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Cell wall

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Plant cells separated by transparent cell walls.

A cell wall is a fairly rigid layer surrounding a cell, located external to the cell membrane, which provides the cell with structural support, protection, and acts as a filtering mechanism. The cell wall also prevents over-expansion when water enters the cell. They are found in plants, bacteria, archaea, fungi, and algae. Animals and most protists do not have cell walls.

The cell wall is constructed from different materials dependent upon the species. In plants, the cell wall is constructed primarily from a carbohydrate polymer called cellulose, and the cell wall can therefore also function as a carbohydrate store for the cell. In bacteria, peptidoglycan forms the cell wall. Archaea have various chemical compositions, including glycoprotein S-layers, pseudopeptidoglycan, or polysaccharides. Fungi possess cell walls of chitin, and algae typically possess walls constructed of glycoproteins and polysaccharides, however certain algal species may have a cell wall composed of silicic acid. Often, other accessory molecules are found anchored to the cell wall.

Properties

Diagram of the plant cell, with the cell wall in green.

The cell wall serves a similar purpose in those organisms that possess them. The wall gives cells rigidity and strength, offering protection against mechanical stress. In multicellular organisms, it permits the organism to build and hold its shape (morphogenesis). The cell wall also limits the entry of large molecules that may be toxic to the cell. It further permits the creation of a stable osmotic environment by preventing osmotic lysis and helping to retain water. The composition, properties, and form of the cell wall may change during the cell cycle and depend on growth conditions.

Rigidity

In most cells, the cell wall is semi-rigid, meaning that it will bend somewhat rather than holding a fixed shape. This flexibility is seen when plants wilt, so that the stems and leaves begin to droop, or in seaweeds that bend in water currents. Wall rigidity seen in healthy plants results from a combination of the wall construction and turgor pressure. As John Howland states it:

Think of the cell wall as a wicker basket in which a balloon has been inflated so that it exerts pressure from the inside. Such a basket is very rigid and resistant to mechanical damage. Thus does the prokaryote cell (and eukaryotic cell that possesses a cell wall) gain strength from a flexible plasma membrane pressing against a rigid cell wall.[1]

The rigidity of the cell wall thus results in part from inflation of the cell contained. This inflation is a result of the passive uptake of water.

Other cell walls are inflexible. In plants, a secondary cell wall is a thicker additional layer of cellulose. Additional layers may be formed containing lignin inside xylem cells, or containing suberin in cork cells. These compounds are rigid and waterproof, making the secondary wall stiff. Both wood and bark cells of trees have secondary walls. Other parts of plants such as the leaf stalk may acquire similar reinforcement to resist the strain of physical forces.

Cerrtain single-cell protists and algae also produce a rigid wall. Diatoms build a frustule from silica extracted from the surrounding water; radiolarians also produce a test from minerals. Many green algae, such as the Dasycladales encase their cells in a secreted skeleton of calcium carbonate. In each case, the wall is rigid and essentially inorganic.

Permeability

The primary cell wall of most plant cells is semi-permeable and permit the passage of small molecules. Key nutrients, especially water and carbon dioxide, are distributed throughout the plant from cell wall to cell wall in apoplastic flow.

Plant cell walls

Composition

Molecular structure of the primary cell wall in plants.

The major carbohydrates making up the primary cell wall are cellulose, hemicellulose and pectin. The cellulose microfibrils are linked via hemicellulosic tethers to form the cellulose-hemicellulose network, which is embedded in the pectin matrix. The most common hemicellulose in the primary cell wall is xyloglucan.

The three primary polymers that make up plant cell walls consist of about 35 to 50% cellulose, 20 to 35 % hemicellulose and 10 to 25% lignin. Lignin fills the spaces in the cell wall between cellulose, hemicellulose and pectin components.

Plant cells walls also incorporate a number of proteins; the most abundant include hydroxyproline-rich glycoproteins (HRGP), also called the extensins, the arabinogalactan proteins (AGP), the glycine-rich proteins (GRPs), and the proline-rich proteins (PRPs). With the exception of glycine-rich proteins, all the previously mentioned proteins are glycosylated and contain hydroxyproline (Hyp). Each class of glycoprotein is defined by a characteristic, highly repetitive protein sequence. Chimeric proteins contain two or more different domains, each with a sequence from a different class of glycoprotein. Most cell wall proteins are cross-linked to the cell wall and may have structural functions. The cell wall may contain suberin and cutin, two waxy substances that protect the cell from herbivores.[2] The relative composition of carbohydrates, secondary compounds and protein varies between plants and between the cell type and age.

