{{Short description|Genus of filamentous green algae}} {{About||the other genus of algae called blanket weed|Spirogyra}} {{Automatic taxobox | image = Cladophora.JPG |image_alt=Cladophora |image_caption=''Cladophora'' | taxon = Cladophora | authority = [[Friedrich Traugott Kützing|Kütz.]], 1843 | type_species = Cladophora oligoclona | type_species_authority = (Kützing) Kützing<ref name=AlgaeBase/> | subdivision_ranks = Species | subdivision = *''[[Cladophora albida]]'' *''[[Cladophora aokii]]'' *''[[Cladophora brasiliana]]'' *''[[Cladophora catenata]]'' *''[[Cladophora columbiana]]'' *''[[Cladophora crispata]]'' *''[[Cladophora dalmatica]]'' *''[[Cladophora fracta]]'' *''[[Cladophora glomerata]]'' *''[[Cladophora graminea]]'' *''[[Cladophora laetevirens]]'' *''[[Cladophora montagneana]]'' *''[[Cladophora prolifera]]'' *''[[Cladophora rivularis]]'' *''[[Cladophora scopaeformis]]'' *''[[Cladophora sericea]]'' *''[[Cladophora socialis]]'' *''[[Cladophora vagabunda]]'' }} [[File:Cladophora glomerata (L.) Kutz. (AM AK310111-1).jpg|thumb|{{center|''Cladophora glomerata'', showing branching filaments and cellular structures}}]]'''''Cladophora''''' is a genus of branching filamentous green algae in the class [[Ulvophyceae]]. They may be referred to as '''reticulated algae''', '''branching algae''',<ref>{{cite web |title=Cladophora sp. Algae |url=https://www.aquasabi.com/aquascaping-wiki_algae_cladophora-sp-algae |website=Aquasabi |access-date=28 August 2024 |language=en}}</ref> or '''blanket weed'''.<ref name=ga>{{cite web |title=The Ultimate Algae Guide |url=https://greenaqua.hu/en/alga-tajekoztato |website=Green Aqua |access-date=28 August 2024 |language=en}}</ref> The genus has a worldwide distribution, mainly found in marine or brackish waters; a few species are found in [[freshwater]].<ref name=Skaloud/> It is harvested for use as a food and medicine.
==Description and appearance== ''Cladophora'' forms thalli consisting of branched, uniseriate (one cell thick) filaments which are usually sparsely to profusely branched (rarely without any branching). The thallus may be attached to a substrate via [[rhizoid]] cells produced by the basal cell, or by a simple disc-like holdfast; alternatively it may be unattached. Filaments are wider at the base and narrower near the tips. Cells produce a branch directly below the cross-wall, or may be pseudo-dichotomously branched. Cells are cylindrical, barrel-shaped or club-shaped.<ref name=Skaloud>{{cite book | last1 = Škaloud | first1 = Pavel | last2 = Rindi | first2 = Fabio | last3 = Boedeker | first3 = Christian | last4 = Leliaert | first4 = Frederik | title = Chlorophyta: Ulvophyceae | series = Süßwasserflora von Mitteleuropa | volume = 13 | publisher = Springer Spektrum | location = Berlin, Germany | year = 2018 | isbn = 978-3-662-55494-4 | doi = 10.1007/978-3-662-55495-1 | url = https://link.springer.com/book/10.1007/978-3-662-55495-1 | pages=i-x, 1–289 }}</ref>
The cells within ''Cladophora'' are large and multinucleate (with multiple [[cell nucleus|nuclei]]) and have many parietal round [[chloroplast]]s each with a single [[pyrenoid]], which usually join into a net-like reticular formation.<ref name=Skaloud/> ''Cladophora'' bright grass-green in color due to [[chlorophyll a]], and [[chlorophyll b]], in ratios similar to those of higher plant; it also contains [[Β-Carotene|β-carotene]] and [[xanthophyll]]s.<ref name=Michalak>Michalak, I., Messyasz, B. Concise review of Cladophora spp.: macroalgae of commercial interest. J Appl Phycol 33, 133–166 (2021). https://doi.org/10.1007/s10811-020-02211-3</ref> Thick-walled swollen cells, called [[akinete]]s, develop under unfavorable conditions.<ref name=Skaloud/> times of short [[Photoperiodism|photoperiods]], low temperatures or nutrient unavailability. The cell walls of ''Cladophora'' species often contain [[cellulose]] and [[pectin]], providing strength and flexibility.<ref name=Michalak/>
The filaments can be quite long and may form dense mats or tufts in aquatic environments. When ''Cladophora'' becomes detached from the rocky substrate that it grows upon, it can then build up on shorelines, making their reticular formations, (dense mats) visible to onlookers. When dense mats form, this can affect light penetration and oxygen levels in aquatic ecosystems. Mats may also serve as habitat and food for various aquatic organisms.<ref name=Michalak/>
Temperature, water currents and waves affect their metabolism and morphology, and branching patterns. At 15–20 °C branches appear alternate, they can also appear completely absent in temperatures below 25 °C.<ref name=Michalak/>
==Life cycle== {{Unreferenced section|date=August 2024}}
=== Asexual reproduction === This plant is annual and can reproduce through both sexual and asexual mechanisms. Some species have isogametic capacities but little is known about this and it seems to be a rare occurrence present in few species. The few cases that have occurred were in apical and sub apical un-specialized cells. Cladophora generally reproduces asexually using 2 flagellated [[zoospore]]s using mitotic division and distal cells of side branches.
