Algal Cell Walls


Algae represent a diverse group of mostly photosynthetic eukaryotes that are profoundly important to Earth's ecosystems and human economy. These organisms possess a variety of extracellular matrix (ECM) components that are critical for multiple life functions. The most well‐studied ECMs are cell walls that are common to the green algae, red algae and brown algae. The typical cell wall consists of crystalline fibrillar polysaccharides (e.g. cellulose) that interact with a surrounding matrix of polysaccharides and proteoglycans. The matrix polysaccharides may be sulfated or acidic and some can complex various cations to form hard surfaces. Other distinct cell wall‐like coverings are found in algae including glycoprotein walls of volvocalean flagellates and silica‐complexed frustules of diatoms. Other noncell wall ECM types of algae include the amphiesma or dinoflagellates and the coccosphere of haptophytes.

Key Concepts

  • Algae possess a variety of extracellular matrices including cell walls.
  • Most green, red and brown algae have cell walls consisting of a composite of a fibrillar polysaccharide framework associated with a polysaccharide/protein matrix.
  • Unique cell walls are found in some algal groups including crystalline glycoprotein cell walls of volvocalean green algae and silica‐complexed cells walls of diatoms.
  • Many algae do not have cell walls but have coverings made of complex scales and plates.
  • The cell wall is synthesised and deposited externally through the coordinated action of the endomembrane and cytoskeletal systems.

Keywords: algae; cell walls; extracellular matrix; scales; coccosphere; frustule; amphiesma

Figure 1. The scaly extracellular covering of the green alga, Mesostigma viride. (a) View through the flattened, biflagellated cell showing the noticeable scale‐covering (arrowheads). Bar, 1 mm. (b) Glancing section of the cell surface. Note the three layers of the scales, the small underlayer of scales (long arrow), the middle layer, oval scales (small arrowhead) and the large basket scales (wide arrow). Bar, 500 nm. (c) View through a preparation of isolated basket scales (arrows). Bar, 500 nm. (d) Magnified view of the basket scale (arrow) revealing the intricate design of the lattice and supporting struts. Bar, 60 nm. (e) A whole cell labelled with a fluorescent‐labelled antibody raised against the basket scale. Note the linear array of scales (arrows) upon the cell surface. Approximately 800 scales can cover a typical cell. Bar, 3 mm. (a–d) were processed for conventional transmission electron microscopy while (e) was processed for immunofluorescence light microscopy.
Figure 2. The cell wall of the chlamydomonad green alga, Gloeomonas kupfferi. (a) Thin section through the wall; (b) deep etch freeze fracture preparation of the cell wall. For details, see Domozych and Dairman . Note the multilayered nature of the wall with a dense, fibrillar, inner layer (IL), a crystalline median layer (arrowheads) and a fibrous outer layer (OL). The chlamydomonad cell wall consists of an aggregation of hydroxyproline‐rich glycoproteins. (a) Bar, 275 nm; (b) Bar, 250 nm.
Figure 3. The cell wall of the CGA taxon, Penium margaritaceum. (a) JIM5, a monoclonal antibody that is specific to high esterified pectin, labels the pectin lattice of the mature wall. (b) CBM3a, a carbohydrate‐binding module that is specific for crystalline cellulose labels the inner wall of the cell wall. (c) JIM7, a monoclonal antibody that binds to high esterified pectins labels a thin band in the cell centre. See Domozych et al. for details. (a) Bar, 5 mm, (b) Bar, 5 mm and (c) Bar, 17 mm.
Figure 4. Transmission electron microscopy of the cell wall of the CGA taxon, Penium margaritaceum. The most distinctive component of the wall is a series of dense aggregation of pectin fibrils that form a unique outer lattice. Bar, 200 nm.
Figure 5. Scanning electron microscopy images of the diatom Didymosphaenia. (a) This diatom produces a silica‐based frustule and a distinct stalk. (b) The frustule (arrow) is highly ornate with multiple openings, several of which are used for the extrusion of EPS. (a) Bar, 50 mm, (b) Bar, 20 mm.


