From Wikipedia, the free encyclopedia
The lineage of algae according to Thomas Cavalier-Smith
. The exact number and placement of endosymbiotic events
is not yet clear, so this diagram can be taken only as a general guide
It represents the most parsimonious way of explaining the three types of endosymbiotic origins of chloroplasts. These types include the endosymbiotic events of cyanobacteria
, red algae
and green algae
, leading to the hypothesis of the supergroups Archaeplastida
respectively. However, the monophyly of Cabozoa has been refuted and the monophylies of Archaeplastida and Chromalveolata are currently strongly challenged. Endosymbiotic events are noted by dotted lines.
Though the prokaryotic Cyanobacteria
(commonly referred to as blue-green algae) were traditionally included as "algae" in older textbooks, many modern sources regard this as outdated
as they are now considered to be closely related to bacteria.
The term algae
is now restricted to eukaryotic
All true algae therefore have a nucleus enclosed within a membrane and chloroplasts
bound in one or more membranes.
Algae constitute a paraphyletic
as they do not include all the descendants of the last universal ancestor
nor do they all descend from a common algal ancestor, although their chloroplasts seem to have a single origin.
Diatoms are also examples of algae.
Algae lack the various structures that characterize land plants, such as phyllids (leaves) and rhizoids
in nonvascular plants, or leaves
, and other organs
that are found in tracheophytes
(vascular plants). Many are photoautotrophic
, although some groups contain members that are mixotrophic
, deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy
, or phagotrophy
. Some unicellular species
rely entirely on external energy sources and have limited or no photosynthetic apparatus.
Nearly all algae have photosynthetic machinery ultimately derived from the Cyanobacteria
, and so produce oxygen
as a by-product of photosynthesis, unlike other photosynthetic bacteria
such as purple
and green sulfur bacteria
. Fossilized filamentous algae from the Vindhya
basin have been dated back to 1.6 to 1.7 billion years ago.
The first alga to have its genome sequenced was Cyanidioschyzon merolae.
Etymology and study
The singular alga
is the Latin word for a particular seaweed and retains that meaning in English.
is obscure. Although some speculate that it is related to Latin algēre
, "be cold",
there is no known reason to associate seaweed with temperature. A more likely source is alliga
, "binding, entwining."
Since Algae has become a biological classification, alga can also mean one classification under Algae, parallel to a fungus being a species of fungi, a plant being a species of plant, and so on.
The ancient Greek
word for seaweed was φῦκος
(fūkos or phykos), which could mean either the seaweed, probably Red Algae, or a red dye derived from it. The Latinization, fūcus
, meant primarily the cosmetic rouge. The etymology is uncertain, but a strong candidate has long been some word related to the Biblical פוך
(pūk), "paint" (if not that word itself), a cosmetic eye-shadow used by the ancient Egyptians and other inhabitants of the eastern Mediterranean. It could be any color: black, red, green, blue.
Accordingly the modern study of marine and freshwater algae is called either phycology
or algology. The name Fucus appears in a number of taxa
have been traditionally included among the Algae, recent works usually exclude them due to large differences such as the lack of membrane-bound organelles
, the presence of a single circular chromosome
, the presence of peptidoglycan
in the cell walls, and ribosomes
different in size and content from those of the Eukaryotes
Rather than in chloroplasts
, they conduct photosynthesis on specialized infolded cytoplasmic membranes called thylakoid membranes
. Therefore, they differ significantly from the Algae despite occupying similar ecological niches.
By modern definitions Algae are Eukaryotes
and conduct photosynthesis within membrane-bound organelles called chloroplasts
. Chloroplasts contain circular DNA
and are similar in structure to Cyanobacteria, presumably representing reduced cyanobacterial endosymbionts
. The exact nature of the chloroplasts is different among the different lines of Algae, reflecting different endosymbiotic events. The table below describes the composition of the three major groups of Algae. Their lineage relationships are shown in the figure in the upper right. Many of these groups contain some members that are no longer photosynthetic. Some retain plastids
, but not chloroplasts, while others have lost plastids entirely.
Phylogeny based on plastid.
not nucleocytoplasmic genealogy:
||These Algae have primary chloroplasts, i.e. the chloroplasts are surrounded by two membranes and probably developed through a single endosymbiotic event. The chloroplasts of Red Algae have chlorophylls a and c (often), and phycobilins, while those of Green Algae have chloroplasts with chlorophyll a and b. Higher plants are pigmented similarly to Green Algae and probably developed from them, and thus Chlorophyta is a sister taxon to the plants; sometimes they are grouped as Viridiplantae.
|Excavata and Rhizaria
These groups have green chloroplasts containing chlorophylls a
and b 
. Their chloroplasts are surrounded by four and three membranes
, respectively, and were probably retained from ingested Green Algae.