=== Sexual reproduction === This occurs when the male and female gametes fuse, usually in water. The fusion of gametes results in the formation of a diploid zygote. The diploid zygote undergoes mitotic divisions to form a diploid sporophyte. The sporophyte phase is often short-lived and less conspicuous. It produces haploid spores through meiosis. Haploid spores are released from the sporophyte and can be dispersed by water currents or other means. Haploid spores germinate to form new haploid gametophytes, completing the life cycle. Reproductive cells, often referred to as zoospores, are typically motile and possess flagella, allowing them to move in water. Unlike ''[[Spirogyra]]'' the filaments of ''Cladophora'' branch and do not undergo conjugation.
There are two multicellular stages in its life cycle – a [[ploidy|haploid]] [[gametophyte]] and a [[ploidy|diploid]] [[sporophyte]] – which look highly similar. The only way to tell the two stages apart is to either count their [[chromosome]]s, or examine their offspring. The haploid gametophyte produces haploid gametes by [[mitosis]] and the diploid sporophyte produces haploid spores by [[meiosis]]. The only visible difference between the gametes and spores of ''Cladophora'' is that the gametes have two [[flagellum|flagella]] and the spores have four.
==Habitat and distribution==
''Cladophora'' exist in multiple ecosystems which include both tropical and moderate climates, freshwater, wastewater, and marine water ecosystems, and in multiple biomes which include lakes, ponds, dam reservoirs, large rivers and the coastal [[Littoral zone|littoral]] zones as well as in areas of oceanic depth (which determines temperatures and light availability as well as oxygen availability, which can become a limiting factor). This taxon needs [[nitrate]]s, and orthophosphates, as well as hard water conditions with pH levels between 7–10, high light intensity and nutrient densities that are rich in nature. ''Cladophora'' may attach to submerged surfaces like rocks or other aquatic substrates, enabling it to grow more rapidly through optimization of space availability, (another limiting factors and source of infraspecific, intraspecies and interspecies competition).<ref>{{Cite journal |last=Zhang |first=Juan |last2=Yao |first2=Xiaojun |last3=Duan |first3=Hongyu |last4=Chu |first4=Xinde |last5=Yang |first5=Chen |last6=Hu |first6=Jiayu |date=2024 |title=Extraction of ''Cladophora'' Blooms in Qinghai Lake Through the Integration of Sentinel-2 MSI Imagery and Deep Learning Techniques |url=https://doi.org/10.1109/jstars.2024.3447886 |journal=IEEE Journal of Selected Topics in Applied Earth Observations and Remote Sensing |volume=17 |pages=19115–19129 |doi=10.1109/jstars.2024.3447886 |issn=1939-1404|doi-access=free }}</ref>
In Europe, only 15 known subtypes of ''Cladophora'' are freshwater. ''Cladophora'' exists in Africa, Asia, the Pacific Islands, Australia, Europe, the Atlantic Islands, North America, New Zealand, as well as the Caribbean. In some Asian countries, almost 5% of human diets are algae, and its consumption is popular in France, Thailand, as well as the Hawaiian Islands.<ref name=Michalak/>
''Cladophora'' balls are formed from the filaments of the algae<ref>Burrows, E.M.1991. ''Seaweeds of the British Isles Volume 2 Chlorophyta.'' Natural History Museum, London. {{ISBN|0-565-00981-8}}</ref> which are photosynthetic. Large numbers of these balls were thrown ashore in Devon, England. They had an average diameter of 2.5 cm and several million balls were found forming a layer.<ref>Bryant, J. and Irvine, Linda. 2016. Marimo, ''Cladophora'', ''Posidonia'' and Other Plant Balls. ''The Linnean.'' '''32''' (2) pp.11–14</ref>
==Ecological impact == The genus ''Cladophora'' is [[Cosmopolitan distribution|cosmopolitan]] and accordingly infestations cannot often be regarded logically as being invasive. Where they occur they may at various times be seen as beneficial, as a nuisance, or an outright pest.{{citation needed|date=May 2023}}
