Cellulose (2024) 31:10165–10190

https://doi.org/10.1007/s10570-024-06220-0

REVIEW PAPER

Applications of regenerated bacterial cellulose: a review

Lucas Rosson · Boon Tan · Wayne Best ·
Nolene Byrne

Received: 11 July 2024 / Accepted: 8 October 2024 / Published online: 26 October 2024
© The Author(s) 2024

Abstract Whilst synthetic polymers have changed found in a variety of commercial products. However,
the world in many important ways, the negative dissolving and regenerating bacterial cellulose is a
impacts associated with these materials are becom- potential avenue to broaden the applications available
ing apparent in waste accumulation and microplas- to this material. The aim of this study is to review the
tic pollution due to lack of biodegradability. Society applications which utilize regenerated bacterial cel-
has become aware of the need to replace or substitute lulose, with a focus on the dissolution/regeneration
environmentally persistent synthetic polymers, and methods used and discussing the associated limita-
cellulose has received a large amount of attention in tions and future outlook.
this respect. The mechanical properties of cellulose,
its renewable nature and biodegradability are advan- Keywords Bacterial cellulose · Cellulose
tageous properties. Drawbacks exist for the use of dissolution · Cellulose regeneration · Sustainability ·
plant cellulose (PC), including the water footprint of Polymer processing
cotton, deforestation associated with wood/dissolving
pulp, and the extensive processing required to refine
plants and wood into pure cellulose. Bacterial cellu- Introduction
lose (BC), also known as microbial cellulose, is gain-
ing momentum in both academic and industry set- Synthetic polymers play a key role in society today,
tings as a potential solution to the many drawbacks of from packaging and building materials to adhesives
plant-based cellulose. Compared to PC, BC has high and paints. However, the environmental impact of
purity, crystallinity and degree of polymerisation, and these polymers has become a mainstream issue with
can be manufactured from waste in a way that yields concern around the fossil-based raw materials as
more cellulose per hectare, per annum, and requires well as pollution due to microplastics and persis-
less intense chemical processing. Native bacterial cel- tence in the environment (Tran et al. 2023). It is for
lulose can be formed and shaped to an extent and is these reasons that academia, industry, and consum-
ers are looking for alternatives. Biopolymers offer
potential solutions via their biodegradability and
L. Rosson (*) · N. Byrne
Institute for Frontier Materials, Deakin University, renewable feedstock (Gowthaman et al. 2021). The
Waurn Ponds, VIC, Australia Earth’s most abundant biopolymer, cellulose, has
e-mail: l.rosson@deakin.edu.au received attention to replace synthetic polymers in
several applications. Cellulose is desirable due to its
B. Tan · W. Best
Nanollose Ltd, Nedlands, Australia good mechanical properties, high thermal stability,

