Duck Weed
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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 from within the arum or aroid family (
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). A ...
), so often are classified as the subfamily Lemnoideae within the family
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). A ...
. 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 Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms ...
" 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 Aerenchyma or aeriferous parenchyma or lacunae, is a modification of the parenchyma to form a spongy tissue that creates spaces or air channels in the leaves, stems and roots of some plants, which allows exchange of gases between the shoot and th ...
) 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 Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or spec ...
), which occurs from a
meristem The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...
enclosed at the base of the frond. Occasionally, three tiny "flowers" consisting of two
stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
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 A pseudanthium (Greek for "false flower"; ) is an inflorescence that resembles a flower. The word is sometimes used for other structures that are neither a true flower nor a true inflorescence. Examples of pseudanthia include flower heads, compos ...
, or reduced
inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
, 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 ''Wolffia'' is a genus of aquatic plants with a cosmopolitan distribution. They include the smallest flowering plants on Earth. Commonly called watermeal or duckweed, these aquatic plants resemble specks of cornmeal floating on the water. Indivi ...
'' 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 angiospe ...
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 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 ...
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, res ...
food source for
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which i ...
. The tiny plants provide cover for fry of many aquatic species. The plants are used as shelter by pond-water species such as
bullfrogs ''Bullfrog'' is a common English language term to refer to large, aggressive frogs, regardless of species. Examples of bullfrogs include: Frog species America *Helmeted water toad (''Calyptocephalella gayi''), endemic to Chile * American bullfr ...
and fish such as bluegills. They also provide shade and, although frequently confused with them, can reduce certain light-generated growths of
photoautotrophic Photoautotrophs are organisms that use light energy and inorganic carbon to produce organic materials. Eukaryotic photoautotrophs absorb energy through the chlorophyll molecules in their chloroplasts while prokaryotic photoautotrophs use chlorophyll ...
algae.


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
soybean The soybean, soy bean, or soya bean (''Glycine max'') is a species of legume native to East Asia, widely grown for its edible bean, which has numerous uses. Traditional unfermented food uses of soybeans include soy milk, from which tofu a ...
s, 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 The Caves of Mars Project was an early 2000s program funded through Phase II by the NASA Institute for Advanced Concepts to assess the best place to situate the research and habitation modules that a human mission to Mars would require. The f ...
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 The Everglades is a natural region of tropical wetlands in the southern portion of the U.S. state of Florida, comprising the southern half of a large drainage basin within the Neotropical realm. The system begins near Orlando with the Kissim ...
, a mostly oligotrophic environment, when excess chemicals (that include fertilizers) are carried by storm runoff, or surface runoff, into its waterways.
Urban runoff Urban runoff is surface runoff of rainwater, landscape irrigation, and car washing created by urbanization. Impervious surfaces (roads, parking lots and sidewalks) are constructed during land development. During rain , storms and other precip ...
and
agricultural pollution Agricultural pollution refers to biotic material, biotic and abiotic byproducts of agriculture, farming practices that result in pollution, contamination or degradation of the environment and surrounding ecosystems, and/or cause injury to human ...
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 Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, 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 ''Spirodela'' is a genus of aquatic plants, one of several genera containing plants commonly called duckweed. ''Spirodela'' species are members of the Araceae under the APG II system. They were formerly members of the Lemnaceae. ''Spirodela' ...
'', '' Landoltia'', ''
Lemna ''Lemna'' is a genus of free-floating aquatic plants referred to by the common name "duckweed". They are morphologically divergent members of the arum family Araceae. These rapidly growing plants have found uses as a model system for studies in ...
'', '' Wolffiella'', and ''
Wolffia ''Wolffia'' is a genus of aquatic plants with a cosmopolitan distribution. They include the smallest flowering plants on Earth. Commonly called watermeal or duckweed, these aquatic plants resemble specks of cornmeal floating on the water. Indivi ...
''. 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 g ...
sizes have a ten-fold range (150~1,500 MB), potentially representing
diploids Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respective ...
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 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 Consortium for the Barcode of Life (CBOL) was an international initiative dedicated to supporting the development of DNA barcoding as a global standard for species identification. CBOL's Secretariat Office is hosted by the National Museum of ...
. The ''atpF-atpH'' non-coding spacer was chosen as a universal
DNA barcoding DNA barcoding is a method of species identification using a short section of DNA from a specific gene or genes. The premise of DNA barcoding is that by comparison with a reference library of such DNA sections (also called " sequences"), an indi ...
marker for species-level identification of duckweeds.


