Caulerpa

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Template:Short description Template:Automatic taxobox

Caulerpa is a genus of seaweeds in the family Caulerpaceae (among the green algae). They are unusual because they consist of only one cell with many nuclei, making them among the biggest single cells in the world.

Referring to the crawling habit of its thallus, the name means 'stem (that) creeps', from the Ancient Greek Script error: No such module "lang". (Script error: No such module "Lang"., ‘stalk’) and Script error: No such module "lang". (Script error: No such module "Lang"., ‘to creep’).[1]

Taxonomy and nomenclature

First described by Jean Vincent Lamouroux in 1809, Caulerpa is the only genus under the family Caulerpaceae, from the order Bryopsidales, class Ulvophyceae, and phylum Chlorophyta.[2] Through the use of tufA gene sequencing, it was revealed that Pseudochlorodesmis F. Børgesen was a sister clade of Caulerpa.[3] Cremen et al.[4] proposed a new classification scheme in Bryopsidales, wherein Caulerpaceae and Halimedaceae were described as sister families.

Species discrimination via morphology-based identification is often hampered by the high degree of variation among traits. Thus, several species are often misidentified without the use of molecular analyses.[5] As of 2019, there are 101 accepted species, with 40 varieties and 67 forms.[6][7]

Biology

Caulerpa species support their large cell size by having the cytoplasm circulate constantly, supported by a network of microtubules. This behavior was known in 1967.[8]

The cytoplasm does not leak out when the cell is cut. Regeneration is directional, with rhizoids at the bottom and fronds at the top.[9]

The genus produces a number of secondary metabolites thought to be related to its toxicity and peppery taste. These include the red pigment alkaloid caulerpin and its derivative caulerchlorin and the amine mixture caulerpicin.

Morphology

External characteristics

File:Caulerpa racemosa algae.jpg
Caulerpa racemosa showing irregular vesiculate ramuli (Hobgood, n.d.)

The genus Caulerpa exhibits several different growth forms. They possess a stolon with rhizoids which grow downward, anchoring the plant to the substrate. The stolons support upright fronds or leaves known as assimilators. The structure of the assimilator may be ligulate (leaf-like, e.g. Caulerpa prolifera), or they may have a central axis known as a rachis. The rachis may have lateral branchlets known as ramuli which themselves come in different forms (terete, turbinate, clavate, peltate, falcate, vesiculate) and arrangements: distichous - ramuli are arranged evenly opposite each other (e.g. Caulerpa taxifolia), irregular - vesiculate ramuli with no distinct arrangement (e.g. Caulerpa racemosa), verticillate - whorled ramuli (e.g. Caulerpa cuppresoides).[2]

Internal anatomy

Caulerpa is coenocytic, meaning it has a multinucleate thallus organization. It is also siphonous, meaning unlike other algae, the thallus and the nuclei are not separated by cell walls. They are instead one long mass of protoplasm surrounded by a single cell wall.[10] The genus also possesses trabeculae, which are inward growing cell wall extensions that pass through the central lumen of siphons. It is hypothesized that these provide the thallus with structural support, facilitate diffusion to the inner cytoplasm, and can possibly help in determining cell shape - which may contribute to the diversity of growth forms found in the genus.[11]

Distribution

Caulerpa is mostly found in tropical regions, however its distribution may also extend up to temperate locations. Diversity is highest in the Caribbean and the Indo-Malay region, as well as in southern Australia, where a majority of Caulerpa species are endemic.[12][2]

Ecology

The genus is typically found in shallow intertidal zones and can reach up to depths of 100 meters. They are known to be able to adapt and thrive in different environmental conditions, which contributes to their potential for becoming invasive species. Some of their traits include having a high tolerance for a wide range of temperatures, their capacity for asexual reproduction through rhizoid extension and fragmentation, their fast growth rate, as well as their capacity for nutrient intake from sediments through their rhizoids.[2][13] This last trait gives them a competitive edge over other macroalgal species who mainly absorb nutrients from the water column.[14]

Life history

Many studies on the life cycle of Caulerpa have been found to contradict each other, leading researchers to conclude that it varies with species and geographical region.[2] Earlier research revealed that Caulerpa has a diplontic life cycle with a diploid vegetative phase and haploid biflagellate gametes. However, it was later found that ploidy status and genome size can vary within and between species.[15][16][17]

