Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of

multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially un ...

algae Algae ( , ; : alga ) are any of a large and diverse group of photosynthetic, eukaryotic organisms. The name is an informal term for a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from ...

, including many seaweeds located in colder waters within the Northern Hemisphere. Brown

are the major seaweeds of the temperate and polar regions. They are dominant on rocky shores throughout cooler areas of the world. Most brown algae live in marine environments, where they play an important role both as food and as a potential

habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...

. For instance, ''

Macrocystis ''Macrocystis'' is a monospecific genus of kelp (large brown algae). This genus contains the largest of all the phaeophyceae or brown algae. ''Macrocystis'' has pneumatocysts at the base of its blades. Sporophytes are perennial and the indiv ...

'', a

kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "under ...

of the order Laminariales, may reach in length and forms prominent underwater

kelp forest Kelp forests are underwater areas with a high density of kelp, which covers a large part of the world's coastlines. Smaller areas of anchored kelp are called kelp beds. They are recognized as one of the most productive and dynamic ecosystems on E ...

s. Kelp forests like these contain a high level of biodiversity. Another example is ''

Sargassum ''Sargassum'' is a genus of brown (class Phaeophyceae) macroalgae ( seaweed) in the order Fucales. Numerous species are distributed throughout the temperate and tropical oceans of the world, where they generally inhabit shallow water and coral ...

'', which creates unique floating mats of seaweed in the tropical waters of the

Sargasso Sea The Sargasso Sea () is a region of the Atlantic Ocean bounded by four currents forming an ocean gyre. Unlike all other regions called seas, it has no land boundaries. It is distinguished from other parts of the Atlantic Ocean by its char ...

that serve as the habitats for many species. Many brown algae, such as members of the order Fucales, commonly grow along rocky seashores. Some members of the class, such as kelps, are used by humans as food. Between 1,500 and 2,000

species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...

of brown algae are known worldwide. Some species, such as ''

Ascophyllum nodosum ''Ascophyllum nodosum'' is a large, common cold water seaweed or brown alga ( Phaeophyceae) in the family Fucaceae, being the only species in the genus ''Ascophyllum''. It is a seaweed that only grows in the northern Atlantic Ocean, also known ...

'', have become subjects of extensive research in their own right due to their commercial importance. They also have environmental significance through

carbon fixation Biological carbon fixation or сarbon assimilation is the process by which inorganic carbon (particularly in the form of carbon dioxide) is converted to organic compounds by living organisms. The compounds are then used to store energy and as ...

. Brown algae belong to the group

Heterokont Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which a ...

ophyta, a large group of

eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bact ...

organisms distinguished most prominently by having

chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it ...

s surrounded by four membranes, suggesting an origin from a

symbiotic relationship Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...

between a basal eukaryote and another eukaryotic organism. Most brown algae contain the pigment

fucoxanthin Fucoxanthin is a xanthophyll, with formula C42H58O6. It is found as an accessory pigment in the chloroplasts of brown algae and most other heterokonts, giving them a brown or olive-green color. Fucoxanthin absorbs light primarily in the blue-green ...

, which is responsible for the distinctive greenish-brown color that gives them their name. Brown algae are unique among heterokonts in developing into multicellular forms with differentiated tissues, but they reproduce by means of flagellated

spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...

s and

gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...

s that closely resemble cells of other heterokonts. Genetic studies show their closest relatives to be the yellow-green algae.

Morphology

Brown algae exist in a wide range of sizes and forms. The smallest members of the group grow as tiny, feathery tufts of threadlike cells no more than a few centimeters (a few inches) long. Some species have a stage in their life cycle that consists of only a few cells, making the entire alga microscopic. Other groups of brown algae grow to much larger sizes. The rockweeds and leathery

s are often the most conspicuous algae in their habitats. Kelps can range in size from the sea palm '' Postelsia'' to the giant kelp ''

Macrocystis pyrifera ''Macrocystis pyrifera'', commonly known as giant kelp or bladder kelp, is a species of kelp (large brown algae), and one of four species in the genus '' Macrocystis''. Despite its appearance, it is not a plant; it is a heterokont. Giant kelp ...

'', which grows to over long and is the largest of all the algae. In form, the brown algae range from small crusts or cushions to leafy free-floating mats formed by species of ''

''. They may consist of delicate felt-like strands of cells, as in '' Ectocarpus'', or of flattened branches resembling a fan, as in '' Padina''. Regardless of size or form, two visible features set the Phaeophyceae apart from all other algae. First, members of the group possess a characteristic color that ranges from an olive green to various shades of

brown Brown is a color. It can be considered a composite color, but it is mainly a darker shade of orange. In the CMYK color model used in printing or painting, brown is usually made by combining the colors orange and black. In the RGB color model use ...

