Cellulose: Structure and Distribution

Bruce Stone, La Trobe University, Bundoora, Victoria, Australia

Cellulose, a (1!4)-b-glucan, is the most abundant carbohydrate polymer in the biosphere, where it may account for 50% of the carbon. Cellulose has a structural and protective function in walls of plant cells and surfaces of other organisms. The insolubility and high textile strength of cellulosic materials arises from the regular, extended, ribbon-like conformation of the individual molecules, their enormous lengths and their ability to aggregate into crystalline microfibrils.

Secondary article
Article Contents
. Introduction . Structure . Distribution

Introduction
Cellulose is a ubiquitous structural component of cell walls of embryophytes and many algae. It is a major component of wood and plant bres used in textiles. Cellulose is also produced by some bacteria and protista, and in the animal kingdom by a solitary group of invertebrates, the tunicates. It is not found in the major Archaea and very rarely in the Fungi. Cellulose is a major carbohydrate and energy source for many microorganisms and herbivorous animals.

polymerization (DPav) of the cellulose molecules depends on the source (Table 1).

Conformation
The spatial organization of the cellulose molecule (Figure 1) is governed by the equatorial congurations of the C1 and C4 hydroxyls, giving a diequatorial (1e!4e), glycosidic linkage. Free rotation around the C1O and C4 O bonds in the glycosidic linkages is limited by van der Waals repulsions between the glucose units. These are minimal when the torsion angles, f and j (Figure 1), for the C1O and C4 O bonds are 2 988 and 2 1438 respectively. The predicted regular chain conformation of a cellulose molecule, determined by these angles, is helical with a repeat of 1.03 nm. The C6 hydroxymethyl groups on alternate glucose units are on opposite sides of the cellulose chain. Thus the molecule may be looked upon as a polymer of the disaccharide cellobiose (Figure 1) although crystallographic studies show that glucose is the true repeating unit. Overall, the cellulose molecule has a twisted-ribbon conformation. Cellulose, although apparently hydrophilic, is completely insoluble in water. This property results from the sideby-side association of individual cellulose molecules to form aggregates in which the molecules are packed regularly with their hydrogen bonding capacities internally

Structure
Chemistry
The molecule of cellulose is described as a (1!4)-b-d glucan (Figure 1). It is an unbranched chain of b-d glucopyranose units in the chair conformation, designated 4 C1 (carbon 4 high and carbon 1 low), which has all its hydroxyl (OH) and hydromethyl (CH2OH) substituents equatorial. The glucose units are joined by glycosidic linkages between the hemiacetal hydroxyl at C1 on one residue and the hydroxyl at C4 on the next residue with the loss of the elements of water. The average degree of

HO HO

CH2OH O OH

OH HO O CH2OH O

4 O HO 3

CH2OH O
5 2

OH HO O
4

OH O CH2OH
HEMIACETAL HYDROXYL

OH 1

NON-REDUCING END

CELLOBIOSE REPEAT
n

REDUCING END

Figure 1 (a) A (1!4)-b-D-glucan molecule terminated by a reducing end bearing a free (unsubstituted) hemiacetal hydroxyl (on the right) and a nonreducing end (on the left). The hydroxymethyl groups at C6 of alternate glucose residues are on opposite sides of the chain. Each carbon atom of the glucose ring carries an axial hydrogen atom. For clarity these are omitted.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

Table 1 Degree of polymerization of celluloses from various sources Acanthamoeba (a single-celled protistan) Embryophyte primary walls Embryophyte secondary walls Boergesenia forbesi (a siphonocladean green alga) 20006000 20006000 700014 000 23 000

satised. X-ray and electron diraction diagrams of native cellulose bres are sharp, indicative of the presence of virtually crystalline regions. The dimensions of the crystallographic unit cell derived from the measured diraction spacings shows a very narrow helix with two glucose units per turn (two-fold helix) and a repeat distance is 1.03 nm as predicted by conformational analysis, with the bre axis passing through the long axis of the glucose units. The polymer chains are packed into sheets which are stabilized by noncovalent interactions within and between adjacent chains (Figure 2a). Intrachain hydrogen bonding, parallel to the chain axis, occurs between O3 and O5 and between O6 and O2 on successive chain units (Figure 2b). There are also interchain hydrogen bonds between O3 and O6 of glucose units on adjacent chains in the sheets (Figure 2b). No interlayer hydrogen bonding occurs, but the sheets associate to form stacks through van der Waals interactions at the hydrophobic surfaces created by the axial hydrogens at carbons 15, above and below the plane of the chain.

