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''Cladophora'' is a genus of reticulated filamentous green algae in the class
Ulvophyceae The Ulvophyceae or ulvophytes are a class of green algae, distinguished mainly on the basis of Ultrastructure, ultrastructural morphology, Biological life cycle, life cycle and molecular phylogenetics, molecular phylogenetic data. The sea lettu ...
. They may be referred to as reticulated algae, branching algae, or blanket weed. The genus has a worldwide distribution and is harvested for use as a food and medicine.


Description and appearance

''Cladophora'' coloring is bright green which reflects the chlorophyll a, and chlorophyll b, which are similar to higher plant ratios and that also contains β-carotene and xanthophylls.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 The thallus branches are smaller than the main axis, dichotomous, rough in texture, and have narrow tips. 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. ''Cladophora'' form a branched filamentous chlorophyte structure with large cylindrical cells forming long, regularly branched growths. 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.


Cellular structure

The cells within ''Cladophora'' have multiple pyrenoids, are large and multiple nucleated and have many parietal round
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
s, which usually join into a net-like reticular formation. Thick walls called akinetes are made out of vegetative cells in times of short photoperiods, low temperatures or nutrient unavailability. The cell walls of ''Cladophora'' species often contain
cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
and pectin, providing strength and flexibility. Thick walls and sheaths are mucilaginous.


Life cycle


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 A zoospore is a motile asexual spore that uses a flagellum for locomotion in aqueous or moist environments. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Certain zoospores are ...
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 ''Spirogyra'' (common names include water silk, mermaid's tresses, and blanket weed) is a genus of filamentous charophyte green algae of the order Zygnematales, named for the helical or spiral arrangement of the chloroplasts that is charact ...
'' the filaments of ''Cladophora'' branch and do not undergo conjugation. There are two multicellular stages in its life cycle – a
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
and a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
sporophyte A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
– which look highly similar. The only way to tell the two stages apart is to either count their
chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
s, or examine their offspring. The haploid gametophyte produces haploid gametes by
mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...
and the diploid sporophyte produces haploid spores by
meiosis Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
. The only visible difference between the gametes and spores of ''Cladophora'' is that the gametes have two
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
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 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 Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
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). 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. ''Cladophora'' balls are formed from the filaments of the algae 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.


Ecological impact

The genus ''Cladophora'' is 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. 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 Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
radiation. ''Cladophora'' species can be an environmental nuisance, however, causing major alterations to
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
conditions linked particularly with increased
phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
loading. Where ''Cladophora'' becomes a pest is generally where special circumstances cause such drastic overgrowth that algal blooms 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 The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
in the United States. 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.


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 Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
and
phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
. Cladophora is known for its blooming activities which are seasonal, (early spring to summer) and indicate their eutrophic abilities as well as that pollution may be present in the environment that they are growing within.


Uses


Food

''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. In
Laos Laos, officially the Lao People's Democratic Republic (LPDR), is the only landlocked country in Southeast Asia. It is bordered by Myanmar and China to the northwest, Vietnam to the east, Cambodia to the southeast, and Thailand to the west and ...
, ''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 Luang Prabang (Lao language, Lao: wikt:ຫຼວງພະບາງ, ຫຼວງພະບາງ, pronounced ), historically known as Xieng Thong (ຊຽງທອງ) and alternatively spelled Luang Phabang or Louangphabang, is the capital of Lu ...
'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 appetizer, with a meal or as a snack with Beer Lao. Luang Prabang kháy sheets kaipen are the most readily available form of Mekong weed and are famous throughout the country and in the neighbouring Isaan, though difficult to find beyond
Vientiane Vientiane (, ) is the capital city, capital and largest city of Laos. Situated on the banks of the Mekong, Mekong River at the Thailand, Thai border, it comprises the five urban districts of Vientiane Prefecture and had a population of 840,000 ...
. Mekong weed can also be eaten raw, in soups, or cooked in steamed curries.


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.


Pharmacology

''Cladophora'' has pharmacological uses as medication. ''Cladophora'' is purported to have antimicrobial,
antihistamine Antihistamines are drugs which treat allergic rhinitis, common cold, influenza, and other allergies. Typically, people take antihistamines as an inexpensive, generic (not patented) drug that can be bought without a prescription and provides ...
, antiviral, antioxidant, anti-sclerosis, anti-inflammatory and weight loss properties as well as
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s and 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.


Other uses

There is an additional demand for ''Cladophora'' for
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
productivity with utilization commercially as material for value added products. It can be used for cosmetic application and used as a fertilizer. Marimo used to be classified in ''Cladophora'', though is now called ''Aegagropila linnaei''. Algae balls are significant in the hobby aquarium trade.


Taxonomy and selected species

''Cladophora'' is a green macroalgae alga classified within the Chlorophyta division. ''Cladophora'' has over 183 species within its genus that are very hard to tell apart and classify, mainly because of the great variation in their appearances, which is affected by habitat, age and environmental conditions.Gestinari, L., et al. (2010)
Distribution of ''Cladophora'' species (Cladophorales, Chlorophyta) along the Brazilian Coast.
''Phytotaxa'' 14 22.
Some species include: *'' Cladophora albida'' *'' Cladophora aokii'' *'' Cladophora brasiliana'' *'' Cladophora catenata'' *'' Cladophora coelothrix'' *'' Cladophora columbiana'' *'' Cladophora crispata'' *'' Cladophora dalmatica'' *'' Cladophora fracta'' *'' Cladophora glomerata'' *'' Cladophora graminea'' *'' Cladophora montagneana'' *'' Cladophora ordinata'' *'' Cladophora prolifera'' *'' Cladophora rivularis'' *'' Cladophora rupestris'' *'' Cladophora scopaeformis'' *'' Cladophora sericea'' *'' Cladophora socialis'' *'' Cladophora vagabunda'' '' Aegagropila linnaei'' (marimo) was formerly placed here as ''Cladophora aegagropila.''


References


Further reading

* *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


Images
of ''Cladophora'' at
AlgaeBase AlgaeBase is a global species database of information on all groups of algae, both seaweed, marine and freshwater algae, freshwater, as well as sea-grass. History AlgaeBase began in March 1996, founded by Michael D. Guiry, Michael Guiry. Text ...
*
''Cladophora'' Index.
Monterey Bay Aquarium {{Authority control Cladophoraceae Cladophorales genera Taxa named by Friedrich Traugott Kützing