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Brown Algae

Christopher E Lane, Dalhousie University, Nova Scotia, Canada Gary W Saunders, University of New Brunswick, Fredericton, Canada
Brown algae form a natural class, the Phaeophyceae, including c. 265 genera with 1500 2000 species. They form rich underwater forests along much of the cold and temperate coastlines of our planet.

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Article Contents
. Introduction . Morphology and Anatomy . Cell Structure . Life Histories . Major Groups . Palaeontology . Ecology

Brown algae (Figure 1) form a natural class, the Phaeophyceae, including c. 265 genera with 15002000 species (van den Hoek et al., 1995). They belong to a larger algal assemblage collectively termed the heterokont chromophytes. Chromophyte refers to the plastid shared by these lineages, which has four bounding membranes, the pigments chlorophyll a and c and various accessory pigments including fucoxanthin, which imparts the brown colour to the Phaeophyceae. Heterokont refers to the agellar arrangement associated with motile cells; the leading agellum is characterized by two rows of sti mastigonemes and the trailing agellum is smooth. Vegetative stages for other heterokont chromophyte classes are predominantly agellate or coccoid unicells and colonies. Brown algal agellate stages are conned to the reproductive gametes and spores, whereas vegetative stages are exclusively multicellular (Figure 1), ranging from microscopic laments to foliose plants up to 50 m in length and 300 kg in weight (Laminariales). Brown algae are, therefore, generally discussed in reference to their ecological and morphological equivalents, those marine members of the Chlorophyta (green algae) and Rhodophyta (red algae) collectively termed seaweeds. See also: Algal agella; Green algae

. Economic Significance

doi: 10.1038/npg.els.0004250

hyphae) form cellular pipelines within the thallus for the distribution of photosynthetic product. Growth may be diuse (i.e. cell divisions occur throughout the plant), or concentrated in meristems that can be intercalary, trichothallic (intercalary meristem positioned at the base of a terminal lament), or apical from a single cell, group of cells or plant margins. In some species, a supercial layer of meristematic cells, the meristoderm, contributes to growth by adding cells to the epidermis, as well as towards the thallus interior contributing to plant girth.

Cell Structure
Vegetative cells (Figure 2) are surrounded by a wall of cellulosic microbrils reinforced by insoluble alginates (e.g. calcium alginate). This structural portion of the wall is embedded in an amorphous mucilaginous component (soluble alginates and the sulfated polysaccharide fucoidan). The plasma membrane lies beneath the wall, encircling the cytoplasm and is continuous between cells via plasmodesmata. Organelles consistent with typical phototrophic eukaryotes are present: a nucleus, with one or more nucleoli, surrounded by the nuclear endoplasmic reticulum (NER) which is continuous with the cytoplasmic ER; mitochondria and Golgi bodies, commonly positioned adjacent to the NER. There are a few ribbon-shaped to numerous discoid plastids that are surrounded by a twomembrane chloroplast ER (CER), the latter continuous with the NER where plastids lie in proximity to the nucleus. Inside the CER is the two-membrane chloroplast envelope, the compartment between these two double-membrane systems containing periplastidial ER. Lamellate thylakoids generally occur in stacks of three with one stack occupying the periphery of the plastid (girdle lamellae). The plastid deoxyribonucleic acid (DNA) is concentrated in a ring-shaped genophore. Pyrenoids (PY) occur in some brown algal plastids and are protuberant, often having a stalked appearance. The major carbohydrate reserve,

Morphology and Anatomy

The simplest brown algae have a heterotrichous organization consisting of a prostrate lamentous system giving rise to free branched or unbranched erect laments (Figure 1b). Additional haplostichous (lamentous construction) forms include loose aggregates of laments enclosed in common mucilaginous mass, solid pseudoparenchymatous aggregations of laments producing crusts (Figure 1c) and branched or unbranched terete and foliose plants. Polystichous construction (cell divisions in many planes to yield true parenchyma) produces terete to foliose thalli in some species (Figure 1d j). Brown algae display the greatest anatomical complexity among the protists. Cells have cytoplasmic continuity via plasmodesmata (analogous to land plant structures) and in the orders Laminariales and Desmarestiales specialized cells (trumpet