. The particular shade depends upon the amount of

present in the alga. Second, all brown algae are

. There are no known species that exist as single cells or as colonies of cells, and the brown algae are the only major group of

seaweed Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of ''Rhodophyta'' (red), ''Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ke ...

s that does not include such forms. However, this may be the result of classification rather than a consequence of evolution, as all the groups hypothesized to be the closest relatives of the browns include single-celled or colonial forms. They can change color depending on salinity, ranging from reddish to brown.

Visible structures

Whatever their form, the body of all brown algae is termed a ''

thallus Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or " twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms ...

'', indicating that it lacks the complex

xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived fr ...

and

phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...

vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...

s. This does not mean that brown algae completely lack specialized structures. But, because some botanists define "true" stems, leaves, and roots by the presence of these tissues, their absence in the brown algae means that the stem-like and leaf-like structures found in some groups of brown algae must be described using different terminology. Although not all brown algae are structurally complex, those that are typically possess one or more characteristic parts. A '' holdfast'' is a rootlike structure present at the base of the alga. Like a root system in plants, a holdfast serves to anchor the alga in place on the ''substrate'' where it grows, and thus prevents the alga from being carried away by the current. Unlike a root system, the holdfast generally does not serve as the primary organ for water uptake, nor does it take in nutrients from the substrate. The overall physical appearance of the holdfast differs among various brown algae and among various substrates. It may be heavily branched, or it may be cup-like in appearance. A single alga typically has just one holdfast, although some species have more than one stipe growing from their holdfast. A '' stipe'' is a stalk or stemlike structure present in an alga. It may grow as a short structure near the base of the alga (as in '' Laminaria''), or it may develop into a large, complex structure running throughout the algal body (as in ''

'' or ''

''). In the most structurally differentiated brown algae (such as '' Fucus''), the tissues within the stipe are divided into three distinct layers or regions. These regions include a central pith, a surrounding cortex, and an outer epidermis, each of which has an analog in the stem of a vascular plant. In some brown algae, the pith region includes a core of elongated cells that resemble the

of vascular plants both in structure and function. In others (such as ''

Nereocystis ''Nereocystis'' (Greek, 'mermaid's bladder') is a monotypic genus of subtidal kelp containing the species ''Nereocystis luetkeana''. Some English names include edible kelp, bull kelp, bullwhip kelp, ribbon kelp, bladder wrack, and variations of ...

''), the center of the stipe is hollow and filled with gas that serves to keep that part of the alga buoyant. The stipe may be relatively flexible and elastic in species like ''

'' that grow in strong currents, or may be more rigid in species like ''

Postelsia palmaeformis ''Postelsia palmaeformis'', also known as the sea palm (not to be confused with the southern sea palm) or palm seaweed, is a species of kelp and classified within brown algae. It is the only known species in the genus Postelsia. The sea palm is ...

'' that are exposed to the atmosphere at low tide. Many algae have a flattened portion that may resemble a leaf, and this is termed a ''blade'', '' lamina'', or ''frond''. The name ''blade'' is most often applied to a single undivided structure, while ''frond'' may be applied to all or most of an algal body that is flattened, but this distinction is not universally applied. The name ''lamina'' refers to that portion of a structurally differentiated alga that is flattened. It may be a single or a divided structure, and may be spread over a substantial portion of the alga. In rockweeds, for example, the lamina is a broad wing of tissue that runs continuously along both sides of a branched ''midrib''. The midrib and lamina together constitute almost all of a rockweed, so that the lamina is spread throughout the alga rather than existing as a localized portion of it. Bladder Wrack (Fucus vesiculosus) - geograph

Bladder Wrack (Fucus vesiculosus) - geograph

In some brown algae, there is a single lamina or blade, while in others there may be many separate blades. Even in those species that initially produce a single blade, the structure may tear with rough currents or as part of maturation to form additional blades. These blades may be attached directly to the stipe, to a holdfast with no stipe present, or there may be an air bladder between the stipe and blade. The surface of the lamina or blade may be smooth or wrinkled; its tissues may be thin and flexible or thick and leathery. In species like '' Egregia menziesii'', this characteristic may change depending upon the turbulence of the waters in which it grows. In other species, the surface of the blade is coated with slime to discourage the attachment of epiphytes or to deter

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthp ...

