Cell wall


A cell wall is a structural layer that surrounds some cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, and functions as a selective barrier. Another vital role of the cell wall is to help the cell withstand osmotic pressure and mechanical stress. Cell walls are found in most prokaryotes, with the exception of mollicute bacteria. Among the eukaryotes, cells walls are prevalent in fungi, algae and plants but absent from animals and many other taxa.
The composition of cell walls varies across taxonomic groups, species, cell type, and the cell cycle. In land plants, the primary cell wall comprises polysaccharides like cellulose, hemicelluloses, and pectin. Often, other polymers such as lignin, suberin or cutin are anchored to or embedded in plant cell walls. Algae exhibit cell walls composed of glycoproteins and polysaccharides, such as carrageenan and agar, distinct from those in land plants. Bacterial cell walls contain peptidoglycan, while archaeal cell walls vary in composition, potentially consisting of glycoprotein S-layers, pseudopeptidoglycan, or polysaccharides. Fungi possess cell walls constructed from the polymer chitin, specifically N-acetylglucosamine. Diatoms have a unique cell wall composed of biogenic silica.

History

A plant cell wall was first observed and named by Robert Hooke in 1665. However, "the dead excrusion product of the living protoplast" was forgotten, for almost three centuries, being the subject of scientific interest mainly as a resource for industrial processing or in relation to animal or human health.
In 1804, Karl Rudolphi and J.H.F. Link proved that cells had independent cell walls. Before, it had been thought that cells shared walls and that fluid passed between them this way.
The mode of formation of the cell wall was controversial in the 19th century. Hugo von Mohl advocated the idea that the cell wall grows by apposition. Carl Nägeli believed that the growth of the wall in thickness and in area was due to a process termed intussusception. Each theory was improved in the following decades: the apposition theory by Eduard Strasburger, and the intussusception theory by Julius Wiesner.
In 1930, Ernst Münch coined the term apoplast in order to separate the "living" symplast from the "dead" plant region, the latter of which included the cell wall.
By the 1980s, some authors suggested replacing the term "cell wall", particularly as it was used for plants, with the more precise term "extracellular matrix", as used for animal cells, but others preferred the older term.

Properties

Cell walls serve similar purposes in those organisms that possess them. They may give cells rigidity and strength, offering protection against mechanical stress. The chemical composition and mechanical properties of the cell wall are linked with plant cell growth and morphogenesis. In multicellular organisms, they permit the organism to build and hold a definite shape. Cell walls also limit the entry of large molecules that may be toxic to the cell. They further permit the creation of stable osmotic environments by preventing osmotic lysis and helping to retain water. Their composition, properties, and form may change during the cell cycle and depend on growth conditions.

Rigidity of cell walls

In most cells, the cell wall is flexible, meaning that it will bend rather than holding a fixed shape, but has considerable tensile strength. The apparent rigidity of primary plant tissues is enabled by cell walls, but is not due to the walls' stiffness. Hydraulic turgor pressure creates this rigidity, along with the wall structure. The flexibility of the cell walls is seen when plants wilt, so that the stems and leaves begin to droop, or in seaweeds that bend in water currents. As John Howland explains
The apparent rigidity of the cell wall thus results from inflation of the cell contained within. This inflation is a result of the passive uptake of water.

Evolution

Cell walls evolved independently in many groups.
The photosynthetic eukaryotes have cellulose cell walls, where the cell wall is closely related to the evolution of multicellularity, terrestrialization and vascularization. The CesA cellulose synthase evolved in Cyanobacteria and was part of Archaeplastida since endosymbiosis; secondary endosymbiosis events transferred it further into brown algae and oomycetes. Plants later evolved various genes from CesA, including the Csl family of proteins and additional Ces proteins. Combined with the various glycosyltransferases, they enable more complex chemical structures to be built.
Fungi use a chitin-glucan-protein cell wall. They share the 1,3-β-glucan synthesis pathway with plants, using homologous GT48 family 1,3-Beta-glucan synthases to perform the task, suggesting that such an enzyme is very ancient within the eukaryotes. Their glycoproteins are rich in mannose. The cell wall might have evolved to deter viral infections. Proteins embedded in cell walls are variable, contained in tandem repeats subject to homologous recombination. An alternative scenario is that fungi started with a chitin-based cell wall and later acquired the GT-48 enzymes for the 1,3-β-glucans via horizontal gene transfer. The pathway leading to 1,6-β-glucan synthesis is not sufficiently known in either case.