Exploitation and cultivation

File:USAID Measuring Impact Conservation Enterprise Retrospective (Philippines; Nagkakaisang Tribu ng Palawan) (26420465388).jpg
Edible fresh latô (Caulerpa lentillifera) sold at a fish market in the Philippines

Some species of Caulerpa are edible. The two most commonly eaten are Caulerpa lentillifera and Caulerpa racemosa, both called "sea grapes" in English. Both are traditionally harvested in the wild and sold in local markets in Southeast Asia, Oceania, and East Asia. They are eaten raw in salads and have a characteristic "sea" flavor and a crunchy texture.[18]

Both species are cultivated in aquaculture. Their cultivation began in the 1950s in Cebu, Philippines, after accidental introduction of C. lentillifera to fish ponds.[19] Cultivation of C. lentillifera continued in Japan in 1986, where it was cultivated in tanks in the tropical waters of Okinawa.[20] Commercial cultivation has since spread to other countries, including Vietnam, Taiwan, and China (in Fujian and Hainan). Most are for domestic consumption, but they are also exported to Japan.[21]

Cultivation of Caulerpa is convenient because they can propagate through fragmentation. There are several farming techniques being used to cultivate Caulerpa, which typically involve tying fragments to different types of infrastructure. Many Pacific countries such as Japan, Philippines, Vietnam, and Samoa use the off-bottom method, where the seaweeds are grown a few meters above the ground on cages or trays.[22] The bottom-planting method is also used in the Philippines, and involves growing the Caulerpa on a substrate.[23] Land-based raceways in hatchery-type facilities offer a more controlled environment for cultures, and have been used more in recent years.[2][22]

Chemical composition

Caulerpa contains a high amount of iron (up to 81.3 mg per 100 g of dry matter in C. racemosa),[24] magnesium, and calcium.[22] Water content is species-specific and generally ranges from 75 – 94%.[22] The genus is known to have a high bioaccumulation rate,[25][18][26] which can make it less than ideal to consume on a regular basis. Carbohydrate content can range from 3.6 - 83.2% of dry matter depending on the species.[22] The main pigments of Caulerpa are chlorophyll a and b.[18] It has a high diversity of chemical compounds which have pharmaceutical potential. Although the genus is known to exhibit high toxicity, it was found to be of low risk to humans.[27]

Utilization

Aside from being a source of food, Caulerpa has several uses from bioremediation, to fertilizer, and health and wellness.[2] The anti-oxidant compounds of Caulerpa have been well-studied, and these are used in treating various diseases and health conditions such as cancer and cardiovascular disorders.[24][28][29][30] Caulerpa has been shown to be effective in filtering water used in culturing fish, mollusks, and shrimp[31] (in particular C. lentillifera[32]). The use of Caulerpa as a biofertilizer has also been studied particularly in India, where fertilizers composed of 25% Caulerpa extracts enhanced the growth and reduced the total sugar content, among other things, of Vigna mungo.[33][34][35]

Invasive behaviour

Another species, Caulerpa taxifolia, has become an invasive species in the Mediterranean Sea, Australia and southern California (where it has since been eradicated). In U.S. waters, the Mediterranean strain of Caulerpa taxifolia is listed as a federal noxious weed, under the Plant Protection Act. The Aquatic Nuisance Species Taskforce has also created a National Management Plan for the Genus Caulerpa. The state of California also prohibits possession of nine different species of Caulerpa.

It is thought that Caulerpa species have such invasive properties in these regions due to their capability to thrive in temperate waters, along with their freedom from natural predators. Most Caulerpa species evolved in tropical waters, where herbivores have immunity to toxic compounds (mainly caulerpicin) within the alga. Temperate water herbivores have no natural immunity to these toxins, allowing Caulerpa to grow unchecked if introduced to temperate waters.

C. racemosa has recently been found in waters around Crete, where it is thought to have contributed to a significant reduction in fisheries. The alga has invaded the area from the warmer waters of the Red Sea.