Polymorphic forms of cellulose

Four crystalline allomorphs of cellulose are recognized from X-ray diraction and infrared spectroscopy in which minimum steric repulsion between chains, combined with maximum hydrogen bonding, provides the best t with the

diraction intensity data. The native cellulose allomorph is referred to as cellulose I. In this allomorph, all chains are oriented in the same direction and are said to be parallel. Two suballomorphs of cellulose I have been recognized by high-resolution, solid-state 13C nuclear magnetic resonance (NMR) spectroscopy and Fourier transform infrared spectroscopy. In allomorph Ia the structure corresponds to a triclinic unit cell with a single chain, whereas in Ib it corresponds to a monoclinic unit cell with two chains (Figures 3 and 4). The Ia allomorph is converted into the Ib allomorph by heating above 2608C. Thus cellulose Ia is a metastable crystalline structure. This interconversion may occur by slipping of the hydrogenbonded cellulose sheets when the intersheet interactions are disrupted. Swelling of cellulose I in concentrated ( 4 14%) sodium hydroxide solution and removal of the base (mercerization) or derivatization with carbon disulphide to form the soluble cellulose xanthate and regeneration in alkali, recrystallizes it to the cellulose II form found in cellophane and rayon. Rayon bres have increased lustre and bind dyes better. In cellulose II, the chains have an antiparallel arrangement and there is extensive intrasheet hydrogen bonding conferring the greatest thermodynamic stability on this allomorph. The green alga Halicystis (Sisson, 1941) and a mutant Acetobacter xylinum (Kuga et al., 1993) produce the cellulose II allomorph. Other allomorphs,

Figure 2 Crystal structure of cellulose I. (a) Projection down the fibre axis (c) showing the layers (sheets) of cellulose chains hydrogen-bonded in the ac plane but lacking intersheet hydrogen bonding. The intersheet bonding involves van der Waals interactions between the hydrophobic faces of the glucose units. (b) View of a layer approximately perpendicular to the ac plane, showing the two intramolecular hydrogen bonds in the direction of the fibre axis (c) and the interchain hydrogen bonds. Reproduced from Kroon-Batenburg LMJ and Kroon J (1995) Carbohydrates in Europe No.12: 15 19.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

Valonia

Micrasterias

Halocynthia

10 nm Ramie Wood Primary wall

Figure 3 Schematic representation of the mode of packing in the unit cell of cellulose I. (a) Triclinic unit cell. (b) Monoclinic unit cell. The monoclinic angle is obtuse. Reproduced from Koyama M, Helbert W, Imai T, Sugiyama J and Henrissat B (1997) Proceedings of the National Academy of Sciences of the USA 94: 9091 9095.

Figure 5 (a) Schematic representations of the cross-sections of typical cellulose microfibrils, ranging from Valonia cellulose to primary wall cellulose. Reproduced from Chanzy H (1990). In: Kennedy JF, Phillips GO and Williams PA (eds) Cellulose: Sources and Exploitation, Industrial Utilization, Biotechnology and Physicochemical Properties, pp. 3 12. New York: Ellis Horwood.

Cellobiose

0.62 nm/0.53 nm

67/63

Monoclinic I

Triclinic I

Figure 4 Schematic diagram showing the differences between the monoclinic and triclinic forms of cellulose I. Each rectangle represents a single glucose unit, with a pair of glucose units (surrounded by a dotted box) constituting the cellobiose repeat due to the two-fold screw symmetry along the chain axis (vertical in the diagram). In the monoclinic form cellobiose units stagger with a shift of a quarter of the c axis period (0.26 nm), whereas the triclinic form exhibits a diagonal shift of the same amount. The different spacings and angles shown depend on which crystallographic face is being viewed. Reproduced from Baker AA, Helbert W, Sugiyama J and Miles MJ (1997) Journal of Structural Biology 119: 129 138.