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biocompatibility, and a chemical structure amenable of the organism. The shape and number of substruc-
to functionalisation. Cellulose is also widely available ture spinnerets within a terminal complex depends
from plants, though extensive processing is usually on the organism (Cousins and Brown 1995). Plant
required to liberate cellulose from other biopolymers cellulose is biosynthesised by ‘rosette’ complexes
present in the plant matrix (Suresh 2018). Bacterial within the plant cell via sequential deposition of glu-
cellulose is naturally extruded by the cell in the form cose units into a molecular chain and simultaneous
of pure nano-scale ribbons and so does not require assembly of these chains into elementary fibrils, the
extensive post-processing to obtain pure cellulose basic unit of supramolecular structure (Garvey et al.
(Gao et al. 2011). The production of BC by the Ace- 2005; Zanchetta et al. 2021). The terminal complexes
tobacter xylinum bacteria was first reported by Brown responsible for assembly of cellulose in vascular
in the late nineteenth century as a “jelly-like translu- plants are organised in sixfold symmetric rosettes
cent mass on the surface of the culture fluid” up to whereas the terminal complexes of bacterial cells are
25 mm in thickness (Brown 1886). Later, Hestrin and arranged in linear rows with single or multiple layers
Schramm reported extracellular cellulose synthesis (Suresh 2018). In bacteria of the Acetobacter genus
in the presence of glucose and oxygen (Hestrin and it is thought that each bacterial cell has only one lin-
Schramm 1954). Figure 1 demonstrates bacterial cel- ear terminal complex consisting of a single row of
lulose fibril extrusion, a network of cellulose nanofi- substructure catalytic sites. Whilst each substructure
brils, and the jelly-like pellicle produced. in plant cellulose terminal complexes are believed
Bacterial cellulose and plant cellulose have an to extrude six cellulose molecular chains (Perez and
identical chemical structure consisting of β-D-glucose Mazeau 2005), the substructures in bacterial cellulose
units connected via (1–4) glycosidic bonds with the are able to extrude 10–15 cellulose chains (Cousins
chemical formula ­ (C6H10O5)n (Gupta et al. 2019) and Brown 1995). Figure 2 shows a conceptualisa-
where each glucose unit is arranged to be inverted to tion of cellulose extrusion and assembly from the
the adjacent glucose unit (Shanks 2014). Cellobiose, terminal complexes of plant and bacterial cells. For
the monomer unit of cellulose, contains two glucose both plant and bacterial cellulose, the organisation of
units and is equivalent in cellulose from plant, algal elementary fibrils into a semicrystalline structure is
and bacterial sources. The differences between bac- thought to be controlled by cellulose synthase com-
terial and plant cellulose arise in the supramolecular plexes located in the cells of the organisms (Rosén
structure and stem from biosynthesis routes, purity, et al. 2020; Suresh 2018; Zanchetta et al. 2021).
crystalline structure and crystallinity, morphology, Plant cellulose biosynthesis involves the elemen-
and degree of polymerisation. tary fibrils and microfibrils being intimately mixed
Biosynthesis of native cellulose occurs via the with other biopolymers such as lignin and hemicel-
extrusion of assembled cellulose chains from spin- luloses (Ahmed et al. 2020). Woody biomass, in
nerets within terminal complexes, located in the cell general, contains 40–50 wt% cellulose, whilst cotton

Fig. 1  Transmission electron microscope (TEM) image of lulose nanofibrils reproduced from (Ding et al. 2021) with
bacterial cell extruding cellulose fibril reproduced from (Tokoh permission from Elsevier (middle), and a photograph of a bac-
et al. 1998) with permission from Springer Nature (left), scan- terial cellulose pellicle provided by Nanollose (right)
ning electron microscope (SEM) image of 3D network of cel-

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secondary cell wall contains three layers, each layer

consisting of parallel microfibrils orientated at some
angle relative to the fibre axis (Heinze 2016). The
morphology of bacterial cellulose is often described
as a reticulated 3D network of cellulose ribbons or
fibrils (Charreau et al. 2020; Zhong 2020; Zuppolini
et al. 2022). Like plant cellulose, the fibres or ribbons
of bacterial cellulose also consist of agglomerations
of fibril structures. The elementary fibrils with diam-
eters of 2–4 nm crystallize into microfibrils, which
form ‘bundles’, and these bundles agglomerate into
the ribbons that make up the 3D network of bacterial
cellulose (Charreau et al. 2020). Cellulose is a semic-
rystalline material containing areas of high structural
order and areas of relatively low order, with regions
of intermediate order (para-crystalline regions)
between or surrounding ordered regions (Ioelovich
2016; Park et al. 2010). The smallest structural unit
Fig. 2  Conceptual schematic of cellulose biosynthesis struc-
tures for plant (top) and bacterial (bottom) cellulose. Created of cellulose, the unit cell, differs between the two
with Chemix (https://​chemix.​org) allomorphs found in native cellulose, cellulose Iα and
Iβ. The dominant crystalline allomorph in a cellulose
sample depends on the source. In plant cellulose, Iβ is
can contain above 90 wt% cellulose (Heinze 2016). the major crystalline form (Ahmed et al. 2020; Zhong
Bacterial cellulose is free from lignin and hemicellu- 2020), whilst cellulose produced by more primitive
loses, leading to a far higher purity than plant-based organisms, such as bacteria, is comprised mainly of
cellulose (Heinze 2016). A raw bacterial cellulose the Iα crystal form (Heinze 2016). The proportion of
pellicle contains bacterial cells, residual nutrients cellulose Iβ in crystalline plant cellulose ranges from
from the culture medium, and other by-products from 60–80% (Ahmed et al. 2020; Somerville 2006) whilst
cellulose synthesis (Zhong 2020). Once the residual the proportion of cellulose Iα in crystalline bacterial
components have been removed with a weak NaOH cellulose has a range of 60–82% (Ahmed et al. 2020;
treatment, the cellulose content of dry bacterial cellu- Singhsa et al. 2018; Zhong 2020). BC is known to
lose pellicle has been reported as 97% (Okiyama et al. have higher crystallinity index, or proportion of crys-
1992). talline material, than many plant celluloses, close to
As biosynthesis of bacterial cellulose differs from that of cotton cellulose (Agarwal et al. 2021). A sum-
that of plant cellulose, it follows that differences mary of structural and morphological differences
in morphology arise. Plant and bacterial celluloses between BC and PC can be found in Table 1.
have notable differences in the shape and size of the Bacterial cellulose is often said to have a higher
fibres. Wood pulp fibres have widths from 10 to 50 degree of polymerisation (DP) than plant cellulose
µm (Chinga-Carrasco 2011) and cotton fibres have (Amr and Ibrahim 2022; Zhong 2020), however
diameters from 8 to 20 µm (Wang and Wang 2009). the opposite is often the case for these celluloses in
The ribbon-shaped fibrils of bacterial cellulose are the native form. For instance, the primary cell wall
nano-scaled having widths in the range of 80–150 nm in plants contains cellulose with a DP of approxi-
(Heinze 2016), and varying in thickness from 3–8 nm mately 8,000 and the secondary cell wall can have a
(Felgueiras et al. 2021; Khajavi et al. 2011). Whilst DP up to 15,000 (Brown Jr 2004), whilst the average
fibres from different plants can have different mor- DP of wood is around 10,000 and the DP of cotton
phologies, for example the twisted-ribbon shape of is 15,000 (Acharya et al. 2021). Values for the DP
cotton fibres and the straight fibres of spruce wood, of bacterial cellulose in literature are lower, ranging
all have a multi-layered cell wall. The primary cell from 2,000–6,000 (Khajavi et al. 2011) and > 8,000
wall consists of amorphous cellulose ‘nets’ whilst the (Hamimed et al. 2020), up to 10,000 (Heinze 2016).