Becker B , Becker D , Kamerling JP and Melkonian M (1991) 2‐keto‐sugar acids in green flagellates: a chemical marker for prasinophycean scales. Journal of Phycology 27: 498–504.

Borowitzka MA and Larkum AW (2008) Calcification in algae: mechanisms and the role of metabolism. Critical Reviews in Plant Sciences 6: 1–45.

Boyer JS (2016) Enzyme‐less growth in Chara and terrestrial plants. Frontiers in Plant Science. DOI:

Brodie J , Chan CX , De Clerck O , et al. (2017) The algal revolution. Trends in Plant Science 22: 726–738.

Brzezinski MA (2008) Mining the genome for the mechanism of biosilicification. Proceedings of the National Academy of Sciences of the USA 105: 1391–1392.

Cock JM , Sterck L , Rouze P , et al. (2010) The Ectocarpus genome and the independent evolution of multicellularity in the brown algae. Nature 645: 617–621.

Cole KM and Sheath RG (1990) Biology of the Red Algae. New York: Cambridge University Press.

Corre G , Templier J , Largeau C , Rousseau B and Berkaloff C (1996) Influence of cell wall composition on the resistance of two Chlorella species (Chlorophyta) to detergents. Journal of Phycology 32: 584–590.

Costaouec TL , Unamunzaga C , Mantecon L and Helbert W (2017) New structural insights into the cell‐wall polysaccharide of the diatom Phaeodactylum tricornutum . Algal Research 26: 172–179.

Craigie JS (1990) Cell walls. In: Cole KM and Sheath RG (eds) Biology of the Red Algae, pp. 221–258. New York: Cambridge University Press.

De Clerck O , Guiry MD , Leliaert F , Samyn Y and Verbruggen H (2013) Algal taxonomy: a road to nowhere? Journal of Phycology 49: 215–225.

Delwiche CF , Graham LE and Thomson N (1989) Lignin‐like compounds and sporopollenin Coleochaete, an algal model for land plant ancestry. Science 245: 399–401.

Deniaud‐Bouet E , Kervarec N , Michel G , et al. (2014) Chemical and enzymatic fractionation of cell walls from Fucales: insights into the structure of the ECM of brown algae. Annals of Botany 114: 1203–1216.

Deniaud‐Bouet E , Hardouin K , Potin P , Kloareg B and Hervé C (2017) A review about brown algal cell walls and fucose‐containing sulfated polysaccharides: cell wall context, biomedical properties and key research challenges. Carbohydrate Polymers 175: 395–408.

Domozych DS , Wells B and Shaw PJ (1991) Basket scales of the green alga, Mesostigma viride: chemistry and ultrastructure. Journal of Cell Science 100: 397–407.

Domozych DS and Dairman M (1993) Synthesis of the inner cell wall of the chlamydomonad flagellate, Gloeomonas kupfferi . Protoplasma 176: 1–13.

Domozych DS , Kort S , Benton S and Yu T (2005) The extracellular polymeric substance of the green alga Penium margaritaceum and its role in biofilm formation. Biofilms 2: 129–144.

Domozych DS , Lambiasse L , Kiemle SN and Gretz MR (2009) Cell‐wall development and bipolar growth in the desmid Penium margaritaceum (Zygnematophyceae, Streptophyta). Asymmetry in a symmetric world. Journal of Phycology 45: 879–893.

Domozych DS , Brechka H , Britton A and Toso M (2011) Cell wall growth and modulation dynamics in a model unicellular green alga, Penium margaritaceum: live cell labeling with monoclonal antibodies. Journal of Botany 8: 632165. DOI: 10.1155/2011/632165.

Domozych DS , Ciancia M , Fangel JU , et al. (2012) The cell walls of green algae: a journey through evolution and diversity. Frontiers in Plant Science. DOI: 10.3389/fpls.2012.00082.

Domozych DS , Sørensen I , Popper ZA , et al. (2014) Pectin metabolism and assembly in the cell wall of the charophyte green alga penium margaritaceum . Plant Physiology 165: 105–118.

Domozych DS (2016) Biosynthesis of the cell walls of the algae. In: Borowitzka MA , Beardall J and Raven JA (eds) The Physiology of Microalgae. New York: Springer.