Fossil record

Extinct free-floating aquatic plants and pollen 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 The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...
) 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 g ...
,
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, th ...
, 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 nom ...
''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
Saskatchewan Saskatchewan ( ; ) is a province in western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on the south by the U.S. states of Montana and North Dak ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
, that were originally described as ''Lemna'' (''Spirodela'') ''scutata'' by
John William Dawson Sir John William Dawson (1820–1899) was a Canadian geologist and university administrator. Life and work John William Dawson was born on 13 October 1820 in Pictou, Nova Scotia, where he attended and graduated from Pictou Academy. Of Scotti ...
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 epoch (geology), 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 mea ...
of the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
. 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 mechani ...
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 (DOE)
Joint Genome Institute The U.S. Department of Energy (DOE) Joint Genome Institute (JGI), first located in Walnut Creek then Berkeley, California, was created in 1997 to unite the expertise and resources in genome mapping, DNA sequencing, technology development, and i ...
announced that the Community Sequencing Program would fund sequencing of the genome of the giant duckweed, ''
Spirodela polyrhiza ''Spirodela polyrhiza'' ( ''S. polyrrhiza'') is a species of duckweed 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 habitat ...
''. This was a priority project for DOE in 2009. The research was intended to facilitate new biomass 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 was ...
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 change in a broader sense also includes previous long-term changes to E ...
. 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 Bioremediation broadly refers to any process wherein a biological system (typically bacteria, microalgae, fungi, and plants), living or dead, is employed for removing environmental pollutants from air, water, soil, flue gasses, industrial effluent ...
because they grow rapidly, absorbing excess mineral nutrients, particularly
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
and
phosphates 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 ...
. 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 Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small Diptera, flies within the family Culicidae (from the Latin ''culex'' meaning "gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish ...
. 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, Spain File:Lemna minor1.jpg, ''
Lemna minor ''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 ...
'' File:Lemna trisulca0.jpg, '' Lemna trisulca'' File:L gibba3.jpg, ''
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 ...
''
Turning the canals of the Poitevin Marsh (
Marais Poitevin The Marais Poitevin () or Poitevin Marsh is a large area of marshland in western France. The name means "Poitou's Marsh" or the "Marsh of the Poitou region". It is a remnant of what was the former . The western zone near the sea (about two-thi ...
, France) into the "Green Venice": File:Spirodela polyrrhiza marais poitevin.jpg, ''
Spirodela polyrhiza ''Spirodela polyrhiza'' ( ''S. polyrrhiza'') is a species of duckweed 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 habitat ...
'' File:Curve of duckweed covered water edged with several bald cypress trees.JPG, Duckweed-covered water edged with several
bald cypress ''Taxodium distichum'' (bald cypress, swamp cypress; french: cyprès chauve; ''cipre'' in Louisiana) is a deciduous conifer in the family Cupressaceae. It is native to the southeastern United States. Hardy and tough, this tree adapts to a wide ...
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 Phytoremediation technologies use living plants to clean up soil, air and water contaminated with hazardous contaminants. It is defined as "the use of green plants and the associated microorganisms, along with proper soil amendments and agronom ...
* Hyperaccumulators table – 3 *
Neuston Neuston, also known as pleuston, are organisms that live at the surface of the ocean or an estuary, or at the surface of a lake, river or pond. Neuston can live on top of the water surface or may be attached to the underside of the water surface. ...


References


External links

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