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Table 1  Differences between plant cellulose (PC) and bacterial cellulose (BC)
Property Plant cellulose (PC) Bacterial cellulose (BC) References

Biosynthesis—structure of terminal Sixfold symmetric ‘rosette’ Linear row Suresh (2018)

complex
Biosynthesis—number of molecu- 6 10–15 Cousins and Brown (1995), Perez and
lar chains extruded per terminal Mazeau (2005)
complex substructure
Purity (% cellulose) Wood: 40–50% 97% Heinze (2016), Okiyama et al. (1992)
Cotton: 90%
Fibre width Wood: 10–50 µm 80–150 nm Chinga-Carrasco (2011), Heinze
Cotton: 8– 20 µm (2016), Wang and Wang (2009)
Crystallinity index Wood: 52–65% 74–89% Agarwal et al. (2011), Liu et al.
Cotton: 85–96% (2019), Nam et al. (2016), Tsouko
et al. (2015)
Dominant crystalline structure Cellulose Iβ Cellulose Iα Ahmed et al. (2020), Heinze (2016),
Zhong (2020)
Degree of polymerisation Wood: 10,000 2,000–10,000 Brown Jr (2004), Cotton Incorporated
Cotton: 15,000 (2023), Heinze (2016), Khajavi et al.
Dissolving pulp: 500–1,500 (2011), Makarov et al. (2019)

