Lemnoideae

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Lemnoideae
Close-up of two different duckweed types:
Spirodela polyrrhiza and Wolffia globosa
: The latter are less than 2 mm long.
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Alismatales
Family: Araceae
Subfamily: Lemnoideae
Genera
Synonyms

Lemnaceae

Lemnoideae is a subfamily of flowering

wetlands. Also known as bayroot, they arose from within the arum or aroid family (Araceae),[1]
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" 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.[2]

Reproduction is mostly by

pistil are produced, by which sexual reproduction occurs. Some view this "flower" as a pseudanthium, or reduced inflorescence, 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

utricle, and a seed
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.[4] 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,[5] 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-

photoautotrophic algae
.

Use as human food crop

Duckweed is consumed in some parts of Southeast Asia, namely Laos, Thailand, and Myanmar.[6] In addition, it is also cultivated as a vegetable in Israel.[7] It produces more protein per square meter than soybeans, so sometimes it is cited as a significant potential food source.[8]

Some initial investigations to what extent duckweed could be introduced in European markets show little consumer objection to the idea.[9] NASA's Caves of Mars Project identified duckweed as a top candidate for growing food on Mars.[10]

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.[11]

slough habitats within the Everglades.[12]

Taxonomy

Duckweeds belong to the order Alismatales and the family Araceae. (a) is a phylogenetic tree based on ribulose-1, 5-bisphosphate carboxylase large-subunit genes. (b) is a schematic ventral view of Spirodela, to show the clonal, vegetative propagation of duckweeds. Daughter fronds (F1) originate from the vegetative node (No), from the mother frond F0 and remain attached to it by the stipule (Sti), which eventually breaks off, thereby releasing a new plant cluster. Daughter fronds may already initiate new fronds (F2) themselves before full maturity. Roots are attached at the prophyllum (P). (c) shows the progressive reduction from a leaf-like body with several veins and unbranched roots to a thallus-like morphology in the Lemnoideae.

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.[13]

Their position within their family has been slightly less clear; however, several twenty-first century studies place them in the position shown below.[13] Although they are in the same family as Pistia, another aquatic plant, they are not closely related.[13]

Araceae

Gymnostachydoideae

Orontioideae (skunk cabbages and golden club)

Lemnoideae (duckweeds)

most of the rest of the family Araceae

The

Landoltia, Lemna, Wolffiella, and Wolffia
.

Duckweed

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).[14] 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.[15]

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.[16] The atpF-atpH non-coding spacer was chosen as a universal DNA barcoding marker for species-level identification of duckweeds.[17]

Fossil record

Extinct free-floating aquatic plants and pollen with affinities to the Lemnoideae first appear in the fossil record during the Late Cretaceous (Maastrichtian) as evidenced by floating leaves described as Aquaephyllum auriculatum from Patagonia, Argentina, and the lemnoid pollen genus Pandaniidites.[18]

Fossils of floating leaves with rootlets from the

anthers that contain Pandaniidites pollen.[20] Occurrences of lemnoid seeds described as Lemnospermum have also been reported.[19]

Research and applications

Research and applications of duckweeds are promoted by two international organizations, The International Lemna Association[21] and the International Steering Committee on Duckweed Research and Applications.[22]

In July 2008, the

U.S. Department of Energy (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 and bioenergy programs.[23] The results were published in February 2014. They provide insight into how this plant is adapted to rapid growth and an aquatic lifestyle.[24]

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

global warming.[27][28] The rapid nature of duckweed has shown that it can double biomass within four and a half days.[29][30][31] Duckweed removes carbon dioxide from the atmosphere, and it may have value for climate change mitigation.[32]

Filtration of contaminants and nutrients

The plants can provide

phosphates. For these reasons, they are touted as water purifiers of untapped value.[33]

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

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.[35][36][37]

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.[38]

