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Algae
A variety of algae growing on the sea bed in shallow waters off the New South Wales coast
A variety of algae growing on the sea bed in shallow waters off the New South Wales coast
Scientific classification
Domain: Eukaryota
Groups included
Cladistically included but traditionally excluded taxa
The lineage of algae according to Thomas Cavalier-Smith. The exact number and placement of endosymbiotic events is currently unknown, so this diagram can be taken only as a general guide[1][2] It represents the most parsimonious way of explaining the three types of endosymbiotic origins of plastids. These types include the endosymbiotic events of cyanobacteria, red algae and green algae, leading to the hypothesis of the supergroups Archaeplastida, Chromalveolata and Cabozoa respectively. However, the monophyly of Cabozoa has been refuted and the monophylies of Archaeplastida and Chromalveolata are currently strongly challenged.[citation needed] Endosymbiotic events are noted by dotted lines.

Algae (/ˈældʒiː/ or /ˈælɡiː/; singular alga /ˈælɡə/, Latin for "seaweed") are a very large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelp (large brown alga), that may grow up to 50 meters in length. Most are photosynthetic and "simple" because they lack many of the distinct cell organelles and cell types found in land plants. The largest and most complex marine forms are called seaweeds.

There is no clear consensus definition of algae. One definition is that algae “have chlorophyll a as their primary photosynthetic pigment and lack a sterile covering of cells around their reproductive cells”.[3] Other authors exclude all prokaryotes [4] and thus do not consider cyanobacteria (blue-green algae) as algae.[5]

Algae constitute a polyphyletic group[4] since they do not include a common ancestor, and although their plastids seem to have a single origin, from cyanobacteria,[1] they were acquired in different ways. Green algae are examples of algae that have primary chloroplasts derived from endosymbiotic cyanobacteria. Diatoms are examples of algae with secondary chloroplasts derived from an endosymbiotic red alga.[6]

Algae exhibit a wide range of reproductive strategies, from simple, asexual cell division to complex forms of sexual reproduction.[7]

Algae lack the various structures that characterize land plants, such as the phyllids (leaf-like structures) of bryophytes, rhizoids in nonvascular plants, and the roots, leaves, and other organs that are found in tracheophytes (vascular plants). Most are phototrophic, although some groups contain members that are mixotrophic, deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy, or phagotrophy. Some unicellular species such as the green algae Prototheca and Helicosporidium have become parasitic heterotrophs, relying entirely on external energy sources and have limited or no photosynthetic apparatus[8] Algae have photosynthetic machinery ultimately derived from cyanobacteria that 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.[9]

Etymology and study

Title page of Samuel Gottlieb Gmelin, Historia Fucorum, dated 1768.

The singular alga is the Latin word for a particular seaweed and retains that meaning in English.[10] The etymology is obscure. Although some speculate that it is related to Latin algēre, "be cold",[11] there is no known reason to associate seaweed with temperature. A more likely source is alliga, "binding, entwining."[12]

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.[13]

Accordingly the modern study of marine and freshwater algae is called either phycology or algology, depending on whether the Greek or Latin root is used. The name Fucus appears in a number of taxa.

Classification

False-color Scanning electron micrograph of the unicellular coccolithophore Gephyrocapsa oceanica.

Most algae except cyanobacteria contain 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 separate lineages 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[14] not nucleocytoplasmic genealogy:

Cyanobacteria

Supergroup affiliation Members Endosymbiont Summary
Primoplantae/
Archaeplastida 		Cyanobacteria 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 Green algae

These groups have green chloroplasts containing chlorophylls a and b.[15] Their chloroplasts are surrounded by four and three membranes, respectively, and were probably retained from ingested green algae.

Chlorarachniophytes, which belong to the phylum Cercozoa, contain a small nucleomorph, which is a relict of the algae's nucleus.

Euglenids, 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 Red algae

These groups have chloroplasts containing chlorophylls a and c, and phycobilins.The shape varies from plant to plant. they may be of discoid, plate-like, reticulate, cup-shaped, spiral or ribbon shaped. They have one or more pyrenoids to preserve protein and starch. The latter chlorophyll type is not known from any prokaryotes or primary chloroplasts, but genetic similarities with red algae suggest a relationship there.[16]

In the first three of these groups (Chromista), the chloroplast has four membranes, retaining a nucleomorph in Cryptomonads, and they likely share a common pigmented ancestor, although other evidence casts doubt on whether the Heterokonts, Haptophyta, and Cryptomonads are in fact more closely related to each other than to other groups.[2][17]

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.[1] The Apicomplexa, a group of closely related parasites, also have plastids called apicoplasts. Apicoplasts are not photosynthetic but appear to have a common origin with Dinoflagellate chloroplasts.[1]

W.H.Harvey (1811—1866) was the first to divide 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.[18]

Relationship to higher plants

The first plants on earth probably evolved from shallow freshwater algae much like Chara almost 500 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.[19][20]

Morphology

The kelp forest exhibit at the Monterey Bay Aquarium. A three-dimensional, multicellular thallus.