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Brown Algae

Figure 1 Morphological diversity among brown algae. (a) Epilithic carpet of intertidal brown algae. (b) Pilayella littoralis: filamentous brown (scale 1 cm). (c) Ralfsia fungiformis: crustose representative (scale 2 cm). (d) Laminaria saccharina: a kelp (scale 10 cm). (e) Alaria nana: a kelp with sporophylls (ruler in cm). (f) Postelsia palmaeformis: a member of the Lessoniaceae (scale 11 cm). (g) Macrocystis integrifolia: a giant kelp; inserts (left to right) holdfast, sporophyll, region of blade production. (h) Hormosira banksii: a southern hemisphere fucoid (scale 3 cm). (i) F. spiralis: a high intertidal fucoid (scale 3 cm). (j) F. distichus ssp. edentatus: a low intertidal inhabitant (scale 2.5 cm) with a close-up (k) of developing receptacles and their characteristic conceptacles (scale 1 cm). The author graciously acknowledges the assistance of Mr Roger Smith (University of New Brunswick) in production of this figure.

Brown Algae



is the rule for brown algae and can be isomorphic or heteromorphic (generations appear similar (Figure 3a) or dierent (Figure 3b), respectively). Meiosis (R!) occurs in unilocular sporangia, resulting in the production and release of meiospores, whereas the mitotic production of gametes and asexual mitospores, usually occurs in plurilocular structures termed gametangia and sporangia, respectively. Isogamy, anisogamy and oogamy are observed for dierent species. See also: Gametophyte and sporophyte


Major Groups


Figure 2 Schematic diagram of vegetative cell anatomy. CW, cell wall; PM, plasma membrane; PL, plasmodesmata; NU, nucleus; N, nucleolus; NER, nuclear endoplasmic reticulum; ER, endoplasmic reticulum; M, mitochondria; GO, Golgi body; CH, discoid plastid; CER, chloroplast endoplasmic reticulum; CE, chloroplast envelope; PER, periplastidial endoplasmic reticulum; T, thylakoid; G, gonophore; PY, pyrenoid; CV, cytoplasmic vacuole; PH, physode. The anatomy of one plastid (CH) was not elaborated. Based on information from van den Hoek et al. (1995).

Brown algae are traditionally separated into orders on the basis of life history pattern, type of growth and thallus structure, number and type of plastid and whether or not pyrenoids are present. As in all algal groups, this system of classication is being revised as a result of molecular systematics. As many as 16 orders have been recognized (see van den Hoek et al., 1995 for detailed descriptions), but consensus is lacking and schemes with fewer orders are recognized (Druehl et al., 1997; Rousseau et al., 2001; Draisma et al., 2001). Five of these orders are discussed. See also: Algal taxonomy: historical overview

A cosmopolitan order with epilithic, epiphytic or partially endophytic species reported, the Ectocarpales has been considered the most primitive group of brown algae based on its lamentous morphology. Taxonomic revisions, considering both the anities of the Ectocarpales relative to other brown algae orders (Rousseau et al., 2001; Draisma et al., 2001) and the generic composition of this order (Rousseau and de Reviers, 1999), have been recommended in light of recent molecular investigations. We present here the Ectocarpales sensu Rousseau and de Reviers (1999), including only taxa possessing exserted, pedunculated pyrenoids.

laminarin, is soluble and stored in cytoplasmic vacuoles commonly positioned adjacent to the CER where the PY is formed. Brown algal cells contain special vesicles called physodes (discussed below). See also: Algal cell walls; Algal chloroplasts

Life Histories
Alternation of generations, i.e. a free-living diploid sporophyte alternates with an independent haploid gametophyte,


Unilocular sporangium R! Male gamete

Plurilocular gametangia

Unilocular sporangium R! Antheridium Male gamete Egg Oogonium

Sporophyte Zygote (a)

Sporophyte Female gamete Gametophyte (b)


Young sporophyte



Figure 3 Common life history types. (a) Isomorphic pattern typical of some Ectocarpales. (b) Heteromorphic pattern of the Laminariales. See text for details. R!, meiosis.