sectional dimensions depending on the source (Figure 5). It has been proposed that the broader brils are aggregates of the narrower brils, for example, the 10-nm onion microbrils are constructed from 2-nm cellulose subunits, containing 1015 chains (Ha et al., 1998), although the broad microbrils of algal cellulose are single crystals. In the large, single microbrils of native cellulose from the alga Valonia, two distinct crystalline allomorphs coexist, in the ratio Ia to Ib of 65 to 35. Microbrils of bacterial and algal celluloses are richer in the Ia phase, and some tunicate celluloses consist wholly of the Ib phase. In embryophytes, primary wall microbrils have similar proportions of the two phases but microbrils in secondary walls of dicotyledons are richer in the Ib phase. Surfaces of microcrystals of Valonia cellulose treated with acid have been imaged using the atomic force microscope (Baker et al., 1997) (Figure 6) revealing the periodicity due to the cellobiose repeat in regions with triclinic characteristics. It has been proposed that the antiparallel structure of cellulose II arises from cellulose I on swelling in alkali when adjacent microbrils, themselves composed of parallel chains, merge to give an antiparallel structure (Sarko, 1986).

termed cellulose III and IV, are derived from cellulose I and II, respectively, by heat or alkali treatment but are not known in native celluloses.

Physical and chemical properties

The physical properties, chemical reactivity and biological functions of cellulose are essentially determined by the regular packing of the cellulose molecules in the microbrils. Crystalline domains may encompass the entire width of the cellulose microbril (Newman et al., 1996; Smith et al., 1998), but chains on the microbril surfaces may noncovalently bond with noncellulosic (1!4)-bglycans, e.g. heteroglucans, heteroxylans and heteromannans in cell walls. These associations are important in stabilizing cell wall structure.
3

Microfibrillar organization
When viewed in the electron microscope, native cellulose preparations appear as brils, referred to as microbrils. Their dimensions are uniform in celluloses from any one source, but vary widely amongst dierent organisms and at dierent stages within one cell type, or developmental stages of the same organism. Microbrils vary in cross-

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

Figure 6 Atomic force microscope image of the surface of an acid-treated Valonia microfibril. The arrows at the top of the image point along the cellulose molecules, which are running almost vertically down the page. The dotted white box highlights an area where bright spots can be seen along the length of the molecules, separated by a distance closely matching the cellobiose repeat interval. The angle of the spots within the box to the molecular axis is 64 + 28. Reproduced from Baker AA, Helbert W, Sugiyama J and Miles MJ (1997) Journal of Structural Biology 119: 129 138.

For its density (1.5 Mg m 2 3) cellulose brils are stier (Young modulus 50130 GPa) and stronger (1 GPa), when measured along the polymer length, than nylon, silk, chitin, collagen, tendon or bone (Ashby et al., 1995). Crystalline cellulose is resistant to hydrolysis by hot dilute acids but is more readily hydrolysed after the chains are made accessible by swelling in concentrated acids. Most celluloses resist dissolution in acetic acidnitric acid (Updegra reagent) although Acetobacter cellulose is partially soluble and chitin does not dissolve, so without supplementary evidence this test is not diagnostic. In the presence of bases, the very weakly acidic, hydroxyl groups (pKa 1112) ionize and due to repulsion between the ionized groups, the regular organization of the chains is disrupted. Thus in 1822% sodium hydroxide cellulose swells but does not dissolve. Aprotic reagents such as N4

methylmorpholino-N-oxide and lithium chloride in dimethylacetamide are good cellulose solvents. In the presence of oxygen, cellulose in alkaline conditions is progressively depolymerized from the reducing end by the alkaline-peeling reaction. The reaction can be blocked by reducing the terminal glucose residue to the acyclic glucitol. The hydroxyl groups on cellulose can be derivatized, to varying degrees, by esterication to form, for example, the thermoplastic acetates and the plastic and explosive nitrates (gun cotton), which are soluble in organic solvents such as acetone. Water-soluble methyl, hydroxyethyl and carboxymethyl esters of cellulose are used widely in commerce as thickeners and emulsiers. Cellulose stains red-violet with zinc chloride in I2/KI (iodine/potassium iodide) and can be recognized in tissues by uorescence induced by ultraviolet (UV) light when

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

associated with uorochromes such as Calcouor White (Tinopal LPW) (a diaminostilbene derivative) or Congo Red (a diphenyldiazo derivative). Again, this test is not diagnostic since other b-glycans such as chitin, callose and certain bacterial extracellular polysaccharides, such as xanthan, also induce uorescence.