Modest growth of ''Cladophora'' is generally harmless; the growth is an important food for many fish and other aquatic animals, as a buffer for the sequestration of nutrients in the water body and for protection of some aquatic organisms from solar [[ultraviolet]] radiation.{{citation needed|date=May 2023}} ''Cladophora'' species can be an environmental nuisance, however, causing major alterations to [[Benthic zone|benthic]] conditions linked particularly with increased [[phosphorus]] loading.<ref name=Michalak/>
Where ''Cladophora'' becomes a pest is generally where special circumstances cause such drastic overgrowth that [[algal bloom]]s develop and form floating mats. Typical examples include where [[hypertrophication]] or high mortality of rival organisms produce high concentrations of dissolved phosphorus. Extensive floating mats prevent circulation that is necessary for the aeration of deeper water and, by blocking the light, they kill photosynthesising organisms growing beneath. The mats interfere with the fishing industry by clogging nets and preventing the use of lines. Where they wash ashore the masses of rotting material reduce shoreline property values along water bodies such as the [[Great Lakes]] in the United States.<ref name="FFBHE">{{Cite web |title=Great Lakes Science Center |url=https://www.glsc.usgs.gov/environmental-health/environmental-health-beach-management-decisions/beach-monitoring-human-health}}</ref>
[[Quagga mussel]] populations have increased tremendously during the same time frame as the blooming of ''Cladophora'', though their ecological relationships are not yet clear and may be complex.<ref>{{cite web |date=June 29, 2008 |title=The beach speaks for itself |url=http://www.jsonline.com/news/wisconsin/29561884.html |archive-url=https://web.archive.org/web/20160303193630/http://www.jsonline.com/news/wisconsin/29561884.html |archive-date=3 March 2016}}</ref>
==Phytoremediation and bioindication==
Some ''Cladophora'' species, including ''Cladophora fascicularis'', have been investigated for their use in [[phytoremediation]]. ''Cladophora'' ''fascicularis'' may contribute to improving water quality in contaminated aquatic ecosystems. This algae is [[biologically active]]. ''Cladophora'' possesses unprecedented capacities to filter toxins from water such as [[heavy metals]] and pollutants and efficiently absorb nutrients, including [[nitrogen]] and [[phosphorus]].<ref name=Michalak/>
Cladophora is known for its blooming activities which are seasonal, (early spring to summer) and indicate their [[Eutrophication|eutrophic]] abilities as well as that pollution may be present in the environment that they are growing within.<ref name=Michalak/>
==Uses==
=== Food === [[Image:Kháy phen appetizer.jpg|thumb|Kháy phen served as an appetizer dish]]''Cladophora'' can be eaten as a food item either dried or fresh, however it loses some of its nutritional value with increased processing and with genetic modification. It is considered a carbohydrate when consumed.<ref name=Michalak/>
In [[Laos]], ''Cladophora'' are commonly eaten as a delicacy and usually known in English under the name "Mekong weed". The algae grow on underwater rocks and thrive in clear spots of water in the [[Mekong]] river basin. They are harvested 1 to 5 months a year and most often eaten in dry sheets similar to Japanese [[nori]], though much cruder in their format. [[Luang Prabang]]'s speciality is dry khai with sesame ([[kaipen]]), while [[Vang Vieng]] is famous for its roasted kháy sheets. They can be eaten in strips as an [[Hors d'oeuvre|appetizer]], with a meal or as a snack with Beer Lao. [[Kaipen]] are the most readily available form of Mekong weed and are famous throughout the country and in the neighbouring [[Isan|Isaan]], though difficult to find beyond [[Vientiane]]. Mekong weed can also be eaten raw, in soups, or cooked in [[steamed curry|steamed curries]].{{citation needed|date=May 2023}}