The notion of BC having a higher DP than PC may waste or byproducts (Felgueiras et al. 2021; Mehro-
refer to the celluloses after processing. The treat- tra et al. 2023). Using waste as the fermentation
ments used for wood cellulose severely degrade the media may also help to combat a major drawback of
DP, with dissolving pulps having a DP between 500 BC production; the high cost of fermentation media
and 1,500 (Cotton Incorporated 2023; Makarov et al. (Felgueiras et al. 2021). Bacterial cellulose is com-
2019). Despite the less intense treatments required monly produced in the form of nata de coco via the
for cotton fibres, processing still causes large reduc- fermentation of coconut water (Charoenrak et al.
tions in the DP of cotton, reducing from around 2023; Nugroho and Aji 2015). Recently, a variety of
9,000–15,000 (Cotton Incorporated 2023) in raw cot- waste streams and by-products have been investigated
ton fibres to 2,000–3,000 in scoured and bleached as nutrient sources for BC production including milk,
cotton fibres (Palme et al. 2016). The low concentra- waste beer yeast, beet molasses, citrus juice, rotten
tion NaOH wash required to remove microbes and fruit, and wastes from the confectionary and bio-
impurities from BC has a smaller effect on lowering diesel industries (Akintunde et al. 2022; Tsouko et al.
the DP, allowing the processed material to maintain a 2015; Zhong 2020). The land use and yield benefits
DP value closer to the native state. of BC compared to wood cellulose were explained
Bacterial cellulose not only has physical advan- by Jozala et al. (2016), with the production of 80 t of
tages over plant celluloses due to high purity, high plant cellulose requiring 7 years on 1 hectare of land
degree of polymerisation and crystallinity, but also and the same mass of bacterial cellulose requiring 22
has potential advantages in terms of sustainabil- days in 500,000 L of fermentation medium. BC may
ity. The environmental advantages of BC compared also have environmental advantages over PC in terms
to PC mentioned in the literature are less pollution of water footprint, with one life cycle assessment
due to the reduced processing requirements for cel- (LCA) study suggesting 98% of the water used can
lulose purification (Bungay et al. 1997; Chen et al. be returned to fresh water after treatment (Forte et al.
2010; Gao et al. 2011; Kalyoncu and Peşman 2020; 2021). However, further comparative LCAs between
Mehrotra et al. 2023; Mona et al. 2019; Potočnik PC and BC are needed to fully elucidate differences
et al. 2023; Zhang and Luo 2011), reduction in land in environmental impact.
area and deforestation required for cultivation (Fel- Bacterial cellulose lends itself to a variety of
gueiras et al. 2021; Mehrotra et al. 2023; Mona et al. applications including most commonly as an addi-
2019), and the ability of BC to be produced from tive for desserts in the form of nata de coco (Iguchi

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et al. 2000), and recently has been applied in com- currently no publications comprehensively review-
mercial applications such as thickener/suspending ing applications of regenerated bacterial cellulose.
agent (Zhong 2020), medicine and cosmetics (Nature The following sections will review the applications
research custom media 2023), wound dressings (Choi of regenerated bacterial cellulose in the biomedical,
et al. 2022; Zhong 2020), textile fibres (Zhong 2020), fibres and textiles, composites, electronics and sen-
vegan leather (Polybion™ 2024), and acoustic dia- sors, 3D printing, filtration and separation, and pack-
phragms (Lee et al. 2014). Properties such as high aging fields, with a focus on solvents used and disso-
purity (Lee et al. 2014), high mechanical strength and lution conditions.
stiffness of nanofibrils (Choi et al. 2022; Hsieh et al.
2008; Yano et al. 2005), high water holding capac- Fibres and textiles
ity (Choi et al. 2022; Lee et al. 2014), and transpar-
ency (Provin et al. 2021) coupled with its sustainabil- Cotton fibres contribute a significant proportion of
ity and biocompatibility make bacterial cellulose a the current fibre production share, comprising 22%
desirable material for many applications (Choi et al. of global fibre production in 2022 (Textile Exchange
2022; Lin et al. 2013). Recent reviews have covered 2023). The demands on scarce resources such as
the different applications of bacterial cellulose, with arable land and water are well known for cotton, and
all of these focusing on the use of BC in the native this is predicted to increase demand for Man-made
form. Previous reviews include biosynthesis and bio- Cellulosic Fibres (MMCFs) as cotton-like alterna-
chemistry of BC (Mishra et al. 2022), production of tives (Haemmerle 2011; Kallio 2021). The majority
BC from waste (Urbina et al. 2021), industrial-scale of MMCFs, or regenerated cellulose fibres (RCFs),
production (Avcioglu 2022; Zhong 2020), biomedical are produced using wood pulp as the source of cel-
applications (Gorgieva and Trček 2019; Horue et al. lulose (Textile Exchange 2023). In 2022, 60–65%
2023; Shrivastav et al. 2022), application in nano- of MMCFs were certified by the Forest Steward-
composites (Revin et al. 2022), BC as a substitute for ship Council (FSC) and/or the Programme for the
plant cellulose (PC) (Sarvananda et al. 2022), as well Endorsement of Forest Certification (PEFC), meaning
as general applications (Choi et al. 2022). Applica- the pulp was sourced from sustainably managed for-
tions specifically involving dissolution and regenera- ests (Textile Exchange 2023). However, the share of
tion of BC are yet to be comprehensively reviewed global forest area certified by these bodies decreased
in the literature. BC dissolution and regeneration can from 2021 to 2022 and is expected to decrease fur-
allow for enhancing the materials morphology to bet- ther, which may put stress on the remainder of the
ter suit certain applications (Rodriguez-Chanfrau world’s forests (Textile Exchange 2023). Further to
et al. 2017), removal of contaminants (Hamid et al. this, the fibre manufacturing processes require puri-
2018), obtaining materials with more structural diver- fied pulp, leading to environmental concern regard-
sity (Horue et al. 2023), and better integration and ing the chemicals and processes used to refine pulp
homogeneity of reinforcement or additives in BC (Soares Silva et al. 2023). One potential application
(Horue et al. 2023; Shah et al. 2013; Ul-Islam et al. for regenerated BC is the substitution of wood pulp in
2019).The smaller number of studies on BC dissolu- these fibre making processes.
tion compared to PC dissolution likely stems from the
lower solubility of BC, making dissolution-regenera- Developing applications
tion more difficult. The limitations in processing and
dissolving BC will therefore also be reviewed. Researchers have been investigating the use of BC
in regenerated cellulose fibres from as early as 2009,
when an article was published on dissolving BC in
Applications of regenerated bacterial cellulose a N,N-dimethyl acetamide/lithium chloride (DMAc/
LiCl) system and regenerating fibres with or with-
Recent reviews have mentioned the importance of out the addition of multi-walled carbon nanotubes
dissolution/regeneration of BC for advanced appli- (MWCNTs) (Chen et al. 2009). Since then, using
cations (Mbituyimana et al. 2021; Raut et al. 2023). BC to substitute wood pulp in these fibres has been
However, to the knowledge of the authors, there are justified by the higher purity and lesser processing