Distribution
Bacteria
Cellulose is produced extracellularly by Gram-negative bacteria such as Acetobacter xylinum, Agrobacterium tumefaciens, Rhizobium spp., Zymomonas mobilis, Achromobacter spp., Azotobacter spp., Aerobacter spp. and Pseudomonas spp. In unaerated, liquid cultures A. xylinum produces cellulose as a thick, translucent pellicle at the air medium interface. Strains of A. xylinum also produce ne cellulose microbrils in aerated, submerged cultures. The cellulose is attached parallel to the longitudinal axis of the cell as a ribbon composed of microbrils (Figure 7). The microbrils are extruded from a linear row of pores along one side of the cell and associate to form the ribbon. Microbrils of cellulose formed by symbiotic and tumorogenic Rhizobiaceae attach bacterial cells to host plant cell surfaces. Cells of the anaerobic, Gram-positive coccus Sarcina ventriculi, a soil- and stomach-inhabiting species, are enveloped in a thick (150200 nm) brous layer of cellulose that cements the cells together in large packets.

Zygnematales) have walls with compositions resembling the primary wall of embryophytes, with cellulose as a prominent component. In the much studied single-celled green alga, Valonia ventricosa (Siphonocladales) and Cladophora and Chaetomorpha (Cladophorales) the broad microbrils are arranged in sheets with near-perfect, parallel arrangement of the microbrils in each layer and in dierent orientations in successive layers (Figure 8). Each microbril consists of 12001400 cellulose chains. Among the conjugating green algae, the single-celled desmids have bilayered walls with cellulose in both layers.

Embryophytes
Cellulose is a component of all but a few specialized embryophyte cell walls. In the thin, semirigid, primary wall that overlays the plasma membrane of dividing, undierentiated cells in apical meristems of growing shoots and roots and lateral meristems (cambium), cellulose microbrils are found embedded in a matrix of noncellulosic, heteropolysaccharides (e.g. pectins, heteroglucans, heteroxylans) and proteins. In the plant, these walls contain  70% water. In some primary walls, for example, endosperm walls of grasses, cellulose contributes as little as 23% of the dry weight of the wall, in other primary walls as much as 40%. In newly divided cells, the microbrils are very narrow (2 nm) (Table 2). The microbrils are deposited in lamellae of which there may be only about four in the thin primary wall. Next to the plasma membrane, the site of microbril synthesis, the preferred orientation is at right angles to the direction of growth and the ordering is not high. In the rst-formed (outer) parts of primary walls, next to the middle lamella, the predominant axis of orientation of the microbrils is along the direction of growth. In supporting tissues of herbaceous stems and petioles, the cells of the soft and exible (pliable) collenchyma tissue have thickened primary walls with layers in which the microbrils have a predominantly transverse orientation alternating with those whose orientation is predominantly longitudinal. Electron microscopy of primary walls of onion epidermal cells (McCann and Roberts, 1992) has revealed that the microbrils are interconnected laterally, possibly by heteroglucans and/or heteroxylans that bond noncovalently with the cellulose microbrils. Pectins and proteins may form additional wall networks. During wall elongation, enzymic and nonenzymic mechanisms are employed to loosen the networks permitting insertion of new wall material. Wall stability is then restored by creating new interconnections. In cells that dierentiate into tracheids and sclerenchyma bres, the principal cellular components of wood, a secondary wall layer is deposited inside the primary wall after the elongation phase of wall growth has been completed. The secondary walls are rich in cellulose (35
5

Protista
In Acanthamoeba (Rhizopoda: Acanthopodina), the single cells form a polyhedral or thickly biconvex cyst having a wall that contains microbrillar cellulose. In the singlecelled coccolithophorids, for example Pleurochrysis scherffelii, the cell surface is covered by calcied coccoliths (scales) with a base plate that includes microbrillar cellulose. The coccoliths are produced in the Golgi apparatus in some species and in a specialized reticulate body in others. In the dinoagellates the cell covering, called an amphisema, is located beneath the plasma membrane. The amphisema is made up of cellulosic plates, and is produced in vesicles lying beneath the plasma membrane. In the water moulds Sarprolegnia and Achlya (Oomycota) the hyphal walls contain cellulose and (1!3)b-glucan.