A range of algal morphologies are exhibited, and convergence of features in unrelated groups is common. The only groups to exhibit three-dimensional multicellular thalli are the reds and browns, and some chlorophytes.[21] Apical growth is constrained to subsets of these groups: the florideophyte reds, various browns, and the charophytes.[21] The form of charophytes is quite different to those of reds and browns, because they have distinct nodes, separated by internode 'stems'; whorls of branches reminiscent of the horsetails occur at the nodes.[21] Conceptacles are another polyphyletic trait; they appear in the coralline algae and the Hildenbrandiales, as well as the browns.[21]

Most of the simpler algae are unicellular flagellates or amoeboids, 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
File:Cyanobacteria Merismopedia.JPG
Cyanobacteria Merismopedia

In three lines even higher levels of organization have been reached, with full tissue differentiation. These are the brown algae,[22]—some of which may reach 50 m in length (kelps)[23]—the red algae,[24] and the green algae.[25] The most complex forms are found among the green algae (see Charales and Charophyta), 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.

Physiology

Many algae, particularly members of the Characeae,[26] have served as model experimental organisms to understand the mechanisms of the water permeability of membranes, osmoregulation, turgor regulation, salt tolerance, cytoplasmic streaming, and the generation of action potentials.

Phytohormones are found not only in higher plants, but in algae too.[27]

Symbiotic algae

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.

Lichens

Main article: Lichens
Rock lichens in Ireland.

Lichens 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."[28] 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.[29] 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.[30] 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.[31]

Coral reefs

Main articles: Coral, Coral reef, and Symbiodinium
Floridian coral reef

Coral reefs are accumulated from the calcareous exoskeletons of marine invertebrates of the order Scleractinia (stony corals). As animals they metabolize sugar and oxygen to obtain energy for their cell-building processes, including secretion of the exoskeleton, with water and carbon dioxide as byproducts. As the reef is the result of a favorable equilibrium between construction by the corals and destruction by marine erosion, the rate at which metabolism can proceed determines the growth or deterioration of the reef.

Dinoflagellates (algal protists) 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 by the host. Stony corals that are reef-building corals (hermatypic corals) require endosymbiotic algae from the genus Symbiodinium to be in a healthy condition.[32] The loss of Symbiodinium from the host is known as coral bleaching, a condition which leads to the deterioration of a reef.

Sea sponges

Main article: Sea sponge

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.[33]

Life-cycle

Rhodophyta, Chlorophyta and Heterokontophyta, 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.[23][34][35]

Further information: Conceptacle

Numbers

Algae on coastal rocks at Shihtiping in Taiwan

The Algal Collection of the US National Herbarium (located in the National Museum of Natural History) consists of approximately 320,500 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).[36] Estimates vary widely. For example, according to one standard textbook,[37] 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"[38]
  • 400 seaweed species for the western coastline of South Africa,[39] and 212 species from the coast of KwaZulu-Natal.[40] Some of these are duplicates as the range extends across both coasts, and the total recorded is probably about 500 species. Most of these are listed in List of seaweeds of South Africa. These exclude phytoplankton and crustose corallines.
  • 669 marine species from California (US)[41]
  • 642 in the check-list of Britain and Ireland[42]

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 microscopic algae, such as phytoplankton.

The most recent estimate suggests a total number of 72,500 algal species worldwide.[43]

Distribution

The topic of distribution of algal species has been fairly well studied since the founding of phytogeography in the mid-19th century AD.[44] 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 of where the spore lands.

The spores of fresh-water algae are dispersed mainly by running water and wind, as well as by living carriers.[45] 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.[46]

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."[47] For example, Clathromorphum is an arctic genus and is not mapped far south of there.[48] On the other hand, scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."[49]

Phytoplankton, Lake Chuzenji

Locations

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).[50]

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.

Cultural associations

In Classical Chinese, the word 藻 is used both for "algae" and (in the modest tradition of the imperial scholars) for "literary talent". The third island in Kunming Lake beside the Summer Palace in Beijing is known as the Zaojian Tang Dao which thus simultaneously means "Island of the Algae-Viewing Hall" and "Island of the Hall for Reflecting on Literary Talent".

