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Vol.25 No.7

Biomimetic engineering of
cellulose-based materials
Tuula T. Teeri1, Harry Brumer III1, Geoff Daniel2 and Paul Gatenholm3
1

Swedish Center for Biomimetic Fiber Engineering, KTH Biotechnology, AlbaNova, SE-10691 Stockholm, Sweden
WURC, Department of Wood Sciences, Swedish University of Agricultural Sciences, Box 7008, SE-75007 Uppsala, Sweden
3
Chalmers University of Technology, Department of Chemical and Biological Engineering, Biopolymer Technology,
SE-412 96 Gothenburg, Sweden
2

Biomimetics is a field of science that investigates

biological structures and processes for their use as
models for the development of artificial systems. Biomimetic approaches have considerable potential in the development of new high-performance materials with low
environmental impact. The cell walls of different plant
species represent complex and highly sophisticated composite materials that can provide inspiration on how to
design and fabricate lightweight materials with unique
properties. Such materials can provide environmentally
compatible solutions in advanced packaging, electronic
devices, vehicles and sports equipment. This review gives
an overview of the structures and interactions in natural
plant cell walls and describes the first attempts towards
mimicking them to develop novel biomaterials.
Trends in biological material science
The development of materials has shaped the fate and
success of mankind since the development of the first tools,
made of wood and stone. Innovations in polymer chemistry
after the World War II launched a new, rapidly changing
era of materials technology. Composite materials, featuring glass, carbon fiber or Kevlar (http://www2.dupont.com/)
embedded in polymer matrices, are widespread throughout modern societies. However, declining oil resources and
global requirements for sustainable development impose a
need to replace petrochemical-based materials with renewable ones. The synthesis of biological materials in nature
occurs by energy efficient processes at moderate temperatures and low pressure, using water as the solvent. The low
energy input of these processes is compensated by the
precise molecular design of the constituent biopolymers,
which drives efficient self-assembly of biomaterials [1].
Owing to their smart design, natural materials outperform
most man-made composites. Classical examples of natural
high-performance composites in which matrix proteins
control the formation of the inorganic component include
abalone shells, bone and enamel [2,3]. Other natural composites with unique properties include cactus spines, with
their high degree of stiffness contributed by an arabinan
cellulose composite [4], and the tunics of sea peaches,
composed of cellulose, proteins and mucopolysaccharides
[5]. Therefore, mimicry of biological structures and the
Corresponding author: Teeri, T.T. (tuula@biotech.kth.se).
Available online 23 May 2007.
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processes controlling their synthesis and self-assembly

has great technical and scientific potential.
Wood is perhaps one of the most complex natural
composite substances and it is widely used as a structural
material. The properties of wood are governed by three
levels of organization [6]: (i) the chemical composition and
nanostructure of the cell walls; (ii) the patterning of the cell
walls and the size and shape of the constituent wood cells;
and (iii) the hierarchical organization of the different cell
types to form higher orders of the wood structure. The
excellent mechanical properties of wood rely on the composite nature of the fiber cell walls, in which the cellulose
microfibrils reinforce a matrix of hemicelluloses and lignin.
Owing to their low density and excellent strength properties, fibers extracted from hemp or linen have been used to
develop lightweight biocomposites, particularly for use in
automobile interiors. However, owing to the different chemical properties of natural fibers and synthetic matrices, poor
interfacial stability of such composites is a common problem
[7,8]. Another problem is the high moisture absorption of
plant fibers, which leads to poor dimensional stability.
Because living plants have reconciled these problems,
the compatibility between phases can be improved by
learning from nature, to produce materials with increased
functionality.
The plant cell wall a versatile biocomposite
The primary cell wall is a dynamic structure that undergoes
many changes during plant development, and it must be
able to accommodate a variety of mechanical requirements
[9]. The basic building blocks of primary cell walls are
cellulose, hemicelluloses, pectin and structural proteins
(Box 1). The strong crystalline cellulose microfibrils encapsulate the cell in a mesh-like structure (Figure 1a), and
hemicelluloses associate with the cellulose microfibrils to
combine strength with extensibility. For example, the xyloglucans that associate intimately with cellulose microfibrils
[10] also act as tethers between adjacent microfibrils,
thereby contributing to the porosity and extensibility of
the primary cell walls of dicotyledons. Pectins [11] are joined
by covalent and ionic cross-links to form structural networks
that are independent but synergistic with the cellulose
hemicellulose networks [12]. The physical properties of
hemicelluloses and pectin are largely determined by the
nature and abundance of side chains, and depend on the
local concentrations of ionically and covalently bound salts