C. cylindracea, which is native to Australia, has also become an invasive species in the Mediterranean.[36]

In New Zealand waters, invasive exotic caulerpa species have been found at Great Barrier Island, Great Mercury Island, and Goat Island Marine Reserve.[37][38]

Use in aquariums

Caulerpa is common in the aquarium hobby as a nitrate absorber because of its rapid growth under relatively adverse conditions. It may also be used in refugiums for a long-term nitrite absorber. Many introductions of invasive Caulerpa to the wild are thought to have occurred via aquarium dumping although there is no proof that this is so. For this reason, some aquarium hobbyists have begun using Chaetomorpha or an algae scrubber instead.[39][40]

Species

File:Feather algae, Caulerpa sertularioides at 11 meters depth on ridge.jpg
Feather algae, Caulerpa sertularioides at 11 metres' depth on ridge
File:Oval sea grapes, Caulerpa racemosa var clavifera, at 5 meters depth.jpg
Oval sea grapes, Caulerpa racemosa var. clavifera, at 5 metres' depth

The following species are currently recognized in the genus Caulerpa:[41]

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  • Caulerpa agardhii Weber-van Bosse, 1898Script error: No such module "Check for unknown parameters".
  • Caulerpa alternans Womersley, 1956Script error: No such module "Check for unknown parameters".
  • Caulerpa ambigua Okamura, 1897Script error: No such module "Check for unknown parameters".
  • Caulerpa andamanensis (W.R.Taylor) Draisma, Prudhomme & Sauvage, 2014Script error: No such module "Check for unknown parameters".
  • Caulerpa antoensis Yamada, 1940Script error: No such module "Check for unknown parameters".
  • Caulerpa articulata Harvey, 1855Script error: No such module "Check for unknown parameters".
  • Caulerpa ashmeadii Harvey, 1858Script error: No such module "Check for unknown parameters".
  • Caulerpa bartoniae G.Murray, 1896Script error: No such module "Check for unknown parameters".
  • Caulerpa bikinensis W.R.Taylor, 1950Script error: No such module "Check for unknown parameters".
  • Caulerpa biserrulata Sonder, 1871Script error: No such module "Check for unknown parameters".
  • Caulerpa brachypus Harvey, 1860Script error: No such module "Check for unknown parameters".
  • Caulerpa brownii (C.Agardh) Endlicher, 1843Script error: No such module "Check for unknown parameters".
  • Caulerpa buginensis E.Verheij & Prud'homme van Reine, 1993Script error: No such module "Check for unknown parameters".
  • Caulerpa cactoides (Turner) C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa chamnitzia
  • Caulerpa chemnitzia (Esper) J.V.Lamouroux, 1809Script error: No such module "Check for unknown parameters".
  • Caulerpa cliftonii Harvey, 1863Script error: No such module "Check for unknown parameters".
  • Caulerpa constricta I.R.Price, J.M.Huisman & M.A.Borowitzka, 1998Script error: No such module "Check for unknown parameters".
  • Caulerpa coppejansii G.Belton & Prud'homme, 2019Script error: No such module "Check for unknown parameters".
  • Caulerpa corynephora Montagne, 1842Script error: No such module "Check for unknown parameters".
  • Caulerpa crispata (Harvey) G.Belton & Gurgel, 2019Script error: No such module "Check for unknown parameters".
  • Caulerpa cupressoides (Vahl) C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa cylindracea Sonder, 1845Script error: No such module "Check for unknown parameters".
  • Caulerpa denticulata Decaisne, 1841Script error: No such module "Check for unknown parameters".
  • Caulerpa dichotoma Svedelius, 1906Script error: No such module "Check for unknown parameters".
  • Caulerpa diligulata Kraft & A.J.K.Millar, 2000Script error: No such module "Check for unknown parameters".
  • Caulerpa ellistoniae Womersley, 1955Script error: No such module "Check for unknown parameters".
  • Caulerpa elongata Weber Bosse, 1898Script error: No such module "Check for unknown parameters".
  • Caulerpa falcifolia Harvey & Bailey, 1851Script error: No such module "Check for unknown parameters".
  • Caulerpa faridii Nizamuddin, 1964Script error: No such module "Check for unknown parameters".
  • Caulerpa fastigiata Montagne, 1837Script error: No such module "Check for unknown parameters".
  • Caulerpa fergusonii G.Murray, 1891Script error: No such module "Check for unknown parameters".