s. Blades are also often the parts of the alga that bear the reproductive structures. Gas-filled floats called '' pneumatocysts'' provide

buoyancy Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the p ...

in many

s and members of the Fucales. These bladder-like structures occur in or near the ''lamina'', so that it is held nearer the water surface and thus receives more light for photosynthesis. Pneumatocysts are most often spherical or

ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface; that is, a surface that may be defined as the ...

al, but can vary in shape among different species. Species such as '' Nereocystis luetkeana'' and '' Pelagophycus porra'' bear a single large pneumatocyst between the top of the stipe and the base of the blades. In contrast, the giant kelp ''Macrocystis pyrifera'' bears many blades along its stipe, with a pneumatocyst at the base of each blade where it attaches to the main stipe. Species of ''

'' also bear many blades and pneumatocysts, but both kinds of structures are attached separately to the stipe by short stalks. In species of '' Fucus'', the pneumatocysts develop within the lamina itself, either as discrete spherical bladders or as elongated gas-filled regions that take the outline of the lamina in which they develop.

Growth

The brown algae include the largest and fastest growing of seaweeds. Fronds of ''

'' may grow as much as per day, and the stipes can grow in a single day. Growth in most brown algae occurs at the tips of structures as a result of divisions in a single ''apical cell'' or in a row of such cells. They are single cellular organisms. As this apical cell divides, the new cells that it produces develop into all the tissues of the alga. Branchings and other lateral structures appear when the apical cell divides to produce two new apical cells. However, a few groups (such as '' Ectocarpus'') grow by a diffuse, unlocalized production of new cells that can occur anywhere on the thallus.

Tissue organization

The simplest browns are filamentous—that is, their cells are elongate and have septa cutting across their width. They branch by getting wider at their tip, and then dividing the widening. These filaments may be haplostichous or polystichous, multiaxial or monoaxial forming or not
pseudoparenchyma
Besides fronds, there are the large in size

parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word π ...

tic kelps with three-dimensional development and growth and different tissues ( meristoderm, cortex and medulla) which could be consider the trees of the sea. There are also the Fucales and Dictyotales smaller than kelps but still parenchymatic with the same kind of distinct tissues. The

cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...

consists of two layers; the inner layer bears the strength, and consists of

cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...

; the outer wall layer is mainly algin, and is gummy when wet but becomes hard and brittle when it dries out. Specifically, the brown algal cell wall consists of several components with alginates and sulphated fucan being its main ingredients, up to 40% each of them. Cellulose, a major component from most plant cell walls, is present in a very small percentage, up to 8%. Cellulose and alginate biosynthesis pathways seem to have been acquired from other organisms through endosymbiotic and horizontal gene transfer respectively, while the sulphated polysaccharides are of ancestral origin. Specifically, the cellulose synthases seem to come from the red alga endosymbiont of the photosynthetic stramenopiles ancestor, and the ancestor of brown algae acquired the key enzymes for alginates biosynthesis from an

actinobacterium The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to ...

. The presence and fine control of alginate structure in combination with the cellulose which existed before it, gave potentially the brown algae the ability to develop complex structurally multicellular organisms like the kelps.

Evolutionary history

Genetic and ultrastructural evidence place the Phaeophyceae among the

heterokont Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which a ...

s (Stramenopiles), a large assemblage of organisms that includes both photosynthetic members with

plastid The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosymbiotic cyanobac ...

s (such as the diatoms) as well as non-photosynthetic groups (such as the slime nets and water molds). Although some heterokont relatives of the brown algae lack plastids in their cells, scientists believe this is a result of evolutionary loss of that organelle in those groups rather than independent acquisition by the several photosynthetic members. Thus, all heterokonts are believed to descend from a single

heterotrophic A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...

ancestor that became photosynthetic when it acquired plastids through

endosymbiosis An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "withi ...

of another unicellular eukaryote. The closest relatives of the brown algae include unicellular and filamentous species, but no unicellular species of brown algae are known. However, most scientists assume that the Phaeophyceae evolved from unicellular ancestors. DNA sequence comparison also suggests that the brown algae evolved from the filamentous Phaeothamniophyceae,

Xanthophyceae Yellow-green algae or the Xanthophyceae (xanthophytes) are an important group of heterokont algae. Most live in fresh water, but some are found in marine and soil habitats. They vary from single-celled flagellates to simple colonial and filamen ...