0167-7799/$ see front matter 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.tibtech.2007.05.002

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Box 1. Main components of plant cell walls

 Cellulose is a linear polymer of b-(1,4)-D-glucose units. During
cellulose biosynthesis the individual polymers stack onto each
other forming crystalline microfibrils that act as the main loadbearing component of the cell wall (Figure Ia).
 Hemicelluloses are a heterogeneous group of branched polysaccharides, including xyloglucan in the primary cell walls of
dicotyledons, glucuronoxylans in the secondary cell walls of
hardwoods, as well as glucomannan and arabinoxylan in the
secondary cell walls of softwoods. Arabinoxylan is also commonly
found in the cell walls of grasses (Figure Ib).
 Pectin constitutes another group of branched polysaccharides,
including homogalacturonans, rhamnogalacturonans, and substi-

tuted galacturonans [10]. Pectin is typically found in the middle

lamella, cementing together the primary walls of adjacent plant
cells (Figure Ic).
 Lignin is a complex phenolic polymer composed of three monomeric building blocks: coniferyl alcohol, sinapyl alcohol and
paracoumaryl alcohol. Lignin fills the spaces in the cell wall
between cellulose, hemicellulose and pectin, thereby conferring
mechanical strength to the cell walls. Lignin is typically present in
the secondary cell walls of trees (wood) (Figure Id).
 Structural proteins commonly found in plant cell walls include, for
example, extensins, arabinogalactan proteins (AGP) as well as
glycine-rich (GRPs), and proline-rich (PRPs) proteins [16] (Figure Ie).

Figure I. Main components of plant cell walls. (a) The repeating unit of a cellulose polymer; (b) the repeating unit of a xyloglucan polymer, one of the cell wall
hemicelluloses; (c) part of the structure of polygalacturonan, a cell wall pectic polysaccharide; (d) the building blocks of lignin; (e) the amino acid sequence of a typical
proline-rich protein (PRP) found in plant cell walls.

Figure 1. Organization of the cellulose structure in the primary (a) and secondary (S2) (b) cell wall layers of spruce wood tracheids, as observed using scanning electron
microscopy. The individual cellulose microfibrils are organized into larger aggregates that have multidirectional orientation in the primary wall but show distinct alignment
in the secondary wall layers. Bars = 1 mm.
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(e.g. calcium and boron) and potential cross-linking agents,

such as ferulic and coumaric acids [11,13]. By varying the
structures or the chemical environment of these polymers,
the mechanical properties of cell walls can be altered during
developmental stages and in different plant tissues [6]. The
diversity of mechanical properties observed in different
parts of thistle flower is an excellent example of such variation [14].
A variety of cell wall-associated enzymes and other
proteins also influence cell wall properties and behavior
[15]. For example, the extensibility of the cellulosexyloglucan network can be regulated by the action of expansins,

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specific cell-wall proteins [16]. In addition, many different

structural proteins (Box 1) are cross-linked to the cell wall
and might influence its mechanical behavior [17].
The basic structural elements of wood the tracheary
elements in softwood and fibers in hardwood are hollow
dead cells surrounded by secondary cell walls (Figure 2). The
secondary walls contain different hemicelluloses to the
primary walls and are lignified (Box 1) in the latter stages
of wood formation. The cellulose microfibrils are deposited
in the secondary walls by a highly organized process, leading
to a hierarchical structure with three layers (S1, S2 and S3).
In the dominant, S2, layer the cellulose microfibrils are well