Eder M and Lutz‐Meindl U (2008) Pectin‐like carbohydrates in the green alga Micrasterias characterized by cytochemical analysis and energy filtering TEM. Journal of Microscopy 231: 201–214.

Estevez JM , Leonardi PI and Alberghina JS (2008) Cell wall carbohydrate epitopes in the green alga Oedogonium bharuchae F. minor (Oedogoniales, Chlorophyta). Journal of Phycology 44: 1257–1268.

Fernandez PV , Ciancia M , Miravalles AB and Estevez JM (2010) Cell‐wall polymer mapping in the coenocytic macroalga Codium vermilara (Bryopsidales, Chlorophyta). Journal of Phycology 46: 456–465.

Finkel ZV (2016) Silicification in the Microalgae. In: Borowitzka MA , Beardall J and Raven JA (eds) The Physiology of Microalgae, pp. 289–299. Cham: Springer.

Graham LE , Graham JM and Wilcox LW (2009) Algae, 2nd edn, p. 616. San Francisco, CA: Benjamin/Cummings Publishing Co.

Hackett JD , Anderson DM , Erdner DL and Bhattacharya D (2004) Dinoflagellates: a remarkable evolutionary experiment. American Journal of Botany 91: 1523–1534.

Hervé C , Siméon A , Jam M , et al. (2016) Arabinogalactan proteins have deep roots in eukaryotes: identification of genes and epitopes in brown algae and their role in Fucus serratus embryo development. New Phytologist 209: 1428–1441.

Higgins MJ , Crawford SA , Mulvaney P and Wetherbee R (2002) Characterization of the adhesive mucilages secreted by live diatom cells using atomic force microscopy. Protist 153: 25–38.

Hoagland KD , Rosowski JR , Gretz MR and Roemer SC (1993) Diatom extracellular polymeric substances: function, fine structure, chemistry and physiology. Journal of Phycology 29: 537–566.

Holzinger A , Callaham DA , Hepler PK and Meindl U (1995) Free calcium in Micrasterias local gradients are not detected in growing lobes. European Journal of Cell Biology 67: 363–371.

Kiemle SN , Domozych DS and Gretz MR (2007) The extracellular polymeric substances of desmids (Conjugatophyceae, Streptophyta): chemistry, structural analyses and implications in wetland biofilms. Phycologia 46: 617–627.

Kimura S and Mizuta S (1995) Cell wall expansion in the marine coenocytic green algae, Chaetomorpha and Boergesenia . Botanica Marina 38: 21–30.

Kloareg B and Quatrano RS (1988) Structure of the cell walls of marine algae and ecophysiological functions of the matrix polysaccharides. Oceanography and Marine Biology. Annual Review 26: 259–315.

Kroger N and Poulsen N (2008) Diatoms – from cell wall biogenesis to nanotechnology. Annual Review of Genetics 42: 83–107.

Kwok ACM and Wong JTY (2003) Cellulose synthesis is coupled to cell cycle progression at G1 in the dinoflagellate Crypthecodinium cohnii . Plant Physiology 131: 1681–1691.

Lee KJ , Marcus SE and Knox JP (2011) Cell wall biology: perspectives from cell wall imaging. Molecular Plant 4: 212–219.

Leliaert F , Smith DR , Moreau H , et al. (2012) Phylogeny and molecular evolution of the green algae. Critical Reviews in the Plant Sciences 31: 1–46.

Lucentinii J (2005) Secondary endosymbiosis exposed. The Scientist 19: 22–23.

Lutz‐Meindl U (2016) Micrasterias as a Model System in Plant Biology. Frontiers in Plant Science. DOI: 10.3389/fpls.2016.00999.

Mann DG , Crawford RM and Round FE (2016) Bacillariophyta. In: Archibald JM , Simpson AGB and Slamovits CH (eds) Handbook of Protists. Springer.

Marsh ME (1996) Polyanion‐mediated mineralization – a kinetic analysis of the calcium‐carrier hypothesis in the phytoflagellate Pleurochrysis carterae . Protoplasma 190: 181–188.