, or the Chrysophyceae between 150 and 200 million years ago. In many ways, the evolution of the brown algae parallels that of the

green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ( Embryophytes) have emerged deep in the Charophyte alg ...

and red algae, as all three groups possess complex multicellular species with an alternation of generations. Analysis of 5S

rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from riboso ...

sequences reveals much smaller evolutionary distances among genera of the brown algae than among genera of red or green algae, which suggests that the brown algae have diversified much more recently than the other two groups.

Fossils

The occurrence of Phaeophyceae as

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

s is rare due to their generally soft-bodied nature, and scientists continue to debate the identification of some finds. Part of the problem with identification lies in the

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...

of morphologies between many brown and red algae. Most fossils of soft-tissue algae preserve only a flattened outline, without the microscopic features that permit the major groups of multicellular algae to be reliably distinguished. Among the brown algae, only species of the genus '' Padina'' deposit significant quantities of minerals in or around their cell walls. Other algal groups, such as the red algae and

, have a number of

calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an a ...

members. Because of this, they are more likely to leave evidence in the fossil record than the soft bodies of most brown algae and more often can be precisely classified. Fossils comparable in morphology to brown algae are known from strata as old as the Upper

Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. ...

, but the taxonomic affinity of these impression fossils is far from certain. Claims that earlier

Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and t ...

fossils are brown algae have since been dismissed. While many

carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon makes ...

aceous fossils have been described from the

Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of th ...

, they are typically preserved as flattened outlines or fragments measuring only millimeters long. Because these fossils lack features diagnostic for identification at even the highest level, they are assigned to fossil form taxa according to their shape and other gross morphological features. A number of

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, w ...

fossils termed ''fucoids'', from their resemblance in outline to species in the genus '' Fucus'', have proven to be inorganic rather than true fossils. The Devonian megafossil '' Prototaxites'', which consists of masses of filaments grouped into trunk-like axes, has been considered a possible brown alga. However, modern research favors reinterpretation of this fossil as a terrestrial

fungus A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...

or fungal-like organism. Likewise, the fossil '' Protosalvinia'' was once considered a possible brown alga, but is now thought to be an early

land plant The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sist ...

. A number of

Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...

fossils have been tentatively classified with the brown algae, although most have also been compared to known red algae species. '' Phascolophyllaphycus'' possesses numerous elongate, inflated blades attached to a stipe. It is the most abundant of algal fossils found in a collection made from Carboniferous strata in

Illinois Illinois ( ) is a state in the Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolitan areas include, Peoria and Roc ...

. Each hollow blade bears up to eight pneumatocysts at its base, and the stipes appear to have been hollow and inflated as well. This combination of characteristics is similar to certain modern genera in the order Laminariales (kelps). Several fossils of '' Drydenia'' and a single specimen of '' Hungerfordia'' from the Upper Devonian of

New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...

have also been compared to both brown and red algae. Fossils of ''Drydenia'' consist of an elliptical blade attached to a branching filamentous holdfast, not unlike some species of '' Laminaria'', ''

Porphyra ''Porphyra'' is a genus of coldwater seaweeds that grow in cold, shallow seawater. More specifically, it belongs to red algae phylum of laver species (from which comes laverbread), comprising approximately 70 species.Brodie, J.A. and Irvine ...

'', or '' Gigartina''. The single known specimen of ''Hungerfordia'' branches dichotomously into lobes and resembles genera like '' Chondrus'' and '' Fucus'' or '' Dictyota''. The earliest known fossils that can be assigned reliably to the Phaeophyceae come from

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...

diatomite deposits of the Monterey Formation in

California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...

. Several soft-bodied brown macroalgae, such as '' Julescraneia'', have been found.

Classification

Phylogeny

Based on the work of Silberfeld, Rousseau & de Reviers 2014.

Taxonomy

This is a list of the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

s in the class Phaeophyceae: * Class Phaeophyceae Hansgirg 1886 ucophyceae; Melanophycidae Rabenhorst 1863 stat. nov. Cavalier-Smith 2006** Subclass Discosporangiophycidae Silberfeld, Rousseau & Reviers 2014 *** Order Discosporangiales Schmidt 1937 emend. Kawai et al. 2007 **** Family Choristocarpaceae Kjellman 1891 **** Family Discosporangiaceae Schmidt 1937 ** Subclass

Ishigeophycidae Ishigeales is an order of brown algae. It includes two families, Ishigeaceae and Petrodermataceae. The genus ''Diplura The order Diplura ("two-pronged bristletails") is one of three orders of non-insect hexapods within the class Entognath ...