Figure 2. Varied structures of wood cell walls. (a, c and e) birch fibers (F) and vessels (V) showing characteristic scalariform perforation plates (SP) that connect axial vessels
and intervessel pitting (IP). (b) Spruce latewood tracheids (T). (d) Douglas fir with typical spiral thickening (SP) and bordered pits (BP) in earlywood tracheids. (f) Pine
bordered pit (BP) with margo (M); (g) Devil tree (Alstonia scholaris) vessel with vestured intervessel pits (VIP). (h) Tension wood fibers from poplar wood with thick cellulose
rich (CR) inner S2 wall layer. Bars: (a) = 100 mm; (b, d and f) = 50 mm; (c, e and h) = 20 mm; (g) = 10 mm.
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aligned (Figure 1b), contributing to the general mechanical

strength of wood [18]. Changes in the chemical composition
or the fine structure of wood cells by developmentally controlled or by externally induced stimuli lead to different
mechanical properties. For example, the secondary walls of
many wood cells display a wide variation in pitting
(Figure 2). The length and width of the cells also vary
between softwoods and hardwoods, and between different
tree species. Furthermore, deviations from the vertical position of the stem are compensated by the formation of
tension wood with high cellulose content in hardwoods
(Figure 2h), and compression wood with increased lignin
content in softwoods. The gelatinous, non-lignified cell walls
of tension wood and flax fibers feature thick S2 layers
(Figure 2h) and a high proportion of galactose-containing
polymers [19]. Understanding the structureproperty
relationships in different types of plant tissues paves the
way for the rational development of novel materials.
Genetic approaches for studying and engineering
plant cell walls
The isolation and characterization of the components of
different plant organs and tissues are not trivial tasks. For
structural analysis, the individual polymers must be
chemically or physically separated from other cell wall
components. Because the different cell wall polymers are
intimately entangled and cross-linked by many noncovalent and covalent interactions they are difficult to
separate without changing their chemical and/or physical
structures. It has been difficult to correlate cell wall mechanical properties with the structural characteristics of the
constituent polymers using traditional techniques.
Screening and characterizing cell wall mutants of thale
cress (Arabidopsis thaliana), combined with mechanical,
spectroscopic and enzymatic characterization of the affected
tissues, have recently provided powerful tools towards understanding cell wall properties [2024]. Furthermore,
recent advances in genomic and proteomic approaches provide rapid and convenient tools to identify previously
unknown enzymes that synthesize plant cell walls [25]. A
recent and elegant example describes how xyloglucan side
chains influence the overall wall strength of Arabidopsis
hypocotyls [26]. By measuring the tensile parameters of
hypocotyls of a series of dark-grown Arabidopsis mutants,
the loss of galactose-containing side chains of xyloglucan
was shown to impair the overall wall strength, whereas loss
of xyloglucan fucosylation only caused a slight decrease of
tensile strength. By contrast, loss of fucose in rhamnogalacuronan II (RGII) reduced the degree of boron-mediated
cross-linking of pectin and resulted in clearly reduced tensile strength [26].
Microbes provide rich sources of enzymes that degrade
plant biomass [24], and several recent examples demonstrate their potential as tools to alter plant cell wall properties. Expression of microbial cellulases or non-catalytic
cellulose-binding modules [27] increased the rate of cellulose
biosynthesis and altered the morphology of the cellulose
produced by transgenic plants [28]. Transgenic tobacco
(Nicotiana tabacum) and rice (Oryza sativa), both expressing microbial cellulases, exhibited swollen cell walls, which
improved the digestibility of this structure [29,30]. In Italian
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ryegrass (Lolium multiflorum) the arabinosyl residues in