Plant cell walls

The walls of plant cells must have sufficient tensile strength to withstand internal osmotic pressures of several times atmospheric pressure that result from the difference in solute concentration between the cell interior and external solutions. Plant cell walls vary from 0.1 to several μm in thickness.
In plants, a secondary cell wall is a thicker additional layer of cellulose which increases wall rigidity. Additional layers may be formed by lignin in xylem cell walls, or suberin in cork cell walls. 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.
The primary cell wall of most plant cells is freely permeable to small molecules including small proteins, with size exclusion estimated to be. The pH is an important abiotic factor governing the transport of molecules through cell walls.

Layers

Up to three strata or layers may be found in plant cell walls:
  • The primary cell wall, generally a thin, flexible and extensible layer formed while the cell is growing.
  • 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. Some cells, such as the conducting cells in xylem, possess a secondary wall containing lignin, which strengthens and waterproofs the wall.
  • The middle lamella, a layer rich in pectins. This outermost layer forms the interface between adjacent plant cells and glues them together.

    Composition

In the primary plant cell wall, the major carbohydrates 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. In grass cell walls, xyloglucan and pectin are reduced in abundance and partially replaced by glucuronoarabinoxylan, another type of hemicellulose. Primary cell walls characteristically extend by a mechanism called acid growth, mediated by expansins, extracellular proteins activated by acidic conditions that modify the hydrogen bonds between pectin and cellulose. This functions to increase cell wall extensibility. The outer part of the primary cell wall of the plant epidermis is usually impregnated with cutin and wax, forming a permeability barrier known as the plant cuticle.
Secondary cell walls contain a wide range of additional compounds that modify their mechanical properties and permeability. The major polymers that make up wood include:
  • cellulose, 35-50%
  • xylan, 20-35%, a type of hemicellulose
  • lignin, 10-25%, a complex phenolic polymer that penetrates the spaces in the cell wall between cellulose, hemicellulose and pectin components, driving out water and strengthening the wall.
Additionally, structural proteins are found in most plant cell walls; they are classified as hydroxyproline-rich glycoproteins, arabinogalactan proteins, glycine-rich proteins, and proline-rich proteins. Each class of glycoprotein is defined by a characteristic, highly repetitive protein sequence. Most are glycosylated, contain hydroxyproline and become cross-linked in the cell wall. These proteins are often concentrated in specialized cells and in cell corners. Cell walls of the epidermis may contain cutin. The Casparian strip in the endodermis roots and cork cells of plant bark contain suberin. Both cutin and suberin are polyesters that function as permeability barriers to the movement of water. The relative composition of carbohydrates, secondary compounds and proteins varies between plants and between the cell type and age. Plant cells walls also contain numerous enzymes, such as hydrolases, esterases, peroxidases, and transglycosylases, that cut, trim and cross-link wall polymers.
Secondary walls - especially in grasses - may also contain microscopic silica crystals, which may strengthen the wall and protect it from herbivores.
Cell walls in some plant tissues also function as storage deposits for carbohydrates that can be broken down and resorbed to supply the metabolic and growth needs of the plant. For example, endosperm cell walls in the seeds of cereal grasses, nasturtium
and other species, are rich in glucans and other polysaccharides that are readily digested by enzymes during seed germination to form simple sugars that nourish the growing embryo.