Silberfeld, Rousseau & Reviers 2014 *** Order Ishigeales Cho & Boo 2004 **** Family Ishigeaceae Okamura 1935 **** Family Petrodermataceae Silberfeld, Rousseau & Reviers 2014 ** Subclass Dictyotophycidae Silberfeld, Rousseau & Reviers 2014 *** Order Dictyotales Bory de Saint-Vincent 1828 ''ex'' Phillips ''et al.'' **** Family Dictyotaceae Lamouroux ex Dumortier 1822 coresbyellaceae Womersley 1987; Dictyopsidaceae*** Order Onslowiales Draisma & Prud’homme van Reine 2008 **** Family Onslowiaceae Draisma & Prud’homme van Reine 2001 *** Order

Sphacelariales Sphacelariales is an order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potent ...

Migula 1909 **** Family

Cladostephaceae Cladostephaceae is a family of brown algae belonging to the order Sphacelariales in the class Phaeophyceae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds locat ...

Oltmanns 1922 **** Family Lithodermataceae Hauck 1883 **** Family Phaeostrophiaceae Kawai et al. 2005 **** Family

Sphacelariaceae Sphacelariaceae is a family of algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Includ ...

Decaisne 1842 **** Family Sphacelodermaceae Draisma, Prud’homme & Kawai 2010 **** Family Stypocaulaceae Oltmanns 1922 *** Order

Syringodermatales Syringodermataceae is a family of brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown alg ...

Henry 1984 **** Family

Syringodermataceae Syringodermataceae is a family of brown algae. It includes two genera, '' Microzonia'' and ''Syringoderma ''Syringoderma'' is a genus in the family Syringodermataceae of the brown algae (class Phaeophyceae). The genus contains four species ...

Henry 1984 ** Subclass Fucophycidae Cavalier-Smith 1986 *** Order

Ascoseirales ''Ascoseira'' is a monotypic genus of seaweed in the brown algae (class Phaeophyceae). The single and type species, ''Ascoseira mirabilis'' Skottsberg, is a large parenchymatous macroalgae, and is endemic to the Antarctic Ocean. ''Ascoseira'' is ...

Petrov1964 emend. Moe & Henry 1982 **** Family Ascoseiraceae Skottsberg 1907 *** Order Asterocladales T.Silberfeld et al. 2011 **** Family Asterocladaceae Silberfeld et al. 2011 *** Order Desmarestiales Setchell & Gardner 1925 **** Family Arthrocladiaceae Chauvin 1842 **** Family Desmarestiaceae (Thuret) Kjellman 1880 *** Order

Ectocarpales Ectocarpales is a very large order in the brown algae (class Phaeophyceae). The order includes families with pseudoparenchymatous ( Splachnidiaceae) or true parenchymatous ( Scytosiphonaceae) tissue. Pseudoparenchymatous refers to a filamento ...

Bessey 1907 emend. Rousseau & Reviers 1999a hordariales Setchell & Gardner 1925; Dictyosiphonales Setchell & Gardner 1925; Scytosiphonales Feldmann 1949**** Family

Acinetosporaceae Acinetosporaceae is a family of brown algae in the order Ectocarpales Ectocarpales is a very large order in the brown algae (class Phaeophyceae). The order includes families with pseudoparenchymatous ( Splachnidiaceae) or true parenchymatou ...

Hamel ex Feldmann 1937 ylaiellaceae; Pilayellaceae**** Family Adenocystaceae Rousseau et al. 2000 emend. Silberfeld et al. 2011 hordariopsidaceae**** Family

Chordariaceae Chordariaceae is a family of brown algae. Members of this family are may be filamentous, crustose with fused cells at the base, or they may be terete and differentiated into a central medulla and an outer photosynthetic cortex. They have a sp ...

Greville 1830 emend. Peters & Ramírez 2001 yrionemataceae**** Family

Ectocarpaceae Ectocarpaceae is a family of brown algae in the order Ectocarpales Ectocarpales is a very large order in the brown algae (class Phaeophyceae). The order includes families with pseudoparenchymatous ( Splachnidiaceae) or true parenchymatous ( ...

Agardh 1828 emend. Silberfeld et al. 2011 **** Family Petrospongiaceae Racault et al. 2009 **** Family

Scytosiphonaceae Scytosiphonaceae is a family of brown algae in the order Ectocarpales. Genera: References Ectocarpales Brown algae families {{Phaeophyceae-stub ...