Brown Algae

Vegetative thalli (Figure 1b) consist of branched uniseriate laments. Thalli are heterotrichous with laments of the prostrate base growing by apical division. Intercalary divisions occur in the erect laments and can be dispersed or concentrated in meristems. Isomorphic alternation of generations is prevalent (Figure 3a). Gametes are formed in plurilocular gametangia on the haploid gametophytes. Morphological similarity (isogametes) belies the true complexity of these gametes, which commonly display behavioural anisogamy, i.e. female gametes settle soon after release and emit pheromones that attract the male gametes. Fusion results in a diploid zygote that grows into the sporophyte. Sporophytes produce unilocular sporangia in which meiosis (R!) and subsequent mitoses occur to produce numerous motile haploid meiospores that will develop into male and female gametophytes. Plurilocular structures can occur on sporophytes, but in this case they produce diploid spores by mitosis that regenerate the sporophyte generation.

Species of the largest genus in this order, Desmarestia, are roughly divided into three groups (Peters et al., 1997). Two of these are present in North America represented by alternately branched perennials and oppositely branched annuals. In the oppositely branched species, sulfuric acid is present in vacuoles. Care in collecting these species is a necessity because if they are stored with other samples they will rapidly destroy, in addition to themselves, the spoils of an entire collecting trip!

Collectively referred to as kelp, these algae form extensive stands in the cold waters of the northern hemisphere in both the Atlantic and Pacic Oceans, reaching their culmination in the luxuriant kelp forests of the northeast Pacic. There have also been sporadic introductions to the southern hemisphere, possibly during recent cooling events (Druehl et al., 1997). Kelp have a strongly heteromorphic alternation of generations (Figure 3b). The dominant sporophytes lend the kelp their distinction as the largest seaweeds and are foliose with a characteristic morphology and anatomy. Plants are anchored by a disc-like, rhizoidal or branched (hapteral) holdfast positioned below a distinct stem-like stipe in most species, which is crowned by a blade (Figure 1d g). In some species there are specialized blades, termed sporophylls, on which unilocular sporangia are produced, and some species have numerous blades resulting from splitting at the stipe to blade transition zone. Six families are currently recognized within the Laminariales, three considered ancestral (Akkesiphycaceae, Chordaceae and Pseudochordaceae (Kawai and Sasaki, 2000; Sasaki et al., 2001) and three derived (Alariaceae, Laminariaceae and Lessoniaceae). Traditionally, the taxonomy of the derived families has been based on the presence (Alariaceae: Figure 1e) or absence of sporophylls, and for the latter case the presence (Lessoniaceae: Figure 1f g) or absence (Laminariaceae: Figure 1d) of splitting. Some species, however, display characteristics of two families (e.g. Macrocystis integrifolia, with splitting and sporophylls, Figure 1g) and this system of taxonomy has been further compromised by molecular investigations (Druehl et al., 1997; Yoon et al., 2001), which clearly show polyphyly among the three traditional families. Growth of sporophytes is from an intercalary meristem and associated meristoderm positioned at the stipe-toblade transition zone. Some species are annuals growing as much as 36 cm per day, whereas other species are perennial, with life expectancies of 317 years. The resulting plants have parenchymatous construction with tissue specialization unequalled among algae. Notably, elongate cells with ared ends at the crosswalls (termed trumpet hyphae because of their morphology) form long tracts of conducting tissue within the medulla. Crosswalls are

A mostly tropical order exhibiting erect, parenchymatous thalli and an isomorphic alternation of generations. Growth is from either an apical cell, or a meristematic marginal row of cells. The modication of the unilocular sporangia in the Dictyotales, which divide to produce four to eight aplanospores, is unique among brown algae. Additionally, while oogomous sexual reproduction is not unusual among brown algae, members of the Dictyotales are the only members of the Phaeophyceae to have uniagellate sperm (a second basal body is present). Like the Ectocarpales, the accepted phylogenetic position of the Dictyotales within brown algae has changed with the application of molecular data. Long thought to be one of the most derived orders of brown algae, molecular data indicate that the Dictyotales branched o prior to the ancestral line that gave rise to the majority of brown algal orders and is thus only a distant relative to these lineages (Rousseau et al., 2001; Draisma et al., 2001).