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requirements for BC (Chen et al. 2010; Gao et al. production is demonstrated in Table 2 below, with the
2011; Zhang and Luo 2011), a lower environmen- average cellulose concentration being approximately
tal impact (Soares Silva et al. 2023) and the ease of 6 wt%. This is lower than cellulose concentrations
production of BC (Wu et al. 2017). The commercial seen for wood pulp used in the major cellulosic fibre
usefulness of BC in textile fibres was demonstrated processes; typically 8–9.5 wt% for the viscose process
by the company Nanollose through an infographic in (Kotek 2006) and 10–18 wt% for the lyocell process
a video on their website (Nanollose Ltd 2023). This (White 2001).
shows the potential ‘field-to-yield’ of bacterial cel- Despite having a lower concentration of cellu-
lulose versus other forms of cellulose such as cot- lose in the spinning solution, researchers were able
ton and bamboo—detailing that BC can achieve the to achieve mechanical properties similar to those of
highest yield of cellulose for a given area of land, per commercial lyocell fibres, as seen in Table 2. Soares
annum. Silva et al. (2023) suggested that the higher DP or
Bacterial cellulose is difficult to dissolve at high broader polydispersity of the BC pulp may have
concentrations, even in solvents capable of dissolving enhanced the mechanical properties of fibres in their
a large amount of plant cellulose (Ruka et al. 2015). study. Both Liang et al. (2021) and Wu et al. (2017)
The factors influencing the low solubility of BC will varied the dissolution time to control the amount of
be discussed in a later section, however one method BC dissolved in the spinning dope. By controlling
employed to improve solubility was grinding BC to this parameter, they claim to have produced a com-
a powdered form (Shen et al. 2010). The relationship posite fibre containing a regenerated cellulose matrix
between the form of BC and solubility is supported by supported by undissolved bacterial cellulose rein-
multiple studies where freeze-drying was employed forcement. Interestingly, these studies produced fibres
to dry the BC pellicle (Chen et al. 2009, 2010; Liang with excellent mechanical properties given the low
et al. 2021; Soares Silva et al. 2023; Wu et al. 2017). cellulose concentration.
Freeze-drying allows the BC to maintain much of Other researchers have focused on imparting
its porous morphology and prevents stiffening or functionality to fibres produced from regenerated
hornification (Andree et al. 2021), likely increasing BC. Chen et al. (2009) increased the electrical con-
penetration of solvents into the solid mass. The diffi- ductivity of BC fibres by incorporating multiwalled
culty in dissolving BC at high concentrations for fibre carbon nanotubes (MWCNT) in a BC- DMAc/LiCl