The plant cell wall has three components:[3]

  • The middle lamella, a layer rich in pectins. This is the outermost layer, forming the interface between adjacent plant cells.
  • The primary cell wall, a wall formed during cell division that expands as the cell grows. It keeps the cell from taking up too much water.
  • The secondary cell wall, a thick layer formed inside the primary cell wall after the cell is fully grown. It is not found in all cell types.

Formation

The middle lamella is laid first, formed from the cell plate during cytokinesis, and the primary cell wall is then expanded inside the middle lamella. The actual structure of the cell wall is not clearly defined and several models exist - the covalently linked cross model, the tether model, the diffuse layer model and the stratified layer model. However, the primary cell wall, can be defined as composed of cellulose microfibrils aligned at all angles. Microfibrils are held together by hydrogen bonds to provide a high tensile strength. The cells are held together and share the gelatinous membrane called the middle lamella, which contains magnesium and calcium pectates (salts of pectic acid). Cells interact though plasmodesma(ta), which are inter-connecting channels of cytoplasm that connect to the protoplasts of adjacent cells across the cell wall.

In some plants and cell types, after a maximum size or point in development has been reached, a secondary wall is constructed between the plant cell and primary wall. Unlike the primary wall, the microfibrils are aligned mostly in the same direction, and with each additional layer the orientation changes slightly. Cells with secondary cell walls are rigid. Cell to cell communication is possible through pits in the secondary cell wall that allow plasmodesma to connect cells through the secondary cell walls.

Trees modify cell walls in their branches to reinforce and support structure.[4] Conifers, such as pine, produce thicker cell walls on the undersides of branches to push their branches upwards. The resulting wood is called compression wood. By contrast, hardwood trees reinforce the walls on the upper sides of branches to pull their branches up. This is known as tension wood. Additional thickening may occur in other parts of the plant in response to mechanical stress.

Algal cell walls

Scanning electron micrographs of diatoms showing the external appearance of the cell wall

Like plants, algae have cell walls.[5] Algal cell walls contain cellulose and a variety of glycoproteins. The inclusion of additional polysaccharides in algal cells walls is used as a feature for algal taxonomy.

  • Manosyl form microfibrils in the cell walls of a number of marine green algae including those from the genera, Codium, Dasycladus, and Acetabularia as well as in the walls of some red algae, like Porphyra and Bangia.
  • Xylanes
  • Alginic acid is a common polysaccharide in the cell walls of brown algae
  • Sulfonated polysaccharides occur in the cell walls of most algae; those common in red algae include agarose, carrageenan, porphyran, furcelleran and funoran.

Other compounds that may accumulate in algal cell walls include sporopollenin and calcium ions.

The group of algae known as the diatoms synthesize their cell walls (also known as frustules or valves) from silicic acid (specifically orthosilicic acid, H4SiO4). The acid is polymerised intra-cellularly, then the wall is extruded to protect the cell. Significantly, relative to the organic cell walls produced by other groups, silica frustules require less energy to synthesize (approximately 8%), potentially a major saving on the overall cell energy budget[6] and possibly an explanation for higher growth rates in diatoms.[7]

Fungal cell walls

Chemical structure of a unit from a chitin polymer chain.

There are several groups of organisms that may be called "fungi". Some of these groups have been transferred out of the Kingdom Fungi, in part because of fundamental biochemical differences in the composition of the cell wall. Most true fungi have a cell wall consisting largely of chitin and other polysaccharides.[8] True fungi do not have cellulose in their cell walls, but some fungus-like organisms do.

True fungi

Not all species of fungi have cell walls but in those that do, the plasma membrane is followed by three layers of cell wall material. From inside out these are:

Fungus-like protists

The group Oomycetes, also known as water molds, are saprotrophic plant pathogens like fungi. Until recently they were widely believed to be fungi, but structural and molecular evidence[9] has led to their reclassification as heterokonts, related to autotrophic brown algae and diatoms. Unlike fungi, oomycetes typically possess cell walls of cellulose and glucans rather than chitin, although some genera (such as Achlya and Saprolegnia) do have chitin in their walls.[10] The fraction of cellulose in the walls is no more than 4 to 20%, far less than the fraction comprised by glucans.[10] Oomycete cell walls also contain the amino acid hydroxyproline, which is not found in fungal cell walls.