Species included in this order are common inhabitants in colder waters of all oceans and form the dominant component of the subtidal ora in the Antarctic region. Their life histories are similar to the Laminariales type (Figure 3b). Growth of the dominant sporophyte is from a trichothallic meristem with pseudoparenchyma developing below the meristem. This basic construction is augmented in some species by activity of a meristoderm contributing parenchyma to the thallus. Gametophytes are microscopic and lamentous with separate males and females producing antheridia and oogonia, respectively, rather than plurilocular gametangia.

Brown Algae

traversed by numerous plasmodesmata, which can form relatively wide connections in larger species, developing the appearance of sieve plates in the phloem of land plants. The analogy to phloem goes further because it has been established that trumpet hyphae facilitate the movement of photosynthetic product within kelp (Schmitz, 1981). The gametophyte bears no resemblance to the sporophyte and is more reminiscent of an ectocarpoid alga. Male and female gametophytes dier slightly in their appearance but are basically microscopic uniseriate laments (sometimes of only a few cells) that grow by apical division. During sexual reproduction (Figure 3b) darkened sori, the site of unilocular sporangia production, are apparent on the vegetative blade (or sporophylls) of the sporophyte. Meiosis, subsequent rounds of mitosis and nally cytokinesis generate numerous haploid motile meiospores, which are released into the water column to nd suitable substrate. Meiospores germinate as male and female gametophytes in an approximately 50:50 ratio. Males produce antheridia, each releasing a single motile male gamete, whereas females develop oogonia, each producing a single egg that remains attached to the oogonium. Male gametes are attracted to eggs by pheromones and one successfully fuses with the egg eecting fertilization. The resulting zygote undergoes mitotic division to regenerate the sporophyte phase, which eventually overgrows the diminutive female gametophyte.

gametangia only, although in some species a few vegetative cells are produced (Clayton, 1984). Mature gametophytes are then extruded within a common wall, and subsequently release gametes into the water column. Reproduction is oogamous with motile male gametes attracted to the nonmotile eggs. This interpretation of fucalean gametophyte development within the meiosporangium is analogous to the condition in land plants.

Fossils are a historical record of the age, nearest relatives and evolutionary progression within and among groups of organisms. Unfortunately, the predominantly soft tissues of brown algae do not lend themselves to fossilization and authentic nds are rare. Reports of 0.81.2 billion year-old brown algal fossils, and corresponding inferences that these algae evolved along with many other eukaryotic lineages towards the end of the Precambrian, are not substantiated (Clayton, 1984). Fossils from the Palaeozoic and Mesozoic bear supercial resemblance to extant brown algae, but are equally comparable to present-day Chlorophyta and Rhodophyta. Bona de brown algal fossils have been reported from 6 to 10 million year-old Miocene deposits (Parker and Dawson, 1965), but are closely aliated with extant genera of the Laminariales and Fucales, providing few insights into brown algal evolution and ancestry. In the absence of a long and diverse fossil record, traditional systematists have relied on intuition, comparing extant species in eorts to understand phylogeny. See also: Algae: phylogeny and evolution; Algal taxonomy: historical overview