Table 2  Solvent used, cellulose concentration, and mechanical properties for regenerated bacterial cellulose fibres prepared by wet
or air-gap wet spinning
Cellulose type and spinning Solvent Cellulose con- Tenacity (cN/dtex) Elongation (%) References
method centration (wt%)

Wood pulp-air-gap spinning NMMO 13–14 4.0–4.8 10–16 Soares Silva et al. (2023)
BC – wet spinning NMMO 7 0.5–1.5 3—8 Gao et al. (2011)
BC—wet spinning Aqueous LiOH/ 6 1.83 11 Zhang and Luo (2011)
BC/Alg—wet spinning urea/thiourea 4.8 BC/1.2 Alg 1.57 10.8
BC—air-gap spinning NMMO 9–12.2 4.64–5.64 8.3–12.1 Soares Silva et al. (2023)
BC—wet spinning DMAc/LiCl 2 6.5 7.1 Wu et al. (2017)
BC– air-gap spinning DMAc/LiCl 4 3.9* 5.6* Chen et al. (2009)
BC/MWCNT—air-gap spin- 4 BC/1 MWCNT 3.4* 3.9*
ning
BC—wet spinning NMMO 8 0.82 12.66 Lu et al. (2013)
BC/HPCS—wet spinning 5 BC/3 HPCS 1.14 2.81
BC—wet spinning Aqueous ZnCl2 5 0.66–0.83 1.95–3.66 Lu and Shen (2011)
BC—air-gap spinning DMAc/LiCl 4 4.33 17.1 Liang et al. (2021)
N-Methylmorpholine N-oxide-monohydrate, NMMO; Bacterial cellulose, BC; Alginate, Alg; Multiwalled carbon nanotubes,
MWCNT; hydroxypropyl chitosan, HPCS. *values determined and converted from stress–strain curve

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solution and regenerating the solution into fibres via Fibres Conference 2023 for their Nullarbor™ fibre
air-gap wet spinning. The same group also created an (Fibre2Fashion News Desk 2023). The Nullarbor fibre
MWCNT nonwoven mat by electro-spinning a solu- is a cellulosic fibre produced via the lyocell process,
tion of BC and the ionic liquid (IL) 1-allyl-3-methyl- using N-Methylmorpholine N-oxide (NMMO) as the
imidazolium (Chen et al. 2010). Dimethyl sulfoxide cellulose solvent and bacterial cellulose in place of
(DMSO) was used as a cosolvent for viscosity adjust- wood pulp. Pilot batches to date have produced over
ment and MWCNT were added to the BC-IL-DMSO one tonne of fibres containing as much as 30% bacte-
system. The addition of MWCNT increased the ten- rial cellulose (Fibre2Fashion News Desk 2023). Two
sile strength and Young’s modulus compared to non- important patents published surrounding this technol-
woven mats prepared from BC alone. The authors ogy include one for producing viscose-type cellulose
suggested applications in medical, mechanical, or solution with bacterial cellulose (Jinzarli et al. 2018),
electrical fields. Another group used hydroxypropyl and another for producing lyocell-type fibres with
chitosan (HPCS) to introduce antibacterial proper- varying proportions of bacterial cellulose in place of
ties in regenerated BC fibres using NMMO as the wood pulp (Gupta et al. 2022). Importantly, both of
solvent (Lu et al. 2013). Compared to pure BC fibres, these patents describe technology that can be assimi-
BC-HPCS fibres showed higher tenacity and modu- lated with current cellulosic fibre manufacturing
lus, with significantly lower elongation at break. As equipment (Felgueiras et al. 2021). Not only does the
a functional fibre, the BC-HPCS blend showed excel- bacterial cellulose used in the Nullarbor fibre replace
lent antibacterial activity against the Staphylococcus wood pulp as the cellulosic feedstock, bacterial cel-
aureus bacteria. lulose pellicles are able to be produced from a variety
of agricultural wastes and by-products such as molas-
Commercial applications ses, fruit juices or wheat straw hydrolysates (Keshk
et al. 2006; Tsouko et al. 2015). As seen in Fig. 3 the
Bacterial cellulose textile fibres have recently been Nullarbor fibres by Nanollose have been transformed
developed commercially. The companies Nanollose into yarn, fabrics and garments, showing that they
and Birla Cellulose recently received the ‘Cellulose can be applied to all areas where traditional lyocell or
Fibre Innovation of the Year’ award at the Cellulose rayon fibres are found (Zhong 2020).

