Lemna

Lemna

Common duckweed (Lemna minor)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Monocots
Order:	Alismatales
Family:	Araceae
Subfamily:	Lemnoideae
Genus:	Lemna L.
Synonyms^[1]^[2]
Staurogeton Rchb. Lenticularia Ség. Lenticula P.Micheli ex Adans. Hydrophace Hallier Telmatophace Schleid. Thelmatophace Godr. Lenticularia P.Micheli ex Montandon

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 community ecology, basic plant biology, ecotoxicology, and production of biopharmaceuticals, and as a source of animal feeds for agriculture and aquaculture. Currently, 14 species of Lemna are recognised.^[3]

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Taxonomy

These duckweeds were previously placed in a separate flowering plant family, the Lemnaceae, but they are now considered to be members of the Araceae.^[4]

Description

Lemna species grow as simple free-floating thalli on or just beneath the water surface. Most are small, not exceeding 5 mm in length, except Lemna trisulca, which is elongated and has a branched structure. Lemna thalli have a single root, which distinguishes this genus from the related genera Wolffia (lacks roots), Spirodela and Landoltia (have multiple roots).

The plants grow mainly by vegetative reproduction: two daughter plants bud off from the adult plant. This form of growth allows very rapid colonisation of new water. Duckweeds are flowering plants, and nearly all of them are known to reproduce sexually, flowering and producing seed under appropriate conditions. Certain duckweeds (such as L. gibba) are long-day plants, while others (such as L. minor) are short-day plants.

When Lemna invades a waterway, it can be removed mechanically, by the addition of herbivorous fish (e.g. grass carp), or, inadvisedly, treated with a herbicide.

The rapid growth of duckweeds finds application in bioremediation of polluted waters, in municipal wastewater treatment ^[5] and as test organisms for environmental studies.^[6] It is also being used as an expression system for economical production of complex biopharmaceuticals.

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^[7] and U.S. Environmental Protection Agency (US EPA)^[8] 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 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 to express proteins of pharmaceutical 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 in preparing such proteins for medical use, promising substantial reductions in manufacturing costs.^[9]^[10] In addition, the host Lemna can be engineered to cause secretion of proteins with human patterns of glycosylation, an improvement over conventional plant gene-expression systems.^[11] Several such products are being developed, including monoclonal antibodies.

Duckweed farming

High yields of duckweed with a high protein content for use in human nutrition, animal and fish feed can be achieved by careful control of growth conditions. Although duckweed can tolerate temperatures ranging from 6 °C (43 °F) to 33 °C (91 °F), the optimal growth range is 20 °C (68 °F) to 28 °C (82 °F). 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 1 foot (30 cm) 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 dry weight. The water may typically contain 60 mg/L of soluble nitrogen and 1 mg/L of phosphorus. 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,^[12] cattle manure, pig waste, biogas plant 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.^[13]

Species

Infrageneric classification following Les et al. 2002.^[14]

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 L. – gibbous duckweed – widespread
Lemna japonica Landolt – Japan, China, Korea, Russian Far East
Lemna minor L. – common duckweed – cosmopolitan
Lemna obscura (Austin) Daubs – United States, Mexico, Bahamas, Colombia, Ecuador
Lemna trisulca L. – ivy duckweed – cosmopolitan
Lemna turionifera Landolt – temperate Europe, Asia, North America

Section Uninerves

Lemna minuta 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 (L.) Schleid. (as L. polyrhiza L.)
Wolffia arrhiza (L.) Horkel ex Wimm. (as L. arrhiza L.)
Wolffia globosa (Roxb.) Hartog & Plas (as L. globosa Roxb.)^[15]

References

^ Kew World Checklist of Selected Plant Families
^ "Genus: Lemna L." Germplasm Resources Information Network. United States Department of Agriculture. 2006-11-03. Archived from the original on 2015-09-24. Retrieved 2013-04-13.
^ "Search results — the Plant List".
^ Bremer, B.; Bremer, K.; Chase, M.W.; Reveal, J.L.; Soltis, D.E.; Soltis, P.S.; Stevens, P.F. (2003). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II". Botanical Journal of the Linnean Society. 141 (4): 399–436. doi:10.1046/j.1095-8339.2003.t01-1-00158.x.
^ Iatrou, Evangelia I.; Kora, Elianta; Stasinakis, Athanasios S. (2019). "Investigation of biomass production, crude protein and starch content in laboratory wastewater treatment systems planted with Lemna minor and Lemna gibba". Environmental Technology. 40 (20): 2649–2656. Bibcode:2019EnvTe..40.2649I. doi:10.1080/09593330.2018.1448002. PMID 29502496. S2CID 3805774.
^ Gatidou et al., 2015. Assessing single and joint toxicity of three phenylurea herbicides using Lemna minor and Vibrio fischeri bioassays. Chemosphere vol. 119, January 2015, p. 569-574. Gatidou et al., 2015
^ SourceOECD: issues
^ "Archived copy" (PDF). Archived from the original (PDF) on 2008-12-17. Retrieved 2007-03-09.{{cite web}}: CS1 maint: archived copy as title (link)
^ "Biolex Corporate Website".
^ Gasdaska, JR; Spencer D; Dickey L (Mar–Apr 2003). "Advantages of Therapeutic Protein Production in the Aquatic Plant Lemna". BioProcessing Journal. 2 (2): 49–56. doi:10.12665/J22.Gasdaska.
^ Cox, KM; Sterling JD; Regan JT; Gasdaska JR; Frantz KK; Peele CG; Black A; Passmore D; Moldovan-Loomis C; Srinivasan M; Cuison S; Cardarelli PM; Dickey LF (December 2006). "Glycan Optimization of a Human Monoclonal Antibody in the Aquatic Plant Lemna Minor". Nature Biotechnology. 24 (12): 1591–1597. doi:10.1038/nbt1260. PMID 17128273. S2CID 1840557.
^ "From urine to protein".
^ Leng, R A; J H Stambolie; R Bell (October 1995). "Duckweed – a potential high-protein feed resource for domestic animals and fish". Livestock Research for Rural Development. 7 (1). Archived from the original on March 24, 2005.^{Scholar search}
^ Les, Donald H.; Crawford, Daniel J.; Landolt, Elias; Gabel, John D.; Kimball, Rebecca T. (2002). "Phylogeny and Systematics of Lemnaceae, the Duckweed Family" (PDF). Systematic Botany. 27 (2): 221–240. Retrieved 2020-04-17.
^ "GRIN Species Records of Lemna". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2020-04-17.

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.