Ugrás a tartalomhoz

Plant Physiology

Ördög Vince, Molnár Zoltán (2011)

Debreceni Egyetem, Nyugat-Magyarországi Egyetem, Pannon Egyetem

Chapter 4. Physiology of plant growth and development

Chapter 4. Physiology of plant growth and development

Cell wall biogenesis and expansion

The structure and biosynthesis of plant cell wall

The cell walls of prokaryotes, fungi, algae, and plants are distinctive from each other in chemical composition and microscopic structure, yet they all serve two common primary functions: regulating cell volume and determining cell shape. Because of these diverse functions, the structure and composition of plant cell walls are complex and variable.

In addition to these biological functions, the plant cell wall is important in human economics. As a natural product, the plant cell wall is used commercially in the form of paper, textiles, fibers (cotton, flax, hemp, and others), charcoal, lumber, and other wood products. Another major use of plant cell walls is in the form of extracted polysaccharides that have been modified to make plastics, films, coatings, adhesives, gels, and thickeners in a huge variety of products.

As the most abundant reservoir of organic carbon in nature, the plant cell wall also takes part in the processes of carbon flow through ecosystems.The organic substances that make up humus in the soil and that enhance soil structure and fertility are derived from cell walls. Finally, as an important source of roughage in our diet, the plant cell wall is a significant factor in human health and nutrition.

The architecture, mechanics and function of plants depend on the structure of the cell wall

In stained sections of plant tissues reveal that the cell wall is not uniform, but varies greatly in appearance and composition in different cell types. Cell walls of the cortical parenchyma are generally thin and have few distinguishing features. In contrast, the walls of some specialized cells, such as epidermal cells, collenchyma, phloem fibers, xylem tracheary elements, and other forms of sclerenchyma have thicker, multilayered walls. Often these walls are intricately sculpted and are impregnated with specific substances, such as lignin, cutin, suberin, waxes, silica, or structural proteins.

Despite this diversity in cell wall morphology, cell walls commonly are classified into two major types: primary walls and secondary walls. Primary walls are formed by growing cells and are usually considered to be relatively unspecialized and similar in molecular architecture in all cell types. Nevertheless, the ultrastructure of primary walls also shows wide variation. Some primary walls, such as those of the onion bulb parenchyma, are very thin (100 nm) and architecturally simple. Other primary walls, such as those found in collenchyma or in the epidermis, may be much thicker and consist of multiple layers (Figure 3.1).

Figure 3.1 Diversity of plant cell wall structure. (A) primary, and (B)-(C) secondary cell walls (source: Taiz L., Zeiger E., 2010)

Secondary walls are the cell walls that form after cell growth (enlargement) has ceased. Secondary walls may become highly specialized in structure and composition, reflecting the differentiated state of the cell. Xylem cells, such as those found in wood, are notable for possessing highly thickened secondary walls that are strengthened by lignin.

A thin layer of material, the middle lamella (plural lamellae), can usually be seen at the junction where the walls of neighboring cells come into contact. The composition of the middle lamella differs from the rest of the wall in that it is high in pectin and contains different proteins compared with the bulk of the wall. Its origin can be traced to the cell plate that formed during cell division.

The cell wall is usually penetrated by tiny membrane-lined channels, called plasmodesmata (singular plasmodesma), which connect neighboring cells. Plasmodesmata function in communication between cells, by allowing passive transport of small molecules and active transport of proteins and nucleic acids between the cytoplasms of adjacent cells.

Primary cell wall is a network of cellulose microfibrils embedded in a matrix of hemicelluloses, pectins, and structural proteins

In primary cell walls, cellulose microfibrils are embedded in a highly hydrated matrix. This structure provides both strength and flexibility. In the case of cell walls, the matrix consists of two major groups of polysaccharides, usually called hemicelluloses and pectins, plus a small amount of structural protein.

Cellulose microfibrils are relatively stiff structures that contribute to the strength and structural bias of the cell wall. The individual glucans that make up the microfibril are closely aligned and bonded to each other to make a highly ordered (crystalline) ribbon that excludes water and is relatively inaccessible to enzymatic attack. As a result, cellulose is very strong and very stable and resists degradation (Figure 3.2).

Figure 3.2 A structural model of a cellulose microfibril (source: Taiz L., Zeiger E., 2010)

Hemicelluloses are flexible polysaccharides that characteristically bind to the surface of cellulose. They may form tethers that bind cellulose microfibrils together into a cohesive network, or they may act as a slippery coating to prevent direct microfibril–microfibril contact. Another term for these molecules is cross-linking glucans. The term hemicellulose includes several different kinds of polysaccharides.

Pectins form a hydrated gel phase in which the cellulose–hemicellulose network is embedded. They act as hydrophilic filler, to prevent aggregation and collapse of the cellulose network. They also determine the porosity of the cell wall to macromolecules. Like hemicelluloses, pectins include several different kinds of polysaccharides.