The dictyostelids are another group formerly classified among the fungi. They are slime moulds that feed as unicellular amoebae, but aggregate into a reproductive stalk and sporangium under certain conditions. Cells of the reproductive stalk, as well as the spores formed at the apex, possess a cellulose wall.[11] The spore wall has been shown to possess three layers, the middle of which is composed primarily of cellulose, and the innermost is sensitive to cellulase and pronase.[11]

Prokaryotic cell walls

Bacterial cell walls

Diagram of a typical gram-negative bacterium, with the thin cell wall sandwiched between the yellow outer membrane and the thin red plasma membrane
Schematic of typical gram-positive cell wall showing arrangement of N-Acetylglucosamine and N-Acetlymuramic acid
Further information: [[:Cell envelope]]

Around the outside of the cell membrane is the bacterial cell wall. Bacterial cell walls are made of peptidoglycan (also called murein), which is made from polysaccharide chains cross-linked by unusual peptides containing D-amino acids.[12] Bacterial cell walls are different from the cell walls of plants and fungi which are made of cellulose and chitin, respectively.[13] The cell wall of bacteria is also distinct from that of Archaea, which do not contain peptidoglycan. The cell wall is essential to the survival of many bacteria. The antibiotic penicillin is able to kill bacteria by inhibiting a step in the synthesis of peptidoglycan.[13]

There are broadly speaking two different types of cell wall in bacteria, called Gram-positive and Gram-negative. The names originate from the reaction of cells to the Gram stain, a test long-employed for the classification of bacterial species.[14]

Gram-positive bacteria possess a thick cell wall containing many layers of peptidoglycan and teichoic acids. In contrast, Gram-negative bacteria have a relatively thin cell wall consisting of a few layers of peptidoglycan surrounded by a second lipid membrane containing lipopolysaccharides and lipoproteins. Most bacteria have the Gram-negative cell wall and only the Firmicutes and Actinobacteria (previously known as the low G+C and high G+C Gram-positive bacteria, respectively) have the alternative Gram-positive arrangement.[15] These differences in structure can produce differences in antibiotic susceptibility, for instance vancomycin can kill only Gram-positive bacteria and is ineffective against Gram-negative pathogens, such as Haemophilus influenzae or Pseudomonas aeruginosa.[16]

Archaeal cell walls

Although not truly unique, the cell walls of Archaea are unusual. Whereas peptidoglycan is a standard component of all bacterial cell walls, all archaeal cell walls lack peptidoglycan,[17] with the exception of one group of methanogens.[1] In that group, the peptidoglycan is a modified form very different from the kind found in bacteria.[17] There are four types of cell wall currently known among the Archaea.

One type of archaeal cell wall is that composed of pseudopeptidoglycan (also called pseudomurein). This type of wall is found in some methanogens, such as Methanobacterium and Methanothermus.[18] While the overall structure of archaeal pseudopeptidoglycan superficially resembles that of bacterial peptidoglycan, there are a number of significant chemical differences. Like the peptidoglycan found in bacterial cell walls, pseudopeptidoglycan consists of polymer chains of glycan cross-linked by short peptide connections. However, unlike peptidoglycan, the sugar N-acetylmuramic acid is replaced by N-acetyltalosaminuronic acid,[17] and the two sugars are bonded with a β,1-3 glycosidic linkage instead of β,1-4. Additionally, the cross-linking peptides are L-amino acids rather than D-amino acids as they are in bacteria.[18]

A second type of archaeal cell wall is found in Methanosarcina and Halococcus. This type of cell wall is composed entirely of a thick layer of polysaccharides, which may be sulfated in the case of Halococcus.[18] Structure in this type of wall is complex and as yet is not fully investigated.

A third type of wall among the Archaea consists of glycoprotein, and occurs in the hyperthermophiles, Halobacterium, and some methanogens. In Halobacterium, the proteins in the wall have a high content of acidic amino acids, giving the wall an overall negative charge. The result is an unstable structure that is stabilized by the presence of large quantities of positive sodium ions that neutralize the charge.[18] Consequently, Halobacterium thrives only under conditions with high salinity.