Uses

Harvesting algae

Agar

Agar, a gelatinous substance derived from red algae, has a number of commercial uses.[51] It is a good medium on which to grow bacteria and fungi as most microorganisms cannot digest agar.

Alginates

Between 100,000 and 170,000 wet tons of Macrocystis are harvested annually in New Mexico for alginate extraction and abalone feed.[52][53]

Energy source

Main articles: Algae fuel, Biological hydrogen production, Biohydrogen, Biodiesel, Ethanol fuel, Butanol fuel, and Vegetable fats and oils

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 is estimated to occur by 2025.[54]

Fertilizer

Seaweed is used as a fertilizer.
Further information: Seaweed fertiliser

For centuries seaweed has been used as a fertilizer; George Owen of Henllys writing in the 16th century referring to drift weed in South Wales:[55]

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.[56] 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.

Nutrition

Seaweed gardens on Inisheer.
See also: Edible seaweed

Naturally growing seaweeds are an important source of food, especially in Asia. They provide many vitamins including: A, B1, B2, B6, niacin and C, and are rich in iodine, potassium, iron, magnesium and calcium.[57] In addition commercially cultivated microalgae, including both algae and cyanobacteria, are marketed as nutritional supplements, such as Spirulina,[58] Chlorella 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;[59] Ireland, dulse; Chile, cochayuyo.[60] Laver is used to make "laver bread" in Wales where it is known as bara lawr; in Korea, gim; in Japan, nori and aonori. 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 and badderlocks are a salad ingredient in Scotland, Ireland, Greenland and Iceland.

Dulse, a type of food.

The oils from some algae have high levels of unsaturated fatty acids. For example, Parietochloris incisa is very high in arachidonic acid, where it reaches up to 47% of the triglyceride pool.[61] Some varieties of algae favored by vegetarianism and veganism contain the long-chain, essential omega-3 fatty acids, Docosahexaenoic acid (DHA) and Eicosapentaenoic acid (EPA). Fish oil contains the omega-3 fatty acids, but the original source is algae (microalgae in particular), which are eaten by marine life such as copepods and are passed up the food chain.[62] Algae has emerged in recent years as a popular source of omega-3 fatty acids for vegetarians who cannot get long-chain EPA and DHA from other vegetarian sources such as flaxseed oil, which only contains the short-chain Alpha-Linolenic acid (ALA).

Pollution control

  • Sewage can be treated with algae, reducing the usage of large amounts of toxic chemicals that would otherwise be needed.
  • Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested, the enriched algae itself can be used as fertilizer.
  • Aquariums and ponds can be filtered using algae, which absorb nutrients from the water in a device called an algae scrubber, also known as an "ATS".[63][64][65][66]

Agricultural Research Service scientists found that 60-90% of nitrogen runoff and 70-100% of phosphorus runoff can be captured from manure effluents using a horizontal algae scrubber, also called an algal turf scrubber (ATS). Scientists developed the ATS, which are shallow, 100-foot raceways of nylon netting where algae colonies can form, and studied its efficacy for three years. They found that algae can readily be used to reduce the nutrient runoff from agricultural fields and increase the quality of water flowing into rivers, streams, and oceans. The enriched algae itself also can be used as a fertilizer. Researchers collected and dried the nutrient-rich algae from the ATS and studied its potential as an organic fertilizer. They found that cucumber and corn seedlings grew just as well using ATS organic fertilizer as they did with commercial fertilizers.[67] Algae scrubbers, using bubbling upflow or vertical waterfall versions, are now also being used to filter aquariums and ponds.

Bioremediation

The algae Stichococcus bacillaris, has been seen to colonize silcone resins used at archaeological sites; biodeterriorating the synthetic substance.[68]

Pigments

The natural pigments produced by algae can be used as an alternative to chemical dyes and coloring agents.[69]

Stabilizing substances

Main articles: Carrageenan and Chondrus crispus

Carrageenan, from the red alga Chondrus crispus, is used as a stabilizer in milk products.