Martone PT , Estevez JM , Fachuang L , et al. (2009) Discovery in seaweed reveals convergent evolution of cell‐wall architecture. Current Biology 19: 169–175.

Mayama S and Kuriyama A (2002) Diversity of mineral coverings and their formation processes: a review focused on the siliceous cell coverings. Journal of Plant Research 115: 289–295.

Michel G , Tonon T , Scornet D , Cock JM and Kloareg B (2010) The cell wall polysaccharide metabolism of the brown alga Ectocarpus siliculosus. Insights into the evolution of extracellular matrix polysaccharides in Eukaryotes. New Phytologist 188: 82–97.

Mihranyan A (2010) Cellulose from cladophorales green algae: from environmental problem to high‐tech composite materials. Journal of Applied Polymer Science 119: 2449–2460.

Mikkelsen MD , Harholt J , Ulvskov P , et al. (2014) Evidence for land plant cell wall biosynthetic mechanisms in charophyte green algae. Annals of Botany 114: 1217–1236.

Moestrup O and Walne PL (1979) Studies on scale morphogenesis in the Golgi apparatus of Pyramimonas tetrarhynchus (Prasinophyceae). Journal of Cell Science 36: 437–459.

Morrill LC and Loeblich AR (1983) Ultrastructure of the dinoflagellate amphiesma. International Review of Cytology 82: 151–180.

Pickett‐Heaps J , Schmid A‐MM and Edgar LA (1990) The cell biology of diatom valve formation. Progress in Phycological Research 7: 1–168.

Pozdnyakov I and Skarlato S (2012) Dinoflagellate amphiesma at different stages of the life cycle. Protistology 7: 108–115.

Popper ZA and Fry SC (2003) Primary cell wall composition of bryophytes and charophytes. Annals of Botany 91 (1): 1–12.

Popper ZA (2008) Evolution and diversity of green plant cell walls. Current Opinion in Plant Biology 11: 286–292.

Popper ZA and Tuohy MG (2010) Beyond the green: understanding the evolutionary puzzle of plant and algal cell walls. Plant Physiology 153: 373–383.

Popper ZA (2011) The Plant Cell Wall. Methods in Molecular Biology, vol. 715. Totowa, NJ: Humana Press.

Popper ZA , Gurvan M , Herve C , et al. (2011) Evolution and diversity of plant cell walls: from algae to flowering plants. Annual Review of Plant Biology 62: 1–24.

Poulsen N , Sumper M and Kroger N (2003) Biosilica formation in diatoms: characterization of native silaffin‐2 and its role in silica morphogenesis. Proceedings of the National Academy of Sciences of the USA 100: 12075–12080.

Ray B and Lahaye M (1995) Cell‐wall polysaccharides from the marine green alga Ulva ‘rigida’ (Ulvales, Chlorophyta). Chemical structure of ulvan. Carbohydrate Research 274: 313–318.

Renn D (1997) Biotechnology and the red seaweed polysaccharide industry: status, needs and prospects. Trends in Biotechnology 15: 9–14.

Rinaudo M (2007) Seaweed polysaccharides. In: Kamerling JP (ed) Comprehensive Glycoscience. From Chemistry to Systems Biology, vol. 2, pp. 691–735. New York: Elsevier.

Roberts AW and Roberts EM (2004) Cellulose synthase (CesA) genes in algae and seedless plants. Cellulose 11: 419–435.

Rossi F and De Philippis R (2016) Exocellular polysaccharides in microalgae and cyanobacteria: chemical features, role and enzymes and genes involved in their biosynthesis. In: Borowitzka MA , Beardall J and Raven JA (eds) The Physiology of Microalgae, pp. 565–590. Cham: Springer.

Rydahl MG , Hansen AR , Kracun SK and Mravec J (2018) Report on the current inventory of the toolbox for plant cell wall analysis: proteinaceous and small molecule probes. Frontiers in Plant Science. DOI: 10.3389/fpls.2018.00581.

Sandgren CD , Smol JP and Kristiansen J (2009) Chrysophyte Algae: Ecology, Phylogeny and Development. New York: Cambridge University Press.