In other Archaea, such as Methanomicrobium and Desulfurococcus, the wall may be composed only of surface-layer proteins,[1] known as an S-layer. S-layers are common in bacteria, where they serve as either the sole cell-wall component or an outer layer in conjunction with peptidoglycan and murein. Most Archaea are Gram-negative, though at least one Gram-positive member is known.[1]

See also

References

  1. ^ a b c d Howland, John L. (2000). The Surprising Archaea: Discovering Another Domain of Life. Oxford: Oxford University Press. pp. 69–71. ISBN 0-19-511183-4. Cite error: The named reference "Howland 2000" was defined multiple times with different content (see the help page).
  2. ^ Laurence Moire, Alain Schmutz, Antony Buchala, Bin Yan, Ruth E. Stark, and Ulrich Ryser (1999). "Glycerol Is a Suberin Monomer. New Experimental Evidence for an Old Hypothesis". Plant Physiol. 119: 1137-1146
  3. ^ Buchanan (2000). Biochemistry & molecular biology of plants (1st ed. ed.). American society of plant physiology. ISBN 0-943088-39-9. {{cite book}}: |edition= has extra text (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  4. ^ Wilson, Brayton F. (1984). The Growing Tree (revised ed. ed.). Amherst: University of Massachusetts Press. pp. 114–115. ISBN 0-87023-424-2. {{cite book}}: |edition= has extra text (help)
  5. ^ Sendbusch, Peter V. (2003-07-31). "Cell Walls of Algae". Botany Online. Retrieved on 2007-10-29.
  6. ^ Raven, J. A. (1983). The transport and function of silicon in plants. Biol. Rev. 58, 179-207.
  7. ^ Furnas, M. J. (1990). "In situ growth rates of marine phytoplankton : Approaches to measurement, community and species growth rates". J. Plankton Res. 12, 1117-1151.
  8. ^ Hudler, George W. (1998). Magical Mushrooms, Mischievous Molds. Princeton, NJ: Princeton University Press, 7. ISBN 0-691-02873-7.
  9. ^ Sengbusch, Peter V. (2003-07-31). "Interactions between Plants and Fungi: the Evolution of their Parasitic and Symbiotic Relations". biologie.uni-hamburg.de. Retrieved on 2007-10-29.
  10. ^ a b Alexopoulos, C. J., C. W. Mims, & M. Blackwell (1996). Introductory Mycology 4. New York: John Wiley & Sons, 687-688. ISBN 0-471-52229-5.
  11. ^ a b Raper, Kenneth B. (1984). The Dictyostelids. Princeton, NJ: Princeton University Press, 99-100. ISBN 0-691-08345-2.
  12. ^ van Heijenoort J (2001). "Formation of the glycan chains in the synthesis of bacterial peptidoglycan". Glycobiology. 11 (3): 25R–36R. PMID 11320055.
  13. ^ a b Koch A (2003). "Bacterial wall as target for attack: past, present, and future research". Clin Microbiol Rev. 16 (4): 673–87. PMID 14557293.
  14. ^ Gram, HC (1884). "Über die isolierte Färbung der Schizomyceten in Schnitt- und Trockenpräparaten". Fortschr. Med. 2: 185–189.
  15. ^ Hugenholtz P (2002). "Exploring prokaryotic diversity in the genomic era". Genome Biol. 3 (2): REVIEWS0003. PMID 11864374.
  16. ^ Walsh F, Amyes S (2004). "Microbiology and drug resistance mechanisms of fully resistant pathogens". Curr Opin Microbiol. 7 (5): 439–44. PMID 15451497.
  17. ^ a b c White, David. (1995) The Physiology and Biochemistry of Prokaryotes, pages 6, 12-21. (Oxford: Oxford University Press). ISBN 0-19-508439-X.
  18. ^ a b c d Brock, Thomas D., Michael T. Madigan, John M. Martinko, & Jack Parker. (1994) Biology of Microorganisms, 7th ed., pages 818-819, 824 (Englewood Cliffs, NJ: Prentice Hall). ISBN 0-13-042169-3.
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