Brown algae

Brown algae
Temporal range: Late Jurassic to present 150–0 Ma[1][2]
Giant kelp (Macrocystis pyrifera)
Giant kelp (Macrocystis pyrifera)
Scientific classification e
Kingdom:Chromista
Phylum:Ochrophyta
Superclass:Fucistia
Class:Phaeophyceae
Kjellman, 1891[3]
Orders

see Classification.

Synonyms

The 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. Most brown algae live in marine environments, where they play an important role both as food and as habitat. For instance, Macrocystis, a kelp of the order Laminariales, may reach 60 m (200 ft) in length and forms prominent underwater kelp forests. Kelp forests like these contain a high level of biodiversity.[4] Another example is Sargassum, which creates unique floating mats of seaweed in the tropical waters of the Sargasso Sea 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 of brown algae are known worldwide.[5] Some species, such as Ascophyllum nodosum, are important in commercial use because they have become subjects of extensive research in their own right. They have environmental significance as well, through carbon fixation.[4]

Brown algae belong to the group Heterokontophyta, a large group of eukaryotic organisms distinguished most prominently by having chloroplasts surrounded by four membranes, suggesting an origin from a symbiotic relationship between a basal eukaryote and another eukaryotic organism. Most brown algae contain the pigment fucoxanthin, 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 spores and gametes 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 long.[6] 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 kelps are often the most conspicuous algae in their habitats.[7] Kelps can range in size from the two-foot-tall (61 cm) sea palm Postelsia to the giant kelp Macrocystis pyrifera, which grows to over 45 m (150 ft) long[8][9] and is the largest of all the algae. In form, the brown algae range from small crusts or cushions[10] to leafy free-floating mats formed by species of Sargassum. They may consist of delicate felt-like strands of cells, as in Ectocarpus, or of foot-long 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. The particular shade depends upon the amount of fucoxanthin present in the alga.[11] Second, all brown algae are multicellular. There are no known species that exist as single cells or as colonies of cells,[11] and the brown algae are the only major group of seaweeds 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.

Visible structures

Two specimens of Laminaria hyperborea, each showing the rootlike holdfast at lower left, a divided blade at upper right, and a stemlike stipe connecting the blade to the holdfast.

Whatever their form, the body of all brown algae is termed a thallus, indicating that it lacks the complex xylem and phloem of vascular plants. 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.[12] 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 Sargassum or Macrocystis). 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 phloem of vascular plants both in structure and function. In others (such as Nereocystis), 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 Macrocystis pyrifera that grow in strong currents, or may be more rigid in species like Postelsia palmaeformis 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.

Species like Fucus vesiculosus produce numerous gas-filled pneumatocysts (air bladders) to increase buoyancy.

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.[6] In other species, the surface of the blade is coated with slime to discourage the attachment of epiphytes or to deter herbivores. Blades are also often the parts of the alga that bear the reproductive structures.

Gas-filled floats called pneumatocysts provide buoyancy in many kelps 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 ellipsoidal, 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 Sargassum 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

Growth in Dictyota dichotoma occurs at each frond tip, where new cells are produced.

The brown algae include the largest and fastest growing of seaweeds.[6] Fronds of Macrocystis may grow as much as 50 centimetres (20 in) per day, and the stipes can grow 6 centimetres (2.4 in) in a single day.[13]

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.[7] 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.[11]

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.[14]

Aside from filamentous forms, there are two main types of tissue organization in the brown algae: pseudoparenchymatous (haplostichous) and parenchymatous (polystichous).[15] The fronds may be multiaxial or monoaxial.

The cell wall consists of two layers; the inner layer bears the strength, and consists of cellulose; the outer wall layer is mainly algin, and is gummy when wet but becomes hard and brittle when it dries out.[15]