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''Lemna'' is a genus of free-floating
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 referred to by the common name "
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 ...
". They are morphologically divergent members of the
arum ''Arum'' is a genus of flowering plants in the family Araceae, native to Europe, northern Africa, and western and central Asia, with the highest species diversity in the Mediterranean region. Frequently called arum lilies, they are not closely ...
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). Also ...
. These rapidly growing plants have found uses as a model system for studies in
community ecology In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological community, ...
, basic plant biology,
ecotoxicology Ecotoxicology is the study of the effects of toxic chemicals on biological organisms, especially at the population, community, ecosystem, and biosphere levels. Ecotoxicology is a multidisciplinary field, which integrates toxicology and ecology. ...
, and production of
biopharmaceutical A biopharmaceutical, also known as a biological medical product, or biologic, is any pharmaceutical drug product manufactured in, extracted from, or semisynthesized from biological sources. Different from totally synthesized pharmaceuticals, t ...
s, and as a source of animal feeds for
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
and
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lot ...
. Currently, 14 species of ''Lemna'' are recognised.


Taxonomy

These duckweeds were previously placed in a separate flowering plant family, the
Lemnaceae 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 ...
, but they are now considered to be members of the
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). Also ...
.


Description

''Lemna'' species grow as simple free-floating
thalli 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 were ...
on or just beneath the water surface. Most are small, not exceeding 5 mm in length, except ''
Lemna trisulca ''Lemna trisulca'' L. (syn. ''Staurogeton trisulcus'' (L.) Schur; star duckweed; ivy-leaved duckweed) is a species of aquatic plants in the arum family Araceae. It has a subcosmopolitan distribution. Unlike other duckweeds, it has submerged ra ...
'', which is elongated and has a branched structure. ''Lemna'' thalli have a single root, which distinguishes this genus from the related
genera 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 nomenclat ...
''
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 ...
'' (lacks roots), ''Spirodela'' and ''Landoltia'' (have multiple roots). The plants grow mainly by
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 ...
: two daughter plants bud off from the adult plant. This form of growth allows very rapid colonisation of new water. Duckweeds are
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, and nearly all of them are known to reproduce sexually,
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 ...
ing and producing
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 ...
under appropriate conditions. Certain duckweeds (such as ''L. gibba'') are long-day plants, while others (such as ''L. minor'') are
short-day plant Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light a ...
s. When ''Lemna'' invades a waterway, it can be removed mechanically, by the addition of herbivorous fish (e.g.
grass carp The grass carp (''Ctenopharyngodon idella'') is a species of large herbivorous freshwater fish in the family Cyprinidae, native to the Pacific Far East, with a native range stretching from northern Vietnam to the Amur River on the Sino-Russian ...
), or, inadvisedly, treated with a
herbicide Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page fo ...
. The rapid growth of duckweeds finds application in
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 ...
of polluted waters, in municipal wastewater treatment and as test organisms for environmental studies. It is also being used as an expression system for economical production of complex
biopharmaceutical A biopharmaceutical, also known as a biological medical product, or biologic, is any pharmaceutical drug product manufactured in, extracted from, or semisynthesized from biological sources. Different from totally synthesized pharmaceuticals, t ...
s. Duckweed meal (dried duckweed) is a good cattle feed. It contains 25–45% protein (depending on the growth conditions), 4.4% fat, and 8–10% fibre, measured by dry weight.


As a bioassay

Organisation for Economic Co-operation and Development The Organisation for Economic Co-operation and Development (OECD; french: Organisation de coopération et de développement économiques, ''OCDE'') is an intergovernmental organization, intergovernmental organisation with 38 member countries ...
and
U.S. Environmental Protection Agency The Environmental Protection Agency (EPA) is an Independent agencies of the United States government, independent executive agency of the United States federal government tasked with environmental protection matters. President Richard Nixon pro ...
(US EPA) guidelines describe toxicity testing using ''L. gibba'' or ''L. minor'' as test organisms. Both of these species have been studied extensively for use in
phytotoxicity Phytotoxicity describes any adverse effects on plant growth, physiology, or metabolism caused by a chemical substance, such as high levels of fertilizers, herbicides, heavy metals, or nanoparticles. General phytotoxic effects include altered plan ...
tests. Genetic variability in responses to toxicants can occur in ''Lemna'', and data are insufficient to recommend a specific clone for testing. The US EPA test uses aseptic technique. The OECD test is not conducted axenically, but steps are taken at stages during the test procedure to keep contamination by other organisms to a minimum. Depending on the objectives of the test and the regulatory requirements, testing may be performed with renewal (semistatic and flow-through) or without renewal (static) of the test solution. Renewal is useful for substances that are rapidly lost from solution as a result of volatilisation, photodegradation, precipitation, or biodegradation.


Production of biopharmaceuticals

''Lemna'' has been transformed by
molecular biologists Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physi ...
to express proteins of
pharmaceutical A medication (also called medicament, medicine, pharmaceutical drug, medicinal drug or simply drug) is a drug used to diagnose, cure, treat, or prevent disease. Drug therapy (pharmacotherapy) is an important part of the medical field and re ...
interest. Expression constructs were engineered to cause ''Lemna'' to secrete the transformed proteins into the growth medium at high yield. Since the ''Lemna'' is grown on a simple medium, this substantially reduces the burden of
protein purification Protein purification is a series of processes intended to isolate one or a few proteins from a complex mixture, usually cells, tissues or whole organisms. Protein purification is vital for the specification of the function, structure and interact ...
in preparing such proteins for medical use, promising substantial reductions in manufacturing costs. In addition, the host ''Lemna'' can be engineered to cause secretion of proteins with human patterns of
glycosylation Glycosylation is the reaction in which a carbohydrate (or ' glycan'), i.e. a glycosyl donor, is attached to a hydroxyl or other functional group of another molecule (a glycosyl acceptor) in order to form a glycoconjugate. In biology (but not al ...
, an improvement over conventional plant gene-expression systems. Several such products are being developed, including
monoclonal antibodies A monoclonal antibody (mAb, more rarely called moAb) is an antibody produced from a cell Lineage made by cloning a unique white blood cell. All subsequent antibodies derived this way trace back to a unique parent cell. Monoclonal antibodies ca ...
.


