg, '' Lemna trisulca'' File:L gibba3.jpg, '' Lemna gibba''
Turning the canals of the Poitevin Marsh ( Marais Poitevin, France) into the "Green Venice": File:Spirodela polyrrhiza marais poitevin.jpg, '' Spirodela polyrhiza'' File:Curve of duckweed covered water edged with several bald cypress trees.JPG, Duckweed-covered water edged with several bald cypress trees A start-up, microTERRA, based in Mexico has attempted to use duckweed as clean water in privately owned aquaculture farms. The plants use nitrogen and phosphorus produced from fish waste as fertilizer, while simultaneously cleaning the water as it grows. The water can then be reused by the aquaculture farmers, and the duckweed, which has a 35-42% protein content, can be harvested as a source of sustainable protein.


See also

* Phytoremediation * Hyperaccumulators table – 3 * Neuston


References


External links

* * * : : * * {{Taxonbar, from=Q161429, from2=Q14293890 Aquatic plants Alismatales subfamilies Phytoremediation plants