=== Biofuel === Trans-esterification of ''Cladophora'' can be used to transform it into biodiesel. Alcohols and enzymes and ''Cladophora'' oils are used to form this reaction, but production is often highly dependent on oil content.<ref name=Michalak/>
=== Pharmacology === ''Cladophora'' has pharmacological uses as medication. ''Cladophora'' is purported to have antimicrobial, [[antihistamine]], antiviral, antioxidant, anti-sclerosis, anti-inflammatory and weight loss properties as well as [[secondary metabolite]]s and [[Bioactive compound|bioactive]] components that have uses in diabetes, hypertension, cancer, and serves as a protection from parasites. However, as of December 2023, ''Cladophora'' has yet to be utilized commercially as a pharmacological agent.<ref name=Michalak/>
=== Other uses === There is an additional demand for ''Cladophora'' for [[biomass]] productivity with utilization commercially as material for value added products. It can be used for cosmetic application and used as a fertilizer.<ref name=Michalak/> [[Marimo]] used to be classified in ''Cladophora'', though is now called ''Aegagropila linnaei''. Algae balls are significant in the hobby aquarium trade.{{citation needed|date=May 2023}}
== Taxonomy and selected species == As of 2025, [[AlgaeBase]] accepts 190 species within the genus ''Cladophora''.<ref name=AlgaeBase>{{AlgaeBase genus | id=37 | title=''Cladophora'' Kützing, 1843, nom. cons. | access-date=2025-07-10 }}</ref>
''Cladophora'' has had a long and complicated taxonomic history. It was first described by [[Friedrich Traugott Kützing]] in 1843, who described many new species and varieties based on minor differences in morphology; subsequent authors did the same, resulting in hundreds of species names.<ref name=Skaloud/> However ''Cladophora'' displays extensive [[phenoplastic plasticity]] in response to different environmental conditions;<ref>{{cite journal|last1=Gestinari |first1=L. M. D. |last2=Pereira |first2=S. M. B. |last3=Yoneshigue-Valentin |first3=Y. |title=Distribution of Cladophora Species (Cladophorales, Chlorophyta) along the Brazilian Coast |journal=Phytotaxa |date=2010 |volume=14 |page=22 |doi=10.11646/phytotaxa.14.1.2}}</ref> the same plant may have a drastically different appearance in fresh or salt water. In the 20th century, [[Christian van den Hoek]] and other authors [[synonymized]] many names, gradually reducing the number of accepted species. On the other hand, molecular data has shown that ''Cladophora'' contains [[cryptic diversity]]. Therefore, it is difficult to know the true number of ''Cladophora'' species.<ref name=Skaloud/>
The simple morphology of ''Cladophora'' has independently evolved multiple times in the order Cladophorales, making it [[polyphyletic]]. Taxonomic studies including [[molecular phylogenetic]]s have begun to split ''Cladophora'' into smaller [[monophyletic]] genera, but the resulting genera often overlap significantly in terms of morphology.<ref name=Skaloud/>
Some species include: {{div col}} *''[[Cladophora albida]]'' *''[[Cladophora aokii]]'' *''[[Cladophora brasiliana]]'' *''[[Cladophora catenata]]'' *''[[Cladophora columbiana]]'' *''[[Cladophora crispata]]'' *''[[Cladophora dalmatica]]'' *''[[Cladophora fracta]]'' *''[[Cladophora glomerata]]'' *''[[Cladophora graminea]]'' *''[[Cladophora graminea]]'' *''[[Cladophora montagneana]]'' *''[[Cladophora prolifera]]'' *''[[Cladophora rivularis]]'' *''[[Cladophora scopaeformis]]'' *''[[Cladophora sericea]]'' *''[[Cladophora socialis]]'' *''[[Cladophora vagabunda]]'' {{div col end}}