arabinoxylans are cross-linked by ferulic acid esters, and the
expression of a fungal ferulic acid esterase gene in transgenic ryegrass made the cell wall more susceptible to enzymatic attack by endoxylanases [31]. These examples aimed
to improve the digestibility of plant biomass by ruminants,
but the expression of enzymes acting to increase crosslinking of the cell wall components could also be used to
improve cell wall mechanical properties.
Bacterial cellulose: towards novel biocompatible
materials
Bacteria of the Gluconacetobacter genus extrude pure
cellulose ribbons into their culture medium [32]. The properties of the resulting cellulosic material can be modulated by
using different bacterial strains, by varying the parameters
of the cultivation, and by adding chemicals and polymers
into the culture medium [33,34]. Access to such nonassembled building blocks opens opportunities towards a
new generation of materials. Cellulose is fully biocompatible, and interesting potential applications have already
emerged in the health-care sector, including wound healing
systems and artificial-skin products [32]. Bacterial cellulose
can also be grown in defined shapes, such as hollow tubes,
that can be used to replace blood vessels in surgical operations [34]. Tubular implants made of bacterial cellulose
with high water content are flexible and elastic but maintain
a constant shape. They are stable against pressure and can
be easily sutured into biological tissues [34,35]. Native
bacterial cellulose has recently also been presented as a
potential, biocompatible scaffold with good mechanical
properties for the engineering of cartilage and blood vessels
[36,37].
Understanding the composite nature of plant
cell walls
Growth of the bacterium Gluconacetobacter xylinus in the
presence of various plant cell wall polysaccharides provides
a convenient model system to investigate the properties of
natural cell walls. The addition of hemicelluloses to the
bacterial fermentation of cellulose affected the size of the
microfibrils [38,39], which supports the hypothesis that
hemicelluloses influence the aggregation pattern of cellulose [40]. The addition of xyloglucan reduced the crystallinity of cellulose, possibly by intercalating into the cellulose
crystals [41]. The galactose side chains of xyloglucan constitute the major determinant of cellulose composite formation, whereas the fucose substitution seems to act as a
secondary modulator [42]. Xyloglucan, glucomannans and
some galactomannans that cross-link cellulose microfibrils
in primary cell walls contributed considerably to the elastic
properties of bacterial cellulose-based composites [43,44].
By contrast, the formation of cellulose and/or pectin networks apparently does not occur by direct binding interactions between cellulose and pectin, but instead depends on
the presence of preformed pectin networks of moderate
strength [45]. Tensile deformation tests carried out with
different combinations of cellulose, xyloglucan and pectin
revealed that the best strength was achieved by cellulose
only, whereas composites including xyloglucan or pectin
were weaker but more extensible [46]. The properties of

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Figure 3. Atomic force microscopy (AFM) images (22 mm) of (a) a film of pure bacterial cellulose; (b) a composite film containing 75% bacterial cellulose and 25% xylan,
and (c) a composite film containing 50% bacterial cellulose and 50% xylan [48].

the composites could be further modulated by adding

hydrolytic enzymes or expansin [47,48].
The structure of secondary cell walls has been mimicked
by mixing bacterial cellulose with glucuronoxylan extracted
by alkali from aspen wood [49]. The mechanical properties,
such as tensile strength and stiffness, of the nanocomposites
reached a maximum at the same cellulose:xylan ratio as in
the secondary cell wall, and their morphology resembled
that of native secondary cell walls (Figure 1, 3ac). Even
closer similarity with secondary cell walls was achieved
when the composites were prepared by using glucuronoxylan extracted from delignified aspen wood chips using
DMSO to preserve the native acetyl groups. Moistureinduced softening due to the plasticization of glucuronoxylan was observed with both the wood fibers and in the
model system. However, the softening behavior of the two
materials was not identical, most probably owing to differences in spatial organization of the components [50]. In
addition to excellent mechanical properties, the cellulose
glucuronoxylan composites had a transparent appearance
similar to plastic films or composites of cellulose nanocrystals mixed with polymer resins [51,52]. Furthermore, the
celluloseglucuronoxylan composites also exhibited remarkable oxygen barrier properties, which make them excellent candidates for future advanced packaging materials
[53].
Biomimetic modification of cellulose surfaces
Most cellulose-based materials used industrially are
chemically treated at various points in the manufacturing
process to alter their properties. When chemically modified
by acetylation, cellulose loses its native structure and
strength [34]. In applications relying on the superb loadbearing properties of cellulose, coating rather than direct
chemical modification is therefore used to alter the surface
and the bulk mechanical properties, but not the structure,
of cellulose. For example, paperboard used as packaging
material for liquids and foodstuffs is often laminated with a
thermoplastic and aluminium to provide an impermeable
barrier to aqueous solutions.
In biocomposites, cellulose is blended in a polymer matrix
in the presence of various compatibilizers, to achieve good
strength properties [7]. With current technology, however,
problems are often encountered in the retention of chemicals
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on the fiber surfaces and the interfacial stability of the