Ardissone & Straforello 1877 hnoosporaceae Setchell & Gardner 1925*** Order Fucales Bory de Saint-Vincent 1827 otheiales Womersley 1987; Durvillaeales Petrov 1965**** Family Bifurcariopsidaceae Cho et al. 2006 **** Family Durvillaeaceae (Oltmanns) De Toni 1891 **** Family Fucaceae Adanson 1763 **** Family Himanthaliaceae (Kjellman) De Toni 1891 **** Family Hormosiraceae Fritsch 1945 **** Family Notheiaceae Schmidt 1938 **** Family

Sargassaceae Sargassaceae is a family of brown algae in the order Fucales. See also *''Carpophyllum maschalocarpum'' *''Halidrys siliquosa ''Halidrys siliquosa'' is a large marine brown algae. Description ''Halidrys siliquosa'' is a large brown alga g ...

Kützing 1843 ystoseiraceae De Toni 1891**** Family Seirococcaceae Nizamuddin 1987 **** Family Xiphophoraceae Cho et al. 2006 *** Order Laminariales Migula 1909 haeosiphoniellales Silberfeld, Rousseau & Reviers 2014 ord. nov. prop.**** Family Agaraceae Postels & Ruprecht 1840 ostariaceae**** Family Akkesiphycaceae Kawai & Sasaki 2000 **** Family Alariaceae Setchell & Gardner 1925 **** Family Aureophycaceae Kawai & Ridgway 2013 **** Family Chordaceae Dumortier 1822 **** Family Laminariaceae Bory de Saint-Vincent 1827 rthrothamnaceae Petrov 1974**** Family Lessoniaceae Setchell & Gardner 1925 **** Family Pseudochordaceae Kawai & Kurogi 1985 *** Order

Nemodermatales ''Nemoderma'' is the only genus in the family Nemodermataceae and order Nemodermatales of the brown algae (class Phaeophyceae). The genus contains only a single species In biology, a species is the basic unit of classification and a taxo ...

Parente et al. 2008 **** Family

Nemodermataceae ''Nemoderma'' is the only genus in the family Nemodermataceae and order Nemodermatales of the brown algae (class Phaeophyceae). The genus contains only a single species In biology, a species is the basic unit of classification and a tax ...

Kuckuck ex Feldmann 1937 *** Order Phaeosiphoniellales Silberfeld, Rousseau & Reviers 2014 **** Family Phaeosiphoniellaceae Phillips et al. 2008 *** Order Ralfsiales Nakamura ex Lim & Kawai 2007 **** Family Mesosporaceae Tanaka & Chihara 1982 **** Family Neoralfsiaceae Lim & Kawai 2007 **** Family Ralfsiaceae Farlow 1881 eterochordariaceae Setchell & Gardner 1925*** Order Scytothamnales Peters & Clayton 1998 emend. Silberfeld et al. 2011 **** Family Asteronemataceae Silberfeld et al. 2011 **** Family Bachelotiaceae Silberfeld et al. 2011 **** Family Splachnidiaceae Mitchell & Whitting 1892 cytothamnaceae Womersley 1987*** Order Sporochnales Sauvageau 1926 **** Family

Sporochnaceae Sporochnacaeae is the only family in the order Sporochnales in the brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within ...

Greville 1830 *** Order Tilopteridales Bessey 1907 emend. Phillips et al. 2008 utleriales Bessey 1907**** Family Cutleriaceae Griffith & Henfrey 1856 **** Family Halosiphonaceae Kawai & Sasaki 2000 **** Family Phyllariaceae Tilden 1935 **** Family Stschapoviaceae Kawai 2004 **** Family

Tilopteridaceae Tilopteridaceae is a brown algae family in the order Tilopteridales Tilopteridales is an order of brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweed ...

Kjellman 1890 Laminaria Life Cycle

Life cycle

Most brown algae, with the exception of the Fucales, perform

sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...

through sporic meiosis. Between generations, the algae go through separate sporophyte (

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. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respecti ...

) and

gametophyte A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...

( haploid) phases. The sporophyte stage is often the more visible of the two, though some species of brown algae have similar diploid and haploid phases. Free floating forms of brown algae often do not undergo

until they attach themselves to substrate. The haploid generation consists of male and female

s. The fertilization of egg cells varies between species of brown algae, and may be isogamous, oogamous, or anisogamous. Fertilization may take place in the water with eggs and motile sperm, or within the oogonium itself. Certain species of brown algae can also perform asexual reproduction through the production of motile diploid

zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Diversity Flagella types Zoospores may possess one or m ...

s. These zoospores form in plurilocular

sporangium A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life c ...