Algae
Cellulose has been identied in the walls of brown algae (Phaeophyta), red algae (Rhodophyta), Chrysophyta, stoneworts (Charophyta) and Xanthophyta. In the diverse green algae (Chlorophyta), some orders (Chlorococcales,

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

Figure 7 Microfibrils of Acetobacter xylinum cellulose associated with the surface of a bacterial cell. (a) Cell surface showing the ribbon of cellulose growing thicker along the cell length as discrete bundles with separate points of origin (arrows) accumulate. (b) Freeze-etch replica of fractured bacterial lipopolysaccharide surface region showing a region of overlap of two separate rows of synthesizing sites. Reproduced from Haigler CH and Benziman M (1982). In: Brown RM Jr (ed.) Cellulose and Other Natural Polymer Systems. Biogenesis, Structure and Degradation, pp. 273 297. New York: Plenum.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

Figure 8 A fracture plane through the wall of an 18-day-old Valonia ventricosa cell showing regular but different orientations of the microfibrils in two lamellae. Reproduced from Itoh T and Brown RM Jr (1984) Planta 160: 372 381.

Table 2 Widths of cellulose microbrils from various sources Wood cambium Quince seed mucilage Acetobacter xylinum Embryophyte primary walls Embryophyte secondary walls Tunicate tests (glomerulocytes) Ramie (Boehmeria nivea) Boergesenia forbesii (a siphonocladean green alga) Valonia ventricosa (a unicellular green alga)  1.5 nm 2 nm 2 nm 2 nm 510 nm 1216 nm 1020 nm 30 nm 50200 nm
S3 S23 S2 S12 S1 P1 P0

60% dry weight) and usually consist of three distinct layers of precisely parallel microbrils, whose orientation is dierent in successive layers (Figure 9). Noncellulosic polysaccharides form the matrix of the secondary wall, and in the later stages of wall dierentiation, lignins are deposited in the matrix of both the primary and secondary wall enveloping the cellulosic microbrils and conferring resistance to compression. The commercially important seed hairs of cotton (Gossypium hirsutum), have unlignied secondary walls that, at maturity, contain more than 95% cellulose. Noncellulosic polysaccharides including the (1!3)-bglucan, callose and waxes make up the remaining material. The microbrils are broad and the DP is extremely high (Table 1). In G. asiaticum and G. arboreum the seed hairs are slightly lignied. The commercial bre kapok consists of the cellulosic fruit hairs of Ceiba pentaudra. Other commercially important cellulose sources are the long, extra-xylary, soft, bast bre bundles from the dicotyledons jute (Corchorus spp.), hemp (Cannabis sativa), ax (Linum usitatissimum), kenaf (Hibiscus cannabinus) and ramie (Boehmeria nivea) and the hard, leaf bres from the

Figure 9 Microfibril orientation in the secondary wall layers of a fibre cell. In S1, the outermost layer (next to the primary wall layer, P) the microfibrils are usually in a slow helix (relatively transverse), whereas in the S2 layer they are relatively longitudinal. The microfibrils in the S3 layer are again more transverse in orientation. The interfaces between the three layers have a series of thin lamellae (one microfibril thick) in which the orientation changes by a small angle in each successive layer. Reproduced from Fujita M and Harada H (1991) Ultrastructure and formation of wood cell wall. In: Hon DN-S and Shirashi N (eds) Wood and Cellulosic Chemistry, pp. 3 57. New York: Dekker.

monocotyledons abaca (Musa textilis), sisal (Agave sisalana) and other agaves.

Slime moulds
In cellular slime moulds (Dictyostelium discoideum, Polyspondylium sp. (Dictyostelia: Acrasiomycota) that dierentiate during reproduction, microbrillar cellulose occurs in the stalk tube surrounding the stalk cell population, in the stalk cell and spore walls, and in the sheath surrounding the slug, which as it migrates leaves the collapsed sheath behind as a slime trail. Nondierentiating amoebae do not
7

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

form cellulose, and mutants lacking cellulose cannot complete their developmental programme (life cycle) as they cannot form normal stalks.

myces and Blastocladiella (Chytridiomycota) are reported also to contain cellulose.