  • Caulerpa filicoides Yamada, 1936Script error: No such module "Check for unknown parameters".
  • Caulerpa filiformis (Suhr) Hering, 1841Script error: No such module "Check for unknown parameters".
  • Caulerpa flagelliformis
  • Caulerpa flexilis J.V.Lamouroux ex C.Agardh, 1823Script error: No such module "Check for unknown parameters".
  • Caulerpa floridana W.R.Taylor, 1960Script error: No such module "Check for unknown parameters".
  • Caulerpa freycinettii
  • Caulerpa geminata Harvey, 1855Script error: No such module "Check for unknown parameters".
  • Caulerpa gracilis (Zan.) Web. V. B.Script error: No such module "Check for unknown parameters".
  • Caulerpa hedleyi Weber-van Bosse, 1910Script error: No such module "Check for unknown parameters".
  • Caulerpa heterophylla I.R.Price, J.M.Huisman & M.A.Borowitzka, 1998Script error: No such module "Check for unknown parameters".
  • Caulerpa hodgkinsoniae J.Agardh, 1887Script error: No such module "Check for unknown parameters".
  • Caulerpa holmesiana G.Murray, 1891Script error: No such module "Check for unknown parameters".
  • Caulerpa integerrima (Zanardini) M.J.Wynne, Verbruggen & D.L.Angel, 2009Script error: No such module "Check for unknown parameters".
  • Caulerpa juniperoides J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa kempfii A.B.Joly & S.M.B.Pereira, 1975Script error: No such module "Check for unknown parameters".
  • Caulerpa laetivirens
  • Caulerpa lagara Carruthers, Walker & Huisman, 1993Script error: No such module "Check for unknown parameters".
  • Caulerpa lamourouxii (Turner) C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa lanuginosa J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa lentillifera J.Agardh, 1837Script error: No such module "Check for unknown parameters".
  • Caulerpa lessonii Bory de Saint-Vincent, 1828Script error: No such module "Check for unknown parameters".
  • Caulerpa longifolia C.Agardh, 1823Script error: No such module "Check for unknown parameters".
  • Caulerpa lucasii Prud'homme, Draisma & G.Belton, 2019Script error: No such module "Check for unknown parameters".
  • Caulerpa macra (Weber-van Bosse) Draisma & Prud'homme, 2014Script error: No such module "Check for unknown parameters".
  • Caulerpa macrodisca Decaisne, 1842Script error: No such module "Check for unknown parameters".
  • Caulerpa manorensis Nizamuddin, 1964Script error: No such module "Check for unknown parameters".
  • Caulerpa matsueana Yamada, 1940Script error: No such module "Check for unknown parameters".
  • Caulerpa megadisca G.S.Belton & C.F.D.Gurgel, 2014Script error: No such module "Check for unknown parameters".
  • Caulerpa mexicana Sonder ex Kützing, 1849Script error: No such module "Check for unknown parameters".
  • Caulerpa microphysa (Weber Bosse) Feldmann, 1955Script error: No such module "Check for unknown parameters".
  • Caulerpa muelleri Sonder, 1853Script error: No such module "Check for unknown parameters".
  • Caulerpa murrayi Weber Bosse, 1898Script error: No such module "Check for unknown parameters".
  • Caulerpa nummularia Harvey ex J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa obscura Sonder, 1845Script error: No such module "Check for unknown parameters".
  • Caulerpa occidentalis (J. Ag.) Boergs.Script error: No such module "Check for unknown parameters".
  • Caulerpa okamurae Weber Bosse, 1897Script error: No such module "Check for unknown parameters".
  • Caulerpa oligophylla Montagne, 1842Script error: No such module "Check for unknown parameters".
  • Caulerpa ollivieri Dostál, 1929Script error: No such module "Check for unknown parameters".
  • Caulerpa opposita Coppejans & Meinesz, 1988Script error: No such module "Check for unknown parameters".
  • Caulerpa papillosa J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa parvifolia Harvey, 1860Script error: No such module "Check for unknown parameters".
  • Caulerpa parvula Svedelius, 1906Script error: No such module "Check for unknown parameters".
  • Caulerpa paspaloides (Bory de Saint-Vincent) Greville, 1830Script error: No such module "Check for unknown parameters".
  • Caulerpa pennata J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa pinnata C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa plumulifera Zanardini, 1878Script error: No such module "Check for unknown parameters".