material blends [8]. In plants, the combination of strength
and interfacial stability is achieved by coating cellulose with
matrix polysaccharides that can interact with other cell wall
polymers and proteins. One idea borrowed from nature is to
use chemically or enzymatically modified xyloglucan to
attach functional groups to cellulose surfaces [54]. Xyloglucan can be isolated in bulk amounts from tamarind seeds
and it binds to cellulose surfaces with high affinity. It can be
easily modified by a step-wise process (Figure 4), starting
with chemical modification of small xyloglucan oligosaccharides, which are not retained on cellulose surfaces but
which can be joined to polymeric xyloglucan by using
the enzyme xyloglucan endo-transglycosylase (XET, E.C.
2.4.1.207). Adsorption of the modified high molecular weight
xyloglucan onto cellulosic materials results in surfacespecific chemical functionalization. Using this approach, a
wide range of different chemical groups (Figure 4) have been
attached onto different types of cellulose surfaces, including
pulp fibers, regenerated cellulose films and filter paper
[54,55]. Of particular interest is the attachment of initiators
of polymerization reactions that permit the construction of
extended polymer architectures onto cellulose surfaces
[56,57]. In this way, the cellulose fibers can be completely
encapsulated in a polymer sheath, which should considerably improve their compatibility with the polymer matrices
commonly used to prepare biocomposites.
In addition to xyloglucan, the affinity of xylan to the
surfaces of cotton linters and bacterial cellulose has been
used to modify their surface properties [58], and modification of pulp fibers by controlled assembly of xylans has
been shown to improve the wettability of the pulp [59].
Pullulan abietate, a compound mimicking the natural
hemicellulose- and lignin-containing interfaces in plant
cell walls, has also been found to rapidly self-assemble
in a desorption-resistant fashion onto cellulose surfaces
[60]. Following enzymatic cross-linking or thermoplastic
consolidation, it might also prove useful for improving the
interfacial stability of cellulose-based composites.
Besides physical binding interactions and entanglement,
plant cell walls are consolidated by covalent or non-covalent
cross-linking of the matrix polysaccharides with each other
or with lignin. Enzymes have crucial roles in creating
these interactions and thus provide interesting tools for

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Figure 4. An overview of xyloglucan-mediated cellulose surface modification [50]. (a) Low molecular weight xylogluco-oligosaccharides (XGO) bearing chemical
functionality (R) are incorporated into high molecular weight xyloglucan (XG) by the enzymatic action of a xyloglucan endo-transglycosylase (XET). (b) The resulting
xyloglucan conjugate (XG-R) is adsorbed from aqueous solution at ambient temperature onto a cellulose surface, for example, cotton fiber, wood pulp fiber or regenerated
cellulose. (c) The installed functional group might produce a desired surface effect itself or might be transformed through further reactions on the cellulose surface.
Examples include fluorescent optical brightening agents (i); nucleophiles such as amino groups (ii); thiol groups for specific, reversible modification (iii); biomolecule
capture agents, including ligands and reactive groups (iv); and initiators for radical polymerization, for example, atom transfer radical polymerization (ATRP), to produce
highly hydrophobic surfaces (v). Xylogluco-oligosaccharides (XGO) are based on a b(14) glucan backbone and have the general composition Glc4Xyl3Gal02. Native
tamarind xyloglucan (XG) comprises multimers of these oligosaccharides and has a molecular weight >200 000 Da. A xyloglucan chain of >5 XGO units is required for
quantitative binding to cellulose. Abbreviations: DTT, dithiothreitol (a disulfide bond reducing agent); FITC, fluorescein isothiocyanate; MTS, methanethiolsulfonate.