The precise role of wall structural proteins is uncertain, but they may add mechanical strength to the wall and assist in the proper assembly of other wall components. The primary wall is composed of approximately 25% cellulose, 25% hemicelluloses, and 35% pectins, with perhaps 1 to 8% structural protein, on a dry-weight basis. However, large deviations from these values may be found.

The primary wall also contains much water. This water is located mostly in the matrix, which is perhaps 75 to 80% water. The hydration state of the matrix is an important determinant of the physical properties of the wall; for example, removal of water makes the wall stiffer and less extensible.

Secondary walls in woody tissues contain more cellulose, xylans, and lignin

After wall expansion ceases, cells sometimes continue to synthesize a wall, known as a secondary wall. Secondary walls are often quite thick, as in tracheids, fibers, and other cells that serve in mechanical support of the plant. Often such secondary walls are multilayered and differ in structure and composition from the primary wall. For example, the secondary walls in wood contain xylans rather than xyloglucans, as well as a higher proportion of cellulose. The orientation of the cellulose microfibrils may be more neatly aligned parallel to each other in secondary walls than in primary walls. Secondary walls are often (but not always) impregnated with lignin.

Lignin is a phenolic polymer with a complex, irregular pattern of linkages that link the aromatic alcohol subunits together. These subunits are synthesized from phenylalanine and are secreted to the wall, where they are oxidized in place by the enzymes peroxidase and laccase. As lignin forms in the wall, it displaces water from the matrix and forms a hydrophobic network that bonds tightly to cellulose and prevents wall enlargement.

Cell wall elongation and degradation

During plant cell enlargement, new wall polymers are continuously synthesized and secreted at the same time that the preexisting wall is expanding.

Microfibril orientation influences growth directionality of cells with diffuse growth

During growth, the loosened cell wall is extended by physical forces generated from cell turgor pressure. Turgor pressure creates an outward-directed force, equal in all directions. The directionality of growth is determined largely by the structure of the cell wall – in particular, the orientation of cellulose microfibrils.

When cells first form in the meristem, they are isodiametric; that is, they have equal diameters in all directions. If the orientation of cellulose microfibrils in the primary cell wall were isotropic (randomly arranged), the cell would grow equally in all directions, expanding radially to generate a sphere (Figure 3.3). In most plant cell walls, however, the arrangement of cellulose microfibrils is anisotropic (nonrandom), or aligned in a preferred direction.

Figure 3.3 The orientation of newly deposited cellulose microfibrils determines the direction of cell expansion (source: Taiz L., Zeiger E., 2010)

Acid-induced cell wall extension is characteristic of primary walls and is mediated by the protein expansin

An important characteristic of growing cell walls is that they extend much faster at acidic pH than at neutral pH. This phenomenon is called acid growth. In living cells, acid growth is evident when growing cells are treated with acid buffers or with the drug fusicoccin, which induces acidification of the cell wall solution by activating an H+-ATPase in the plasma membrane. Auxin-induced growth is also associated with wall acidification, but it is probably not sufficient to account for the entire growth induction by this hormone, and other wall-loosening processes may be involved.

The idea that proteins are required for acid growth was confirmed in reconstitution experiments, in which heat inactivated walls were restored to nearly full acid growth responsiveness by addition of proteins extracted from growing walls. The active components proved to be a group of proteins that were named expansins. These proteins catalyze the pH-dependent extension and stress relaxation of cell walls.

Plant cell walls play a major role in carbon flow through ecosystems

Most plant cell walls are constructed in a way to resist enzymatic digestion – a defense against pathogen invasion – and so the recycling of the carbon and energy locked in the cell wall is mostly carried out by saprophytic fungi and bacteria armed with a suite of specialized enzymes capable of digesting cell walls. Some animals, such as ruminants and termites, are also able to partake in this fibrous feast, with the aid of gut microbes similarly equipped with cell-wall digesting enzymes. The organic substances that make up humus in the soil and that enhance soil structure and fertility are derived from cell wall residues – one of many legacies of plants to their environment. Finally, as the major source of dietary fiber, the plant cell wall is a significant factor in human health and nutrition.

Cell wall degradation and plant defense

The plant cell wall is not simply an inert, static exoskeleton. In addition to acting as a mechanical restraint, the wall serves as an extracellular matrix that interacts with cell surface proteins, providing positional and developmental information. It contains numerous enzymes and smaller molecules that are biologically active and that can modify the physical properties of the wall, sometimes within seconds. In some cases, wall-derived molecules can also act as signals to inform the cell of environmental conditions, such as the presence of pathogens. This is an important aspect of the defense response of plants.

Walls may also be substantially modified long after growth has ceased. For instance, the cell wall may be selectively degraded, such as occurs in ripening fruit or in the endosperm or cotyledons of germinating seeds. In cells that make up the abscission zones of leaves and fruits, the middle lamella is digested, with the result that the cells become unglued and separate. Cells may also separate selectively during the formation of intercellular air spaces, during the emergence of the root from germinating seeds, and during other developmental processes. Plant cells may also modify their walls during pathogen attack as a form of defense.