Fungi
Exceptionally amongst Fungi the chitinous walls of the parasitic Rhizidomyces (Hypochytridiomycota), Allo-

Tunicates
The body of adult tunicates (Tunicata: Urochordata: Animalia) is covered by a thick external supportive and protective skeleton, the tunic or test, which is secreted by a

Figure 10 Coiled microfibril bundles in the glomerulocytes in the epidermis beneath the test of the ascidian Metandrocarpa uedai. Reproduced from Kimura S and Itoh T (1995) Protoplasma 186: 24 33.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Cellulose: Structure and Distribution

single layer of epidermal cells. The tunic is 90% water and the major constituents are cellulose (60%) and protein (27%). The microbrils are wide and highly crystalline and appear to provide strength to the tunic. The microbrils are not arranged in lamellae as in embryophyte walls. In Polyandrocarpa misakiensis, the tunic cord connecting the dorsal tunic and internal tunic of the siphon is constructed of bundled, highly crystalline microbrils covered by a single layer of epidermal cells which are the site of their synthesis (Kimura and Itoh, 1998). In compound, styelid ascidians a network of cellulose microbrils is also found in the haemocoel, a connective tissue layer lying beneath the tunic, between the epidermis and aerial epithelium (Kimura and Itoh, 1997). Here the microbrils may play the same structural role as collagen bres in the connective tissue of higher chordates. Glomerulocytes found in the epidermis beneath the test of the ascidian Metandrocarpa uedai are responsible for the synthesis and deposition of cellulose (Kimura and Itoh, 1995). Bundles of microbrils seen in the glomerulocyte (Figure 10) have the diraction pattern of cellulose I.

Kimura S and Ito T (1995) Evidence for the role of glomerulocytes in cellulose synthesis in the tunicate, Melandrocarpa keda. Protoplasma 186: 2433. Kimura S and Itoh T (1997) Cellulose network of hemocoel in selected compound styelid ascidians. Journal of Electron Microscopy 46: 327 335. Kimura S and Itoh T (1998) A new cellulosic structure, the tunic cord in the ascidian Polyandrocarpa misakiensis. Protoplasma 204: 94102. Kuga S, Takagi S and Brown RM Jr (1993) Native folded chain cellulose II. Polymer 34: 32933297. McCann MC and Roberts K (1992) Changes in cell wall architecture during cell elongation. Journal of Experimental Botany 45: 16831691. Newman RH, Davies LM and Harris PJ (1996) Solid-state 13C nuclear magnetic resonance characterization of cellulose in the cell walls of Arabidopsis thaliana leaves. Plant Physiology 111: 475485. Sarko A (1986) Recent X-ray crystallographic studies on cellulose. In: Young RA and Rowell RM (eds) Cellulose: Structure, Modication and Hydrolysis, pp. 2949. New York: Wiley-Interscience. Sisson WA (1941) Some X-ray observations regarding the membrane structure of Haliocystis. Contributions of the Boyce Thompson Institute 12: 3144. Smith BG, Harris PJ, Melton LD and Newman RH (1998) Crystalline cellulose in hydrated primary cell walls of three monocotyledons and one dicotyledon. Plant and Cell Physiology 39: 711720.

References
Ashby MF, Gibson LJ, Wegst U and Olive R (1995) The mechanical properties of natural materials I. Material property charts. Proceedings of the Royal Society (London) A 450: 123140. Baker AA, Helbert W, Sugiyama J and Miles M J (1997) High-resolution atomic force microscopy of native Valonia cellulose I microcrystals. Journal of Structural Biology 119: 129138. Ha M-A, Apperley DC, Evans BW et al. (1998) Fine structure in cellulose microbrils: NMR evidence from onion and quince. The Plant Journal 16: 183190.

Further Reading
French AD (2000) The structure and biosynthesis of cellulose. In: Kung SD and Yang SF (eds) The Discoveries in Plant Biology Series, vol. 3, pp. 163197. Hong Kong: World Science Press. Krassig H, Steadman RG, Schliefer K and Albrecht W (1986) Cellulose. In: Ullmans Encyclopedia of Industrial Chemistry, 5th edn, pp. 375 418. Weinheim: VCH Verlagsgesellschaft. Richmond PA (1991) Occurrence and functions of native cellulose. In: Haigler CH and Weimer PJ (eds) Biosynthesis and Biodegradation of Cellulose, pp. 523. New York: Dekker.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net