  • Caulerpa prolifera (Forsskål) J.V.Lamouroux, 1809Script error: No such module "Check for unknown parameters".
  • Caulerpa pusilla (Kützing) J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa qureshii Nizamuddin, 1964Script error: No such module "Check for unknown parameters".
  • Caulerpa racemosa (Forsskål) J.Agardh, 1873Script error: No such module "Check for unknown parameters".
  • Caulerpa remotifolia Sonder, 1853Script error: No such module "Check for unknown parameters".
  • Caulerpa reniformis G.R.South & Skelton, 2003Script error: No such module "Check for unknown parameters".
  • Caulerpa reyesii Meñez & Calumpong, 1982Script error: No such module "Check for unknown parameters".
  • Caulerpa scalpelliformis (R.Brown ex Turner) C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa sedoides C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa selago (Turner) C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa serrulata (Forsskål) J.Agardh, 1837Script error: No such module "Check for unknown parameters".
  • Caulerpa sertularioides (S.G.Gmelin) M.Howe, 1905Script error: No such module "Check for unknown parameters".
  • Caulerpa seuratii Weber-van Bosse, 1910Script error: No such module "Check for unknown parameters".
  • Caulerpa simpliciuscula (R.Brown ex Turner) C.Agardh, 1823Script error: No such module "Check for unknown parameters".
  • Caulerpa spathulata Womersley & A.Bailey, 1970Script error: No such module "Check for unknown parameters".
  • Caulerpa subserrata Okamura, 1897Script error: No such module "Check for unknown parameters".
  • Caulerpa taxifolia (M.Vahl) C.Agardh, 1817Script error: No such module "Check for unknown parameters".
  • Caulerpa tongaensis Papenfuss, 1943Script error: No such module "Check for unknown parameters".
  • Caulerpa trifaria Harvey, 1863Script error: No such module "Check for unknown parameters".
  • Caulerpa turbinata (J. Ag.) EubankScript error: No such module "Check for unknown parameters".
  • Caulerpa urvilleana Montagne, 1845Script error: No such module "Check for unknown parameters".
  • Caulerpa veravalensis Thivy & V.D.Chauhan, 1963Script error: No such module "Check for unknown parameters".
  • Caulerpa verticillata J.Agardh, 1847Script error: No such module "Check for unknown parameters".
  • Caulerpa vesiculifera (Harvey) Harvey, 1863Script error: No such module "Check for unknown parameters".
  • Caulerpa vickersiae Børgesen, 1911Script error: No such module "Check for unknown parameters".
  • Caulerpa vieillardii Kützing, 1863Script error: No such module "Check for unknown parameters".
  • Caulerpa vitifolia (Bonpland) J.V.LamourouxScript error: No such module "Check for unknown parameters".
  • Caulerpa webbiana Montagne, 1837Script error: No such module "Check for unknown parameters".
  • Caulerpa wysorii T.Sauvage & M.J.Wynne, 2021Script error: No such module "Check for unknown parameters".
  • Caulerpa zeyheri Kützing, 1857Script error: No such module "Check for unknown parameters".


See also

References

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  15. ↑ Script error: No such module "Citation/CS1".
  16. ↑ Collado-Vides L, Ruesink J (2002)Morphological plasticity and invasive potential. In: Williams E, Grosholz E (eds) International Caulerpa taxifolia Conference Proceedings. San Diego, California, pp 88–118
  17. ↑ Script error: No such module "Citation/CS1".
  18. ↑ a b c Script error: No such module "Citation/CS1".
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  24. ↑ a b Script error: No such module "Citation/CS1".
  25. ↑ Script error: No such module "Citation/CS1".
  26. ↑ Script error: No such module "Citation/CS1".
  27. ↑ Script error: No such module "Citation/CS1".
  28. ↑ Script error: No such module "Citation/CS1".
  29. ↑ Script error: No such module "Citation/CS1".
  30. ↑ Script error: No such module "Citation/CS1".
  31. ↑ Script error: No such module "Citation/CS1".
  32. ↑ Script error: No such module "Citation/CS1".
  33. ↑ Script error: No such module "Citation/CS1".
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  40. ↑ Script error: No such module "citation/CS1".
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Further reading

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External links

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