See also

References

  1. PMID 21653404
    .
  2. .
  3. ^ Landolt, Elias (1986). Biosystematic investigations in the family of duckweeds (Lemnaceae) Vol. 2: The family of Lemnaceae: a monographic study. – Morphology, karyology, ecology, geographic distribution, nomenclature, descriptions. Zürich: Eidgenössische Technische Hochschule Zürich.
  4. . Retrieved 7 May 2012.
  5. ^ Hutchinson, G. Evelyn (1975). A Treatise on Limnology. Vol. 3: Limnological botany. New York, NY: John Wiley & Sons.
  6. S2CID 6617534
    .
  7. ^ "Lemna Gibbous Duckweed, Swollen duckweed, Windbags, Duckweed PFAF Plant Database". pfaf.org. Retrieved 4 January 2023.
  8. ^ Landesman, Louis. "Dr. Wastewater's duckweed application page". Archived from the original on 27 October 2009. Retrieved 31 January 2012.
  9. .
  10. ^ "Flat crops for Mars". The caves of Mars (highmars.org). Archived from the original on 1 July 2007. Retrieved 11 January 2022.
  11. .
  12. . Retrieved 1 January 2021.
  13. ^ a b c Cabrera, Lidia I.; Salazar, Gerardo A.; Chase, Mark W.; Mayo, Simon J.; Bogner, Josef; Dávila, Patricia (2008). "Phylogenetic relationships of aroids and duckweeds (Araceae) inferred from coding and noncoding plastid DNA".
    PMID 21632433
    .
  14. .
  15. .
  16. . Retrieved 2 August 2012.
  17. .
  18. .
  19. ^ .
  20. .
  21. ^ "The International Lemna Association (ILA)" (official website).
  22. ^ "International Steering Committee on Duckweed Research and Applications (ISCDRA)" (official website).
  23. ^ "Duckweed genome sequencing has global implications". E! Science News (Esciencenews.com). 8 July 2008. Retrieved 13 November 2011.
  24. PMID 24548928
    .
  25. ^ Michael, Todd P. (2008). "Genome sequencing of the duckweed Spirodela polyrhiza: A biofuels, bioremediation, and carbon cycling crop" (PDF). Rutgers University. Archived from the original (PDF) on 3 October 2011. Retrieved 7 May 2012.
  26. ^ "Researchers Find Fuel in Odd Places". Ncsu.edu. Retrieved 13 November 2011.
  27. ^ Sims, Bryan (n.d.). "Duckweed quacks volumes of potential". Biomassmagazine.com. Retrieved 13 November 2011.
  28. ^ "Duckweed a possible solution to energy needs, researchers say". Pressofatlanticcity.com. 3 May 2010. Retrieved 13 November 2011.
  29. PMID 33424309
    .
  30. .
  31. .
  32. ^ "Carbon Neutral Energy". American Energy Independence. Retrieved 13 November 2011.
  33. ^ "Duckweed wastewater treatment and reuse for fodder (West Bank)". Idrc.ca. Archived from the original on 8 June 2011. Retrieved 13 November 2011.
  34. ^ Iqbal, Sascha (March 1999). "Duckweed aquaculture: Potentials, possibilities, and limitations for combined wastewater treatment and animal feed production in developing countries" (PDF). SANDEC Report. 6 (99). Retrieved 31 January 2012.
  35. ^ "Duckweed genome sequencing has global implications. Pond scum can undo pollution, fight global warming and alleviate world hunger" (Press release). Rutgers University. 8 July 2008. Retrieved 10 March 2022.
  36. ^ Cross, John W. "Practical duckweed: Application areas and sponsors". Mobot.org. Retrieved 13 November 2011.
  37. ^ Knibb, Wayne (July 2001 – June 2004). "Bioremediation of aquaculture waste and degraded waterways using finfish". Department of Primary Industries & Fisheries Tools. Australia: Queensland Government. Archived from the original on 20 October 2007. Retrieved 6 February 2012.
  38. ^ "3 Eco-Innovators Tackling Pollution and Climate Change". Columbia Magazine. Retrieved 24 November 2021.

External links

derived from
Watson, L. & Dallwitz, M.J. (3 May 2006) [1992]. The Families of Flowering Plants: Descriptions, illustrations, identification, information retrieval. Archived from the original on 3 January 2007 – via delta-intkey.com.