See also

References

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  32. ^ Taylor, Dennis L (1983). "The coral-algal symbiosis". In Goff, Lynda J (ed.). Algal Symbiosis: A Continuum of Interaction Strategies. CUP Archive. pp. 19–20. ISBN 978-0-521-25541-7.
  33. ^ http://uwsp.edu/cnr/UWEXlakes/laketides/vol26-4/vol26-4.pdf
  34. ^ Christopher S. Lobban; Paul J. Harrison (1994). Seaweed Ecology and Physiology. Cambridge : Cambridge University Press. ISBN 978-0-521-40897-4.
  35. ^ Algae II
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  37. ^ John (2002), page 1.
  38. ^ Huisman (2000), page 25.
  39. ^ Stegenga (1997).
  40. ^ Clerck, Olivier (2005). Guide to the seaweeds of KwaZulu-Natal. ISBN 978-90-72619-64-8.
  41. ^ Abbott and Hollenberg (1976), page 2.
  42. ^ Hardy and Guiry (2006).
  43. ^ Guiry, M. D. (2012). "HOW MANY SPECIES OF ALGAE ARE THERE?". Journal of Phycology. 48 (5): 1057–1063. doi:10.1111/j.1529-8817.2012.01222.x.
  44. ^ Round (1981), Chapter 8, Dispersal, continuity and phytogeography.
  45. ^ Round (1981), page 360.
  46. ^ Round (1981), page 362.
  47. ^ Round (1981), Page 357.
  48. ^ Round (1981), page 371.
  49. ^ Round (1981), page 366.
  50. ^ Round (1981), page 176.
  51. ^ Lewis, J G; Stanley, N F; Guist, G G (1988). "9 Commercial production of algal hydrocolloides". In Lembi, C.A.; Waaland, J.R. (eds.). Algae and Human Affairs. Cambridge: Cambridge University Press. ISBN 978-0-521-32115-0.
  52. ^ "Macrocystis C. Agardh 1820: 46". AlgaeBase. Archived from the original on 4 January 2009. Retrieved 28 December 2008. {{cite web}}: Unknown parameter |deadurl= ignored (|url-status= suggested) (help)
  53. ^ "Secondary Products of Brown Algae". Algae Research. Smithsonian National Museum of Natural History. Retrieved 29 December 2008.
  54. ^ An Outlook on Microalgal Biofuels René H. Wijffels and Maria J. Barbosa Science 13 August 2010: 329 (5993), 796-799. [DOI:10.1126/science.1189003]
  55. ^ Read, Clare Sewell (1849). "On the Farming of South Wales: Prize Report". Journal of the Royal Agricultural Society of England. 10. London: John Murray: 142–143. Downloadable Google Books.
  56. ^ McHugh, Dennis J. (2003). "9, Other Uses of Seaweeds". A Guide to the Seaweed Industry: FAO Fisheries Technical Paper 441. Rome: Fisheries and Aquaculture Department, Food and Agriculture Organization (FAO) of the United Nations. ISBN 978-92-5-104958-7.
  57. ^ Simoons, Frederick J (1991). "6, Seaweeds and Other Algae". Food in China: A Cultural and Historical Inquiry. CRC Press. pp. 179–190. ISBN 978-0-936923-29-1.
  58. ^ Morton, Steve L. "Modern Uses of Cultivated Algae". Ethnobotanical Leaflets. Southern Illinois University Carbondale. Archived from the original on 23 December 2008. Retrieved 26 December 2008.
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Bibliography

General

  • Chapman, V.J. (1950). Seaweeds and their Uses. London: Methuen & Co. Ltd. ISBN 978-0-412-15740-0.
  • Fritsch, F.E. (1945). The Structure and Reproduction of the Algae. I. and II. Cambridge, England: Cambridge University Press
  • van den Hoek, C., D.G. Mann, and H.M. Jahns (1995). Algae: an introduction to phycology. Cambridge University Press (623 pp).
  • Lembi, C.A.; Waaland, J.R. (1988). Algae and Human Affairs. Cambridge: Cambridge University Press. ISBN 978-0-521-32115-0.
  • Mumford, T F; Miura, A (1988). "Porphyra as food: cultivation and economic". In Lembi, C A; Waaland, J R (eds.). Algae and Human Affairs. Cambridge University Press. pp. 87–117. ISBN 978-0-521-32115-0..
  • Round, F E (1981). The Ecology of Algae. London: Cambridge University Press. ISBN 978-0-521-22583-0.
  • Smith, G.M. (1938). Cryptogamic Botany, vol. 1. McGraw-Hill, New York.