Fucoids (Figure 1h j) are common components of the coastal ora and dominate intertidal biomass in many parts of the world (Figure 1a). In the northern hemisphere species diversity is low, but in the southern hemisphere diversity is rich and the Fucales are considered the ecological equivalent to the Laminariales of the North. The diploid generation, foliose plants constructed of true parenchyma derived from the action of one or more apical cells and their derivative meristoderm, dominates the life cycle. Unilocular structures are produced in conceptacles, which appear as protuberances on the plant surface, and the conceptacles can be aggregated on swollen portions of the plant to produce characteristic receptacles (Figure 1k). In diplontic life histories the diploid generation is technically a gametophyte and produces gametes by meiosis (haploid generation is absent), whereas in an alternation of generations the haploid generation is the gametophyte (Figure 3). Fucoid life histories appear diplontic, but the diploid generation is, nonetheless, considered a sporophyte because life histories are interpreted as a special case of alternation of generations (pseudodiplontic). Meiosis occurs, but free-swimming spores are not released and diminutive gametophytes develop within the unilocular sporangium. Gametophytes are generally reduced to

DNA alternative
More recently, the tools of molecular biology have provided systematists with new perspectives on brown algal evolution. On the basis of molecular data it has been proposed that the brown algae are a recent group of only 200 million years (Saunders and Druehl, 1992), further compromising reports of Precambrian brown algae. Brown algae were considered most closely allied to the Chrysomeridales, an order of heterokont chromophytes, based on the uncommon lateral insertion of agella in these groups (apical/subapical in other heterokonts). This proposal has received weak support from molecular systematic analyses and until recently, the closest relative to the Phaeophyceae in the molecular trees is the heterokont chromophyte class Xanthophyceae (Saunders et al., 1997). The recently described Schizocladophyceae (Kawai et al., 2003) now resolves as the closest ally to the brown algae in molecular trees. Similarly, long-standing views on

Brown Algae

evolution within and among brown algal lineages are under increasing pressure. Consult Druehl et al. (1997) for a molecular perspective on the relationships among extant brown algae, and van den Hoek et al. (1995) for an intuitive synthesis of available data. See also: Molecular phylogeny reconstruction

Brown algae occur in most coastal environments ranging from tropical to polar waters, but display their richest diversity and greatest biomass in the cold temperate waters of the northern and southern hemispheres. The narrow ribbon of coastline to which most brown algae are conned accounts for less than 1% of the earths surface. Nevertheless, the most productive areas on the planet are reportedly brown algal stands (cf. productivity of 0.52.5 kg carbon m22 per year versus 0.42.0 kg carbon m22 per year for tropical rainforests; Dawes, 1998). Availability of suitable substrate generally solid rock is a major factor inuencing the local abundance and distribution of brown algae. In addition to epilithic habitat, brown algae grow on other solid substrate, e.g. concrete, wood and metal, as epiphytes on other plants, partially endophytic and some species even grow as free-oating populations (e.g. Sargassum in the Sargasso Sea). Brown algae can be so abundant that they modify, and in fact become, the habitat for many other algae and animals (Figure 1a). See also: Algal ecology; Biogeography of marine algae

Laminariales (Figure 1d) dominate the upper subtidal and lowest intertidal zones. These are replaced by Fucus distichus ssp. edentatus (Figure 1j ) in the lower intertidal, a zone of Ascophyllum nodosum and F. vesiculosus occupies the mid-intertidal and nally a high intertidal zone dominated by F. spiralis (Figure 1i) is evident. The upper limit of a species vertical distribution is generally controlled by abiotic factors: tolerance to exposure, temperature and desiccation; whereas, lower limits are established by biotic interactions, notably competition (Dawes, 1998). In short, stress-tolerant species are conned to higher-stress environments by relatively more-competitive, less-stress-tolerant species. It should be noted that algal reproductive stages are dispersed to, and probably initiate development in, all zones, but algae are only successful where the balance of environmental stress and competition is in their favour.

Environment and development

Growth, development and reproduction in brown algae is commonly linked to environmental cues including temperature and light (quality, quantity and day length). Kelp gametophytes, for example, can overwinter at the singlecell stage, initiating vegetative growth and subsequent gametogenesis only when a critical day length is reached (also a minimum quantity of blue light is required). Gametogenesis only occurs, however, when water temperatures are below 158C, thus limiting this response to the spring. Sporophytes of perennial kelp (Laminariales) display a seasonal pattern of vegetative growth and reproduction that is regulated by a combination of day length and temperature. The intercalary meristem at the stipe blade transition zone is active in the spring and initiates development of a new blade. As it grows, the new blade displaces the previous years blade, growth of the former being sustained by energy accumulated during photosynthesis in the previous summer and stored in the latter. The day-length response is a true photoperiodic short-day response similar to the phytochrome response observed for land plants.