Siver PA (2003) Synurophyte algae. In: Wehr JD and Sheath RG (eds) Freshwater Algae of North America. Ecology and Classification, pp. 523–557. New York: Academic Press.

Sørensen I , Domozych DS and Willats WGT (2010) How have plant cell walls evolved? Plant Physiology 153: 366–372.

Sørensen I , Pettolino FA , Bacic A , et al. (2011) The charophycean green algae provide insaights into the early origins of plant cell walls. The Plant Journal 68: 201–211.

Synytsya A , Čopíková J , Kim WJ and Park YI (2015) Cell wall polysaccharides of marine algae. In: Kim SK (ed) Springer Handbook of Marine Biotechnology. Berlin, Heidelberg: Springer.

Sze P (1998) A Biology of the Algae. New York: WCB/McGraw‐ Hill.

Taylor AR and Brownlee C (2016) Calcification. In: Borowitzka MA , Beardall J and Raven JA (eds) The Physiology of Microalgae, pp. 301–320. Cham: Springer.

Terauchi M , Nagasato C , Inoue A , Ito T and Motomura T (2015) Distribution of alginate and cellulose and regulatory role of calcium in the cell wall of the brown alga Ectocarpus siliculosus (Ectocarpales, Phaeophyceae). Planta 244: 361–377.

Terauchi M , Nagasato IA , Ito T and Motomura T (2016) Distribution of alginate and cellulose and regulatory role of calcium in the cell wall of the brown alga Ectocarpus siliculosis (Ectocarpales, Phaeophyceae). Planta 244: 361–377.

Terauchi M , Yamagishi T , Hanyuda T and Kawai H (2017) Genome‐wide computational analysis of the secretome of brown algae (Phaeophyceae). Marine Genomics 32: 49–59.

Tsuji Y and Yoshida M (2017) Biology of haptophytes: complicated cellular processes driving the global carbon cycle. Advances in Botanical Research 84: 219–261.

Vardi A , Thamatrakoln K , Bidle KD and Falkowski PG (2009) Diatom genomes come of age. Genome Biology 9: 245.

Voigt J and Frank R (2003) 14‐3‐3 proteins are constituents of the insoluble glycoprotein framework of the Chlamydomonas cell wall. The Plant Cell 15: 1399–1413.

Voigt J , Woestemeyer J and Frank R (2007) The chaotrope‐soluble glycoprotein GP2 is a precursor of the insoluble glycoprotein network of the Chlamydomonas cell wall. Journal of Biological Chemistry 282: 30381–30392.

Wee JL (1997) Scale biogenesis in synurophycean protists: phylogenetic implications. CRC Critical Reviews in Plant Science 16: 497–534.

Wizemann A , Meyer FW and Westphal H (2014) A new model for the calcification of the green macro‐alga Halimeda opuntia (Lamouroux). Coral Reefs 33: 951–964.

Ye N , Zhang X , Miao M , et al. (2015) Saccharina genomes provide novel insight into kelp biology. Nature Communications 6: 6986.

Zurzolo C and Bowler C (2001) Exploring bioorganic pattern formation in diatoms. a story of polarized trafficking. Plant Physiology 127: 1339–1345.

Further Reading

Domozych DS (2015) Cell wall evolution and diversity. In: Ramawat KG and Merillon J‐M (eds) Polysaccharides, pp. 55–79. Switzerland: Springer.

Graham LE , Graham JM and Wilcox LW (2009) Algae, 2nd edn, p. 616. San Francisco, CA: Benjamin/Cummings Publishing Co.

Popper ZA , Gurvan M , Herve C , et al. (2011c) Evolution and diversity of plant cell walls: from algae to flowering plants. Annual Review of Plant Biology 62: 1–24.

Preisig HR , Anderson OR and Corliss JO (1994) Terminology and nomenclature of protist cell surface structures. Protoplasma 181: 1–28.

Round FE , Crawford RM and Mann DG (1990) The Diatoms. New York: Cambridge University Press.

Winter A and Siessner WG (1994) Coccolithophores. New York: Cambridge University Press.

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Domozych, David(Jan 2019) Algal Cell Walls. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1002/9780470015902.a0000315.pub4]