, and can mature into the sporophyte phase immediately. In a representative species '' Laminaria'', there is a conspicuous

generation and smaller haploid generations. Meiosis takes place within several unilocular

along the algae's blade, each one forming either haploid male or female

s. The spores are then released from the sporangia and grow to form male and female gametophytes. The female gametophyte produces an egg in the oogonium, and the male gametophyte releases motile sperm that fertilize the egg. The fertilized zygote then grows into the mature diploid sporophyte. In the order Fucales, sexual reproduction is oogamous, and the mature diploid is the only form for each generation. Gametes are formed in specialized conceptacles that occur scattered on both surfaces of the receptacle, the outer portion of the blades of the parent plant. Egg cells and motile sperm are released from separate sacs within the conceptacles of the parent algae, combining in the water to complete fertilization. The fertilized zygote settles onto a surface and then differentiates into a leafy

and a finger-like holdfast. Light regulates differentiation of the zygote into blade and holdfast. Saccharina latissima NOAA

Ecology

Brown algae have adapted to a wide variety of marine ecological niches including the tidal splash zone, rock pools, the whole intertidal zone and relatively deep near shore waters. They are an important constituent of some brackish water ecosystems, and have colonized freshwater on a maximum of six known occasions. A large number of Phaeophyceae are intertidal or upper littoral, and they are predominantly cool and cold water organisms that benefit from nutrients in up welling cold water currents and inflows from land; ''

'' being a prominent exception to this generalisation. Brown algae growing in brackish waters are almost solely asexual.

Chemistry

Brown algae have a value in the range of , in contrast with red algae and greens. This reflects their different metabolic pathways. They have cellulose walls with alginic acid and also contain the polysaccharide fucoidan in the amorphous sections of their cell walls. A few species (of ''Padina'') calcify with

aragonite Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including pre ...

needles. In addition to alginates, fucoidan and cellulose, the carbohydrate composition of brown algae consist of

mannitol Mannitol is a type of sugar alcohol used as a sweetener and medication. It is used as a low calorie sweetener as it is poorly absorbed by the intestines. As a medication, it is used to decrease pressure in the eyes, as in glaucoma, and to low ...

, laminarin and

glucan A glucan is a polysaccharide derived from D-glucose, linked by glycosidic bonds. Glucans are noted in two forms: alpha glucans and beta glucans. Many beta-glucans are medically important. They represent a drug target for antifungal medications ...

. The photosynthetic system of brown algae is made of a P700 complex containing chlorophyll a. Their plastids also contain chlorophyll c and carotenoids (the most widespread of those being

). Brown algae produce a specific type of tannin called phlorotannins in higher amounts than red algae do.

Importance and uses

Brown algae include a number of

edible seaweed Edible seaweed, or sea vegetables, are seaweeds that can be eaten and used for culinary purposes. They typically contain high amounts of fiber. They may belong to one of several groups of multicellular algae: the red algae, green algae, and br ...

s. All brown algae contain

alginic acid Alginic acid, also called algin, is a naturally occurring, edible polysaccharide found in brown algae. It is hydrophilic and forms a viscous gum when hydrated. With metals such as sodium and calcium, its salts are known as alginates. Its colour ...

(alginate) in their cell walls, which is extracted commercially and used as an industrial thickening agent in food and for other uses. One of these products is used in

lithium-ion batteries A lithium-ion or Li-ion battery is a type of rechargeable battery which uses the reversible reduction of lithium ions to store energy. It is the predominant battery type used in portable consumer electronics and electric vehicles. It also see ...

. Alginic acid is used as a stable component of a battery

anode An anode is an electrode of a polarized electrical device through which conventional current enters the device. This contrasts with a cathode, an electrode of the device through which conventional current leaves the device. A common mnemoni ...

. This

polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with ...

is a major component of brown algae, and is not found in land plants. Alginic acid can also be used in

aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lotus ...

. For example, alginic acid enhances the immune system of rainbow trout. Younger fish are more likely to survive when given a diet with alginic acid. Brown algae including

beds also fix a significant portion of the earth's carbon dioxide yearly through photosynthesis. Additionally, they can store a great amount of carbon dioxide which can help us in the fight against climate change. Sargachromanol G, an extract of

Sargassum siliquastrum ''Sargassum siliquastrum'' is a brown alga species in the genus ''Sargassum''. Sargachromanols are bio-active isolates of ''S. siliquastrum''. It has anti-inflammatory effect. See also * List of Sargassum species The genus ''Sargassum'' co ...

, has been shown to have anti-inflammatory effects.