Regional

Britain and Ireland
  • Brodie, Juliet; Burrows, Elsie M; Chamberlain, Yvonne M.; Christensen, Tyge; Dixon, Peter Stanley; Fletcher, R.L.; Hommersand, Max H; Irvine, Linda M; Maggs, Christine A (1977–2003). Seaweeds of the British Isles: A Collaborative Project of the British Phycological Society and the British Museum (Natural History). London, Andover: British Museum (Natural History), HMSO, Intercept. ISBN 978-0-565-00781-2. {{cite book}}: Unknown parameter |displayauthors= ignored (|display-authors= suggested) (help)
  • Cullinane, John P (1973). Phycology of the South Coast of Ireland. Cork: Cork University Press.
  • Hardy, F G; Aspinall, R J (1988). An Atlas of the Seaweeds of Northumberland and Durham. The Hancock Museum, University Newcastle upon Tyne: Northumberland Biological Records Centre. ISBN 978-0-9509680-5-6.
  • Hardy, F G; Guiry, Michael D; Arnold, Henry R (2006). A Check-list and Atlas of the Seaweeds of Britain and Ireland (Revised ed.). London: British Phycological Society. ISBN 978-3-906166-35-3.
  • John, D M; Whitton, B A; Brook, J A (2002). The Freshwater Algal Flora of the British Isles. Cambridge, UK; New York: Cambridge University Press. ISBN 978-0-521-77051-4.
  • Knight, Margery; Parke, Mary W (1931). Manx Algae: An Algal Survey of the South End of the Isle of Man. Liverpool Marine Biology Committee (LMBC) Memoirs on Typical British Marine Plants & Animals. Vol. XXX. Liverpool: University Press.
  • Morton, Osborne (1994). Marine Algae of Northern Ireland. Belfast: Ulster Museum. ISBN 978-0-900761-28-7.
  • Morton, Osborne (1 December 2003). "The Marine Macroalgae of County Donegal, Ireland". Bulletin of the Irish Biogeographical Society. 27: 3–164.
Australia
  • Huisman, J M (2000). Marine Plants of Australia. University of Western Australian (UWA) Press. ISBN 978-1-876268-33-6.
New Zealand
  • Chapman, Valentine Jackson; Lindauer, VW; Aiken, M; Dromgoole, FI (1900, 1956, 1961, 1969, 1970). The Marine algae of New Zealand. London; Lehre, Germany: Linnaean Society of London; Cramer. {{cite book}}: Check date values in: |year= (help)CS1 maint: year (link)
Europe
  • Cabioc'h, Jacqueline; Floc'h, Jean-Yves; Le Toquin, Alain; Boudouresque, Charles-François; Meinesz, Alexandre; Verlaque, Marc (1992). Guide des algues des mers d'Europe: Manche/Atlantique-Méditerranée (in French). Lausanne, Suisse: Delachaux et Niestlé. ISBN 978-2-603-00848-5.
  • Gayral, Paulette (1966). Les Algues de côtes françaises (manche et atlantique), notions fondamentales sur l'écologie, la biologie et la systématique des algues marines (in French). Paris: Doin, Deren et Cie.
  • Guiry, M.D.; Blunden, G. (1991). Seaweed Resources in Europe: Uses and Potential. John Wiley & Sons. ISBN 978-0-471-92947-5.
  • Míguez Rodríguez, Luís (1998). Algas mariñas de Galicia: bioloxía, gastronomía, industria (in Galician). Vigo: Edicións Xerais de Galicia. ISBN 978-84-8302-263-4.{{cite book}}: CS1 maint: unrecognized language (link)
  • Otero, J. (2002). Guía das macroalgas de Galicia (in Galician). A Coruña: Baía Edicións. ISBN 978-84-89803-22-0.{{cite book}}: CS1 maint: unrecognized language (link)
  • Bárbara, I.; Cremades, J. (1993). Guía de las algas del litoral gallego (in Spanish). A Coruña: Concello da Coruña - Casa das Ciencias.
Arctic
  • 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. Vol. 20. Stockholm: Kungl. Svenska vetenskapsakademiens handlingar. pp. 1–350.
Greenland
  • Lund, Søren Jensen (1959). The Marine Algae of East Greenland. Kövenhavn: C.A. Reitzel. 9584734.
Faroe Islands
  • Børgesen, Frederik (1903, 1970 reprint). "Marine Algae". In Warming, Eugene (ed.). Botany of the Faröes Based Upon Danish Investigations. Part II. København: Det nordiske Forlag. pp. 339–532. {{cite book}}: Check date values in: |year= (help)CS1 maint: year (link).
Canary Islands
  • Børgesen, Frederik (1925, 1926, 1927, 1929, 1930, 1936). Marine Algae from the Canary Islands. København: Bianco Lunos. {{cite book}}: Check date values in: |year= (help)CS1 maint: year (link)
Morocco
  • Gayral, Paulette (1958). Algues de la côte atlantique marocaine (in French). 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 978-0-7992-1793-3.
North America
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