Brown algae and zonation

Tidal action in the oceans divides the coastal habitat into a subtidal and an intertidal zone. The subtidal zone remains submerged even at low tide, and the lower limit of brown algal growth is dependent on light penetration (abiotic factor) and to a lesser extent on biotic factors (competition and grazing). In clear tropical waters some browns can grow to depths of 100 m, whereas in turbid waters few brown algae grow below the low water mark. Tidal cycles expose intertidal seaweeds to atmospheric conditions that can be harsher and more variable than adjacent aquatic conditions. Exposure carries the risk of desiccation and intertidal algae are well adapted to their unique environment. For example, many fucoid algae can lose up to 90% of their water and subsequently recover normal photosynthesis upon rehydration. Organisms that inhabit the intertidal zone have a restricted vertical distribution manifested as distinct bands. In cold temperate waters brown algae dominate the intertidal zone (Figure 1a) and, therefore, can serve as signature species for the various vertical zones. In the lower Bay of Fundy (New Brunswick, Canada), for example, representatives of the

Chemical ecology
Algae, in particular Phaeophyceae, accumulate a broad spectrum of secondary metabolites that are generally regarded as grazing deterrents. Brown algae produce volatile brominated organocompounds that are released into the atmosphere in amounts equal to the industrial output of like compounds. These compounds contribute to ozone destruction and it has been suggested that holes in the ozone over the Arctic in the spring may be linked to enhanced seasonal production of these compounds by brown algae (van den Hoek et al., 1995). These observations do not serve to mitigate the damage or responsibility that humans bear as a result of industrial production of

Brown Algae

ozone-damaging compounds and continued eorts to control use remain essential to the health of the atmosphere. See also: Algal metabolism; Secondary metabolites: deterring herbivores A second class of compounds, collectively termed phlorotannins, are stored in the physodes of brown algal cells (Figure 2). Physodes may simply be repositories of waste metabolites, which also serve an antibiotic role. Specically, these compounds probably serve to deter herbivores and to inhibit the growth of epiphytic plants and animals on an algas surface. A role in regulating exposure to sunlight has also been advanced, notably in ltering out harmful ultraviolet radiation, a notion consistent with the abundant occurrence of physodes in surface tissue of intertidal fucoids (Clayton, 1984). The role or roles, however, are yet to be determined with certainty.

Economic Significance
The alginates produced by brown algae are used by the food and pharmaceutical industries as stabilizing agents in emulsions and suspensions owing to their colloidal properties (Jensen, 1995). New uses are being developed at an ever-increasing rate, notably in the biotechnology and medical elds. Approximately 575 000 tonnes of brown algae are harvested per year to provide 21 500 tonnes of alginates for the market (van den Hoek et al., 1995). Over half of this is from Europe where Ascophyllum (Fucales) and Laminaria (Laminariales) are harvested. Another third is acquired through the mechanical harvesting of Macrocystis and Nereocystis (Laminariales) from the rich kelp forests of the Pacic coast of North America. Kelp are not only harvested from natural stands, but are cultivated in many countries both as a source of alginates and for food. Kelp species of the genera Laminaria and Undaria are particularly popular food crops in many coastal Asian countries. Brown algae are also commonly used in agriculture and horticulture as fertilizers and soil conditioners, and are recognized to have growth promoting qualities. More recently, polyculture the farming of seaweed in proximity to sh cages is being explored as a means of reducing the impact of aquaculture activities on estuarine ecosystems. See also: Fisheries management