Edible Brown Algae

Kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "under ...
(Laminariales)

* Arame (''Eisenia bicyclis'') * Badderlocks (''

Alaria esculenta ''Alaria esculenta'' is an edible seaweed, also known as dabberlocks or badderlocks, or winged kelp. It is a traditional food along the coasts of the far north Atlantic Ocean. It may be eaten fresh or cooked in Greenland, Iceland, Scotland and I ...

'') * Cochayuyo ('' Durvillaea antarctica'') * '' Ecklonia cava'' * Kombu (''

Saccharina japonica ''Saccharina japonica'' is a marine species of the Phaeophyceae (brown algae) class, a type of kelp or seaweed, which is extensively cultivated on ropes between the seas of China, Japan and Korea. It has the common name sweet kelp. It is wi ...

'') * Oarweed ('' Laminaria digitata'') * Sea palm ''

'' * Sea whip ('' Nereocystis luetkeana'') * Sugar kelp ('' Saccharina latissima'') * Wakame (''Undaria pinnatifida'') * Hiromi (''

Undaria undarioides ''Undaria'' is a genus of kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group o ...

'') * Grapestone (''

Mastocarpus papillatus ''Mastocarpus papillatus'', sometimes called Turkish washcloth, black tar spot, or grapestone is a species of red algae in the family Phyllophoraceae. It is sometimes confused with the distantly related ''Turkish towel'' ('' Chondracanthus ex ...

'')

Fucales

* Bladderwrack ('' Fucus vesiculosus'') * Channelled wrack (''

Pelvetia canaliculata ''Pelvetia canaliculata'', the channelled wrack, is a very common brown alga (Phaeophyceae) found on the rocks of the upper shores of Europe. It is the only species remaining in the monotypic genus ''Pelvetia''. In 1999, the other members of th ...

'') * Hijiki or Hiziki (''Sargassum fusiforme'') * Limu Kala (''

Sargassum echinocarpum ''Sargassum'' is a genus of brown (class Phaeophyceae) macroalgae ( seaweed) in the order Fucales. Numerous species are distributed throughout the temperate and tropical oceans of the world, where they generally inhabit shallow water and coral ...

'') * ''

'' ** '' Sargassum cinetum'' ** ''

Sargassum vulgare ''Sargassum'' is a genus of brown (class Phaeophyceae) macroalgae (seaweed) in the order Fucales. Numerous species are distributed throughout the temperate and tropical oceans of the world, where they generally inhabit shallow water and coral re ...

'' ** '' Sargassum swartzii'' ** ''

Sargassum myriocysum ''Sargassum'' is a genus of brown (class Phaeophyceae) macroalgae (seaweed) in the order Fucales. Numerous species are distributed throughout the temperate and tropical oceans of the world, where they generally inhabit shallow water and coral re ...

'' * Spiral wrack ('' Fucus spiralis'') * Thongweed ('' Himanthalia elongata'')

Ectocarpales Ectocarpales is a very large order in the brown algae (class Phaeophyceae). The order includes families with pseudoparenchymatous ( Splachnidiaceae) or true parenchymatous ( Scytosiphonaceae) tissue. Pseudoparenchymatous refers to a filamento ...

* Mozuku (''

Cladosiphon okamuranus is a type of edible seaweed in the genus '' Cladosiphon'', naturally found in Okinawa, Japan. Most mozuku is now farmed by locals, and sold to processing factories. The main use of mozuku is as food, and as a source of one type of sulfated ...

'')

References

External links

Monterey Bay Flora

University of California Museum of Paleontology The University of California Museum of Paleontology (UCMP) is a paleontology museum located on the campus of the University of California, Berkeley. The museum is within the Valley Life Sciences Building (VLSB), designed by George W. Kelham and ...

Phaeophyceae

National University of Ireland, Galway The University of Galway ( ga, Ollscoil na Gaillimhe) is a public research university located in the city of Galway, Ireland. A tertiary education and research institution, the university was awarded the full five QS stars for excellence in 201 ...

{{Authority control Biological oceanography Tithonian first appearances Extant Late Jurassic first appearances

Morphology

Visible structures

Growth

Tissue organization

Evolutionary history

Fossils

Classification

Phylogeny

Taxonomy

Life cycle

Ecology

Chemistry

Importance and uses

Edible Brown Algae

Kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "under ... (Laminariales)

Fucales

Ectocarpales Ectocarpales is a very large order in the brown algae (class Phaeophyceae). The order includes families with pseudoparenchymatous ( Splachnidiaceae) or true parenchymatous ( Scytosiphonaceae) tissue. Pseudoparenchymatous refers to a filamento ...

See also

References

External links

Kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "under ...
(Laminariales)