, which belong to the phylum Euglenozoa
, live primarily in freshwater and have chloroplasts with only three membranes. It has been suggested that the endosymbiotic Green Algae were acquired through myzocytosis
rather than phagocytosis
|Chromista and Alveolata
These groups have chloroplasts containing chlorophylls a and d, and phycobilins. The latter chlorophyll type is not known from any prokaryotes or primary chloroplasts, but genetic similarities with the Red Algae suggest a relationship there.
In the first three of these groups (Chromista
), the chloroplast has four membranes, retaining a nucleomorph
, and they likely share a common pigmented ancestor, although other evidence casts doubt on whether the Heterokonts
, and Cryptomonads
are in fact more closely related to each other than to other groups.
The typical dinoflagellate
chloroplast has three membranes, but there is considerable diversity in chloroplasts within the group, and it appears there were a number of endosymbiotic events.
, a group of closely related parasites, also have plastids
. Apicoplasts are not photosynthetic but appear to have a common origin with Dinoflagellate
(1811—1866) was the first to divide the Algae into four divisions based on their pigmentation. This is the first use of a biochemical criterion in plant systematics. Harvey's four divisions are: Red Algae (Rhodophyta), Brown Algae (Heteromontophyta), Green Algae (Chlorophyta) and Diatomaceae.
Relationship to higher plants
The first plants on earth evolved from shallow freshwater algae much like Chara
some 400 million years ago. These probably had an isomorphic alternation of generations
and were probably filamentous. Fossils of isolated land plant spores suggest land plants may have been around as long as 475 million years ago.
forest exhibit at the Monterey Bay Aquarium. A three-dimensional, multicellular thallus.
Most of the simpler algae are unicellular flagellates
, but colonial and non-motile forms have developed independently among several of the groups. Some of the more common organizational levels, more than one of which may occur in the life cycle
of a species, are
- Colonial: small, regular groups of motile cells
- Capsoid: individual non-motile cells embedded in mucilage
- Coccoid: individual non-motile cells with cell walls
- Palmelloid: non-motile cells embedded in mucilage
- Filamentous: a string of non-motile cells connected together, sometimes branching
- Parenchymatous: cells forming a thallus with partial differentiation of tissues
In three lines even higher levels of organization have been reached, with full tissue differentiation. These are the brown algae
—some of which may reach 50 m in length (kelps
—the red algae
and the green algae
The most complex forms are found among the green algae (see Charales
), in a lineage that eventually led to the higher land plants
. The point where these non-algal plants begin and algae stop is usually taken to be the presence of reproductive organs with protective cell layers, a characteristic not found in the other alga groups.
Some species of algae form symbiotic relationships
with other organisms. In these symbioses
, the algae supply photosynthates (organic substances) to the host organism providing protection to the algal cells. The host organism derives some or all of its energy requirements from the algae. Examples are as follows.
are defined by the International Association for Lichenology to be "an association of a fungus
and a photosynthetic symbiont
resulting in a stable vegetative body having a specific structure."
The fungi, or mycobionts, are from the Ascomycota
with a few from the Basidiomycota
. They are not found alone in nature but when they began to associate is not known.
One mycobiont associates with the same phycobiont species, rarely two, from the Green Algae
, except that alternatively the mycobiont may associate with the same species of Cyanobacteria
(hence "photobiont" is the more accurate term). A photobiont may be associated with many specific mycobionts or live independently; accordingly, lichens are named and classified as fungal species.
The association is termed a morphogenesis because the lichen has a form and capabilities not possessed by the symbiont species alone (they can be experimentally isolated). It is possible that the photobiont triggers otherwise latent genes in the mycobiont.
Algae of the Dinoflagellate
phylum are often endosymbionts
in the cells of marine invertebrates, where they accelerate host-cell metabolism by generating immediately available sugar and oxygen through photosynthesis
using incident light and the carbon dioxide produced in the host. Endosymbiont algae in the Stony Corals are described by the term zooxanthellae
, with the host Stony Corals called on that account hermatypic corals
, which although not a taxon
are not in healthy condition without their endosymbionts. Zooxanthellae belong almost entirely to the genus Symbiodinium
The loss of Symbiodinium
from the host is known as coral bleaching
, a condition which unless corrected leads to the deterioration and loss of the reef.