Duckweed farming

High yields of duckweed with a high protein content for use in
human nutrition Human nutrition deals with the provision of essential nutrients in food that are necessary to support human life and good health. Poor nutrition is a chronic problem often linked to poverty, food security, or a poor understanding of nutritiona ...
,
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
and
fish feed Manufactured feeds are an important part of modern commercial aquaculture, providing the balanced nutrition needed by farmed fish. The feeds, in the form of granules or pellets, provide the nutrition in a stable and concentrated form, enabling th ...
can be achieved by careful control of growth conditions. Although duckweed can tolerate temperatures ranging from to , the optimal growth range is to . The acceptable pH range is 5 to 9, but better growth is obtained in the pH range of 6.5 to 7.5. A minimum water depth of is desirable to prevent excessive temperature swings. High nitrogen levels, for example 20 mM urea, have provided a protein content in the range of 45% by weight. The water may typically contain 60 mg/L of soluble
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 1 mg/L of
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
. Fertiliser is required on a daily basis for optimal growth. Duckweed can be farmed organically, with nutrients being supplied from a variety of sources, for example human urine, cattle manure, pig waste,
biogas plant Biogas is a mixture of gases, primarily consisting of methane, carbon dioxide and hydrogen sulphide, produced from raw materials such as agricultural waste, manure, municipal waste, plant material, sewage, green waste and food waste. It is a r ...
slurry, or other organic matter in slurry form. Because of the rapid growth of duckweed, daily harvesting is necessary to achieve optimal yields. Harvesting is done such that less than 1 kg/m2 of duckweed remains. Under optimal conditions, a duckweed farm can produce 10 to 30 tons of dried duckweed per hectare per year.Scholar search
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Species

Infrageneric classification following Les et al. 2002. ;Section ''Alatae'' *'' Lemna aequinoctialis'' Welw. – lesser duckweed – tropical and subtropical *'' Lemna perpusilla'' Torr. – minute duckweed – eastern United States, Quebec ;Section ''Biformes'' *'' Lemna tenera'' Kurz – Indochina, Sumatra, Northern Territory of Australia ;Section ''Lemna'' *'' Lemna disperma'' Hegelm. *'' Lemna ecuadoriensis'' Landolt *''
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 ...
'' L. – gibbous duckweed – widespread *'' Lemna japonica'' Landolt – Japan, China, Korea, Russian Far East *''
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 ...
'' L. – common duckweed – cosmopolitan *'' Lemna obscura'' (Austin) Daubs – United States, Mexico, Bahamas, Colombia, Ecuador *''
Lemna trisulca ''Lemna trisulca'' L. (syn. ''Staurogeton trisulcus'' (L.) Schur; star duckweed; ivy-leaved duckweed) is a species of aquatic plants in the arum family Araceae. It has a subcosmopolitan distribution. Unlike other duckweeds, it has submerged ra ...
'' L. – ivy duckweed – cosmopolitan *'' Lemna turionifera'' Landolt – temperate Europe, Asia, North America ;Section ''Uninerves'' *''
Lemna minuta ''Lemna minuta'' is a species of duckweed known by the common name least duckweed. It is the smallest ''Lemna'' species. It is native to parts of the Americas, and naturalized in others; the exact native range is not known. It is found on other c ...
'' Kunth – least duckweed – North + South America *'' Lemna valdiviana'' Phil. – Valdivia duckweed – North and South America *'' Lemna yungensis'' Landolt – Bolivia ;Formerly placed here *'' Landoltia punctata'' (G.Mey.) Les & D.J.Crawford (as ''L. oligorrhiza'' Kurz and ''L. punctata'' G.Mey.) *''
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 ...
'' (L.) Schleid. (as ''L. polyrhiza'' L.) *''
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 vascul ...
'' (L.) Horkel ex Wimm. (as ''L. arrhiza'' L.) *''
Wolffia globosa ''Wolffia globosa'' is a species of flowering plant known by the common names Asian watermeal and duckweed. It is native to Asia and is found in parts of the Americas and Africa, where it is an introduced species. It grows in mats on the surface ...
'' (Roxb.) Hartog & Plas (as ''L. globosa'' Roxb.)


References


General readings

*Cross, J.W. (2006)
The Charms of Duckweed
*Landolt, E. (1986) Biosystematic investigations in the family of duckweeds (Lemnaceae). Vol. 2. The family of Lemnaceae – A monographic study. Part 1 of the monograph: Morphology; karyology; ecology; geographic distribution; systematic position; nomenclature; descriptions. Veröff. Geobot. Inst., Stiftung Rübel, ETH, Zurich. Image:LemnaTrisulca.jpg, Ivy Duckweed (''Lemna trisulca'') Image:LemnaGibba.jpg, Gibbous Duckweed (''Lemna gibba'') Image:Common Duckweed (Lemna minor ).jpg, Common Duckweed (''Lemna minor'')


External links



* ttp://titania.sourceoecd.org/vl=5981511/cl=43/nw=1/rpsv/ij/oecdjournals/1607310x/v1n2/s22/p1 OECD Guideline for Lemna Test
USDA Plants Profile: North American SpeciesJepson Manual Treatment: ''Lemna''
{{Taxonbar, from=Q161207 Lemnoideae Araceae genera Aquatic plants