Related genera with species formerly included in ''Cladophora'' are: * ''[[Pseudocladophora]]''<ref name=Brito>{{cite journal|last1=Brito |first1=Jhullyrson Osman Ferreira de |last2=Alves |first2=Aigara Miranda |last3=Portela |first3=Luane do Carmo |last4=Van Den Berg |first4=Cassio |last5=Pereira |first5=Sonia Maria Barreto |last6=Gestinari |first6=Lísia Monica |last7=Cassano |first7=Valéria |last8=Moura |first8=Carlos Wallace do Nascimento |last9=Gama |first9=Watson Arantes |title=Phylogenetic analysis of the ''Cladophora coelothrix'' complex, including the description of the new genus ''Leliaertia'' (Cladophorales, Ulvophyceae) |journal=Phycologia |date=2024 |volume=63 |pages=11–23 |doi=10.1080/00318884.2023.2282921}}</ref> * ''[[Aegagropila]]'' (''[[Marimo|Aegagropila linnaei]]'' (marimo) was formerly placed here as ''Cladophora aegagropila'') * ''[[Aegagropilopsis]]''<ref>{{cite journal|last1=Boedeker |first1=Christian |last2=O'Kelly |first2=Charles J. |last3=Star |first3=Wim |last4=Leliaert |first4=Frederik |title=Molecular phylogeny and taxonomy of the ''Aegagropila'' clade (Cladophorales, Ulvophyceae), including the description of ''Aegagropilopsis'' gen. nov. and ''Pseudocladophora'' gen. nov. |journal=Journal of Phycology |date=2012 |volume=48 |issue=3 |pages=808–825 |doi=10.1111/j.1529-8817.2012.01145.x |pmid=27011097 }}</ref> * ''[[Leliaertia]]''<ref name=Brito/> * ''[[Lychaete]]''<ref name=Brito/> * ''[[Vandenhoekia]]''<ref name=Boedeker2023/> * ''[[Willeella]]''<ref name=Boedeker2016>{{cite journal|last1=Boedeker |first1=Christian |last2=Leliaert |first2=Frederik |last3=Zuccarello |first3=Giuseppe C. |title=Molecular phylogeny of the Cladophoraceae (Cladophorales, Ulvophyceae), with the resurrection of ''Acrocladus'' Nägeli and ''Willeella'' Børgesen, and the description of ''Lurbica'' gen. nov. and ''Pseudorhizoclonium'' gen. nov. |journal=Journal of Phycology |date=2016 |volume=52 |issue=6 |pages=905–928 |doi=10.1111/jpy.12457 |pmid=27535014 }}</ref> * ''[[Lurbica]]''<ref name=Boedeker2016/> * ''[[Rama (alga)|Rama]]''<ref name=Boedeker2023>{{cite journal|last1=Boedeker |first1=Christian |last2=Wynne |first2=Michael J. |last3=Zuccarello |first3=Giuseppe C. |title=Hidden diversity in high-latitude Southern Hemisphere environments: Reinstatement of the genus ''Rama'' and description of ''Vandenhoekia'' gen. nov. (Cladophoraceae, Ulvophyceae, Chlorophyta), two highly variable genera |journal=Journal of Phycology |date=2023 |volume=59 |issue=6 |pages=1284–1298 |doi=10.1111/jpy.13394 |pmid=37795849 |doi-access=free }}</ref>
==References== {{Reflist}}
==Further reading== *{{cite book |last1=Garrison |first1=Paul J. |last2=Greb |first2=Stephen R. |last3=LaLiberte |first3=Gina |title=Western Lake Michigan Nearshore Survey Of Water Chemistry And ''Cladophora'' Distribution, 2004-2007 |date=May 2008 |publisher=Wisconsin Department of Natural Resources, Bureau of Science Services |series=PUB-SS-1038 2008 |url=https://dnr.wisconsin.gov/sites/default/files/topic/GreatLakes/CladophoraReport2008.pdf}} *Marsin, P. and J. Tomasz. (2005). Introductory studies on the morphology of the genus ''Cladophora'' from the Gulf of Gdańsk. ''Ocean. Hydrob. Studies'', 34 (Supl.3): 187–193
==External links== *[http://www.algaebase.org/search/pictures/?currentMethod=imgs&fromSearch=yes&displayCount=20&query=Cladophora Images] {{Webarchive|url=https://web.archive.org/web/20170224120711/http://www.algaebase.org/search/pictures/?currentMethod=imgs&fromSearch=yes&displayCount=20&query=Cladophora |date=2017-02-24 }} of ''Cladophora'' at [[AlgaeBase]] *{{cite web |url=http://www.algaebase.org/generadetail.lasso?genus_id=37 |title=''Cladophora'' Kützing 1843: 262 |work=[[AlgaeBase]] |author=T. J. Entwisle|author1-link=Timothy John Entwisle|author2=M. D. Guiry |date=2005-11-12}} *[https://web.archive.org/web/20110516162453/http://www.mbari.org/staff/conn/botany/greens/angie/index.htm ''Cladophora'' Index.] Monterey Bay Aquarium
{{Taxonbar|from=Q311264}} {{Authority control}}
[[Category:Cladophoraceae]] [[Category:Cladophorales genera]] [[Category:Taxa named by Friedrich Traugott Kützing]] [[Category:Freshwater algae]]