Green Algae live close to the surface of some sponges, for example, breadcrumb sponge (Halichondria panicea
). The alga is thus protected from predators; the sponge is provided with oxygen and sugars which can account for 50 to 80% of sponge growth in some species.
, the three main algal Phyla
, have life-cycles which show tremendous variation with considerable complexity. In general there is an asexual phase where the seaweed's cells are diploid
, a sexual phase where the cells are haploid
followed by fusion of the male and female gametes
. Asexual reproduction is advantageous in that it permits efficient population increases, but less variation is possible. Sexual reproduction allows more variation, but is more costly. Often there is no strict alternation between the sporophyte and also because there is often an asexual phase, which could include the fragmentation of the thallus.
The Algal Collection of the U.S. National Herbarium
(located in the National Museum of Natural History
) consists of approximately 320500 dried specimens, which, although not exhaustive (no exhaustive collection exists), gives an idea of the order of magnitude of the number of algal species (that number remains unknown).
Estimates vary widely. For example, according to one standard textbook,
in the British Isles the UK Biodiversity Steering Group Report
estimated there to be 20000 algal species in the UK. Another checklist reports only about 5000 species. Regarding the difference of about 15000 species, the text concludes: "It will require many detailed field surveys before it is possible to provide a reliable estimate of the total number of species ...."
Regional and group estimates have been made as well: 5000—5500 species of Red Algae worldwide, "some 1300 in Australian Seas,"
400 seaweed species for the western coastline of South Africa
669 marine species from California (U.S.A.),
642 in the check-list of Britain and Ireland,
and so on, but lacking any scientific basis or reliable sources, these numbers have no more credibility than the British ones mentioned above. Most estimates also omit the microscopic Algae, such as the phytoplankta
The topic of distribution of algal species has been fairly well studied since the founding of phytogeography
in the mid-19th century AD.
Algae spread mainly by the dispersal of spores
analogously to the dispersal of Plantae by seeds and spores. Spores are everywhere in all parts of the Earth: the waters fresh and marine, the atmosphere, free-floating and in precipitation or mixed with dust, the humus
and in other organisms, such as humans. Whether a spore is to grow into an organism depends on the combination of the species and the environmental conditions.
The spores of fresh-water Algae are dispersed mainly by running water and wind, as well as by living carriers.
The bodies of water into which they are transported are chemically selective. Marine spores are spread by currents. Ocean water is temperature selective, resulting in phytogeographic zones, regions and provinces.
To some degree the distribution of Algae is subject to floristic discontinuities caused by geographical features, such as Antarctica
, long distances of ocean or general land masses. It is therefore possible to identify species occurring by locality, such as "Pacific Algae" or "North Sea Algae". When they occur out of their localities, it is usually possible to hypothesize a transport mechanism, such as the hulls of ships. For example, Ulva reticulata
and Ulva fasciata
travelled from the mainland to Hawaii
in this manner.
Mapping is possible for select species only: "there are many valid examples of confined distribution patterns."
For example, Clathromorphum
is an arctic genus and is not mapped far south of there.
On the other hand, scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."
Algae are prominent in bodies of water, common in terrestrial environments and are found in unusual environments, such as on snow
and on ice
. Seaweeds grow mostly in shallow marine waters, under 100 metres (330 ft); however some have been recorded to a depth of 360 metres (1,180 ft)
The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live suspended in the water column (phytoplankton
) provide the food base for most marine food chains
. In very high densities (algal blooms
) these algae may discolor the water and outcompete, poison, or asphyxiate other life forms.
Algae are variously sensitive to different factors, which has made them useful as biological indicators in the Ballantine Scale
and its modification.
, an Algae derivative, has a number of commercial uses.
To be competitive and independent from fluctuating support from (local) policy on the long run, biofuels should equal or beat the cost level of fossil fuels. Here, algae based fuels hold great promise, directly related to the potential to produce more biomass per unit area in a year than any other form of biomass. The break-even point for algae-based biofuels should be within reach in about ten years.
This kind of ore they often gather and lay on great heapes, where it heteth and rotteth, and will have a strong and loathsome smell; when being so rotten they cast on the land, as they do their muck, and thereof springeth good corn, especially barley ... After spring-tydes or great rigs of the sea, they fetch it in sacks on horse backes, and carie the same three, four, or five miles, and cast it on the lande, which doth very much better the ground for corn and grass.