In: Bhattacharya D (ed.) Origins of Algae and their Plastids, pp. 221 235. Vienna, New York: Springer. Jensen A (1995) Production of alginate. In: Wiessner W, Schnepf E and Starr RC (eds) Algae, Environment and Human Aairs, pp. 7992. Bristol: Biopress. Kawai H and Sasaki H (2000) Molecular phylogeny of the brown algal genera Akkesiphycus and Halosiphon (Laminariales), resulting in the circumscription of the new families Akkesiphycaceae and Halosiphonaceae. Phycologia 39: 416428. Parker BC and Dawson EY (1965) Non-calcareous marine algae from California Miocene deposits. Nova Hedwigia 10: 273295. Peters AF, van Oppen MJH, Wiencke C, Stam WT and Olsen JL (1997) Phylogeny and historical ecology of the Desmarestiaceae (Phaeophyceae) support a southern hemisphere origin. Journal of Phycology 33: 294309. Rousseau F, Burrowes R, Peters AF, Kuhlenkamp R and de Reviers B (2001) A comprehensive phylogeny of the Phaeophyceae based on nrDNA sequences resolves the earliest divergences. Compte Rendu Hebdomadaire des Seances de lAcademie des Sciences, Paris, Sciences de la Vie 324: 305319. Rousseau F and de Reviers B (1999) Circumscription of the order Ectocarpales (Phaeophyceae): bibliographical synthesis and molecular evidence. Cryptogamie Algologie 20: 518. Sasaki H, Flores-Moya A, Henry EC, Mu ller D and Kawai H (2001) Molecular phylogeny of Phyllariaceae, Halosiphonaceae and Tilopteridales (Phaeophyceae). Phycologia 40: 123134. Saunders GW and Druehl LD (1992) Nucleotide sequences of the smallsubunit ribosomal RNA genes from selected Laminariales (Phaeophyta): implications for kelp evolution. Journal of Phycology 28: 544 549. Saunders GW, Potter D and Andersen RA (1997) Phylogenetic anities of the Sarcinochrysidales and Chrysomeridales (Heterokonta) based on analyses of molecular and combined data. Journal of Phycology 33: 310318. Schmitz K (1981) Translocation. In: Lobban CS and Wynne MJ (eds) The Biology of Seaweeds, pp. 534558. Berkeley: University of California Press. van den Hoek C, Mann DG and Jahns HM (1995) Algae: An Introduction to Phycology. Cambridge: Cambridge University Press. Yoon HS, Lee JY, Boo SM and Bhattacharya D (2001) Phylogeny of the Alariaceae, Laminariaceae, and Lessoniaceae (Phaeophyceae) based on plastid-encoded RuBisCo spacer and nuclear-encoded ITS sequences comparisons. Molecular Phylogeny and Evolution. 21: 231243.

Further Reading
Bold HC and Wynne MJ (1985) Introduction to the Algae: Structure and Reproduction, 2nd edn. Englewood Clis, NJ: Prentice-Hall. Clayton MN (1988) Evolution and life histories of brown algae. Botanica Marina 31: 379387. De Reviers B and Rousseau F (1999) Towards a new classication of the brown algae. Progress in Phycological Research 13: 107201. Fletcher RL (1987) Seaweeds of the British Isles, Vol. 3: Fucophyceae (Phaeophyceae), part 1. London: British Museum (Natural History). Lobban CS and Harrison PJ (1997) Seaweed Ecology and Physiology. Cambridge: Cambridge University Press. Lu ning K (1990) Seaweeds. Their Environment, Biogeography, and Ecophysiology. New York: Wiley. Womersley HBS (1987) The Marine Benthic Flora of Southern Australia, Part II. Adelaide: South Australian Government Printing Division.

Clayton MN (1984) Evolution of the Phaeophyta with particular reference to the Fucales. Progress in Phycological Research 3: 1146. Dawes CJ (1998) Marine Botany, 2nd edn. New York: Wiley. Draisma SGA, Prudhomme van Reine WF, Stam WT and Olsen JL (2001) A reassessment of phylogenetic relationships within the Phaeophyceae based on Rubisco large subunit and ribosomal DNA sequences. Journal of Phycology 37: 586603. Druehl LD, Mayes C, Tan IH and Saunders GW (1997) Molecular and morphological phylogenies of kelp and associated brown algae.