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fiber modification. The wood particles and fibers used to

manufacture boards are traditionally treated with synthetic
formaldehyde-based resins or subjected to high pressures
and temperatures to improve strength properties. In wood,
oxidative enzymes catalyze the formation of monolignol
radicals, which undergo chemical coupling to form lignin
polymers. Oxidative enzymes have been used to activate
surface exposed lignin and lignin precursors on wood fiber
surfaces, leading to enhanced bonding strength of medium
density fiber boards [61,62]. Pilot-scale testing of fiberboards made of enzyme-activated wood fibers demonstrated
strength properties comparable to boards bonded by synthetic resins [63,64].
Cross-linking proteins for interfacial stability
Natural or engineered cross-linking proteins might also
become useful tools to enhance binding interactions between
natural polymers, thereby leading to improved strength
properties of the resulting composites [65]. Extensins
(Box 1) represent natural cross-linking proteins that probably form covalent networks in plant cell walls by their
tyrosyl and lysyl residues cross-linked by extensin peroxidases [66]. Because complex natural cross-linking proteins
are difficult to produce in large quantities, biomimetic
approaches might provide a practical alternative. For
example, the addition of engineered double-headed cellulose-binding proteins improved the mechanical properties of
papers and increased the water-repellence of paper surfaces
[67]. Furthermore, bi-functional proteins consisting of cellulose- and starch-binding modules promoted interactions
between cellulose and granular starch and improved the
binding of soluble starch onto cellulose surfaces [65]. By
building on the natural diversity of protein-binding interactions as well as protein engineering, a wide range of
molecular architectures can be designed. Small protein
domains can also be engineered to bind metals, semi-conducting oxides and other inorganic compounds to fabricate
different types of functional nanostructures [68]. When
linked to cellulose-binding modules, such molecular assemblies can be anchored to cellulose surfaces to give rise to a
whole range of new types of functions. Potential applications
of such surfaces extend far beyond the traditional uses of
cellulose.
Conclusions and future perspectives
The interest in entirely bio-based materials is steadily
increasing in response to growing environmental awareness
and declining supplies of raw oil. Cellulose microfibrils are
natural reinforcing elements with excellent mechanical
properties but, nevertheless, poor performance in technical
applications. This is mainly due to insufficient dispersion of
the microfibrils into commonly used polymeric matrices in
traditional processing equipment, differences in polarity
between cellulose and hydrophobic polymers that result
in poor interfacial adhesion, and poor dimensional stability
of cellulose fibers when subjected to moisture. As summarized in this review, it might be possible to solve some of these
bottlenecks by learning how cellulose interacts with other
biopolymers in plant tissues. In particular, the lack of
interfacial interactions with other polymers can be overcome by coating cellulose with (bio)polymers that bind to
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cellulose surfaces and can, in turn, be chemically activated.

This opens the whole range of composite manufacturing
technologies relying on in situ polymerizing matrices.
Natural cellulose fibers that exhibit a range of different
sizes and shapes are not suitable building blocks for nanostructured composites. However, recent technology to prepare cellulose nanocrystals of defined dimensions is
overcoming this limitation [69], thus paving the way for
the development of cellulose-based nanomaterials for different types of coating applications and packaging materials.
In addition, cellulose-based materials with an excellent
capacity to hold water have interesting applications as
biocompatible materials that can be used in contact with
the human body, including tissue scaffolds and implants,
wound dressings, biocompatible coatings and drug-release
formulations. The biomimetic approaches described in this
review have great potential to tailor the functionality and to
improve the performance of these new materials.
Acknowledgements
We thank the Knut and Alice Wallenberg Foundation (http://
www.wallenberg.org) (G.D., P.G. and T.T.) and the Swedish Research
Council (http://www.vr.se) (H.B.) for financial support.

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