Today Algae are used by humans in many ways; for example, as fertilizers
, soil conditioners
and livestock feed.
Aquatic and microscopic species are cultured in clear tanks or ponds and are either harvested or used to treat effluents pumped through the ponds. Algaculture
on a large scale is an important type of aquaculture
in some places. Maerl
is commonly used as a soil conditioner.
Naturally growing seaweeds are an important source of food, especially in Asia. They provide many vitamins including: A, B1
, and are rich in iodine
In addition commercially cultivated microalgae
, including both Algae and Cyanobacteria
, are marketed as nutritional supplements, such as Spirulina
and the Vitamin-C supplement, Dunaliella
, high in beta-carotene
Algae are national foods of many nations: China
consumes more than 70 species, including fat choy
, a cyanobacterium
considered a vegetable; Japan
, over 20 species; Ireland
is used to make "laver bread" in Wales
where it is known as bara lawr
; in Korea
; in Japan
. It is also used along the west coast of North America from California
to British Columbia
, in Hawaii
and by the Māori
of New Zealand
. Sea lettuce
are a salad ingredient in Scotland
- Sewage can be treated with algae, reducing the need for greater amounts of toxic chemicals than are already used.
- Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested, the enriched algae itself can be used as fertilizer.
The natural pigments
produced by algae can be used as an alternative to chemical dyes
and coloring agents.
Carrageenan, from the red alga Chondrus crispus, is used as a stabiliser in milk products.
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- Gayral, Paulette (1966) (in French). Les Algues de côtes françaises (manche et atlantique), notions fondamentales sur l'écologie, la biologie et la systématique des algues marines. Paris: Doin, Deren et Cie.
- Guiry, M.D.; Blunden, G. (1991). Seaweed Resources in Europe: Uses and Potential. John Wiley & Sons. ISBN 0471929476.
- Míguez Rodríguez, Luís (1998) (in Galician). Algas mariñas de Galicia: bioloxía, gastronomía, industria. Vigo: Edicións Xerais de Galicia. ISBN 84-8302-263-X.
- Otero, J. (2002) (in Galician). Guía das macroalgas de Galicia. A Coruña: Baía Edicións. ISBN 84-89803-22-6.
- Bárbara, I.; Cremades, J. (1993) (in Spanish). Guía de las algas del litoral gallego. A Coruña: Concello da Coruña - Casa das Ciencias.
- Kjellman, Frans Reinhold (1883). The algae of the Arctic Sea: a survey of the species, together with an exposition of the general characters and the development of the flora. 20. Stockholm: Kungl. Svenska vetenskapsakademiens handlingar. pp. 1–350.
- Lund, Søren Jensen (1959). The Marine Algae of East Greenland. Kövenhavn: C.A. Reitzel. ISBN 9584734.
- Faroe Islands
- Børgesen, Frederik (1903, 1970 reprint). "Marine Algae". in Warming, Eugene. Botany of the Faröes Based Upon Danish Investigations. Part II. Det nordiske Forlag. pp. 339–532. .
- Canary Islands
- Børgesen, Frederik (1925, 1926, 1927, 1929, 1930, 1936). Marine Algae from the Canary Islands. København: Bianco Lunos.
- Gayral, Paulette (1958) (in French). Algues de la côte atlantique marocaine. Casablanca: Rabat [Société des sciences naturelles et physiques du Maroc].
- South Africa
- Stegenga, H.; Bolton, J.J.; Anderson, R.J. (1997). Seaweeds of the South African West Coast. Bolus Herbarium, University of Cape Town. ISBN 079921793x.
- North America
- Abbott, I.A.; Hollenberg, G.J. (1976). Marine Algae of California. California: Stanford University Press. ISBN 0804708673.
- Greeson, Phillip E. (1982). An annotated key to the identification of commonly occurring and dominant genera of Algae observed in the Phytoplankton of the United States. Washington, D.C.: U.S. Department of the Interior, Geological Survey. http://www.archive.org/details/annotatedkeytoid00gree. Retrieved 2008-12-19.
- Taylor, William Randolph (1937, 1957, 1962, 1969). Marine Algae of the Northeastern Coast of North America. Ann Arbor: University of Michigan Press. ISBN 0472049046.
- Wehr, J D; Sheath, R G (2003). Freshwater Algae of North America: Ecology and Classification. USA: Academic Press. ISBN 0127415505.