Fig. 3  A Nanollose lyocell

garment being modelled

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Other commercial operations have also reported in the biomedical industry, indicating the solvent used
the use of bacterial cellulose in textile fibres. HeiQ as well as any additives. As with dissolution for fibres
AeoniQ, a subsidiary of the chemical company HeiQ, and textile applications, the concentration of BC in
has reported the production of a cellulosic textile solutions for biomedical applications was also low.
fibre made from different cellulosic sources includ- The concentration of BC ranged from 1.5 wt% in the
ing bacterial cellulose (HeiQ AeoniQ 2023). The ionic liquid 1-butyl-3-methylimidazolium chloride
HeiQ AeoniQ website reports the fibres can match (BmimCl) (Shao et al. 2016) to 5 wt% in concentrated
the properties of polyester or nylon fibres, “endlessly trifluoroacetic acid (TFA) (Jayani et al. 2020).
recyclable” and biodegradable in soil. The Hong
Kong Research Institute of Textiles and Apparel Wound healing
(HKRITA) webpage ‘Development of Regenerated
Bacterial Cellulose Fibres from Sustainable Source’ Imparting antibacterial properties to regenerated BC
also reports the use of bacterial cellulose in textile for wound healing applications was a common aim
fibres (The Hong Kong Research Institute of Textiles in literature. This can be achieved by incorporation
and Apparel (HKRITA) 2022). The bacterial cellu- of nanoparticles (NPs) such as copper NPs (Shao
lose, obtained from Kombucha, is said to be dissolved et al. 2016), titanium dioxide NPs (Khan et al. 2015a,
in a novel solvent system free of toxic chemicals, and b) and zinc oxide NPs (Ul-Islam et al. 2014), drugs
spun into textile fibres. such as amoxicillin (Ye et al. 2018), or polymers
such as ε-Polylysine (He et al. 2022) and polyethyl-
Biomedical eneimine (PEI) (Wahid et al. 2020). These materials
were mostly obtained by mixing the active compo-
Due to BC’s moisture properties, non-toxicity and nent (NP, drug, or polymer) with BC in the solution
biocompatibility, applications such as wound healing, state. Exceptions to this included a study by Shao
drug delivery, tissue engineering, bone tissue engi- et al. (2016) where the BC was dissolved and regener-
neering, nerve and dental implants, artificial cornea ated prior to being immersed in a solution containing
and retina, and scaffolds for cell-enzyme immobilisa- ­CuCl2 for copper NP insertion. The membranes, for
tion have been extensively investigated (Blanco et al. potential use as wound dressings, showed the inhi-
2018; Choi et al. 2022; Lin et al. 2013; Zhong 2020). bition zones against five different strains of bacteria
Predominantly, these materials are produced from increased in size with an increase in Cu NP loading.
hydrogels or membranes of native bacterial cellulose The motivation behind the dissolution-regeneration
without dissolution. Many applications of BC in the route in the study by Shao and colleagues was to
biomedical industry require antimicrobial properties, have a higher degree of control over the shape and
and this can be achieved in the form of additives to pore size of the regenerated films. The other excep-
BC membranes or dispersions. Through the dissolu- tion, where active ingredients were included after
tion-regeneration route, the integration and composi- regeneration, was a paper by Ye et al. (2018), who
tion of additives in BC materials can be more easily fabricated amoxicillin-grafted cellulosic sponges for
controlled (Horue et al. 2023; Shah et al. 2013). Bio- wound dressings. In other work by He et al. (2022),
medical applications of regenerated BC also benefit BC was dissolved in ionic liquid (BmimCl) and
from the purity of BC, with less processing required regenerated, before being dissolved again along with
compared to plant-based cellulose. Regenerated BC ε-Polylysine (ε-PL) and hyaluronic acid (HA). The
materials in literature for biomedical applications casting and regeneration of this composite solution
have focused on imparting antibacterial properties resulted in a unique morphology, as the ε-PL and
through additives such as nanoparticles (Khan et al. HA were self-assembled into microspheres dispersed
2015a, b; Shao et al. 2016) or polymers (Wahid et al. through the regenerated BC aerogel. These aerogels
2020), changing the structure for oral drug delivery showed excellent antibacterial properties against
(Badshah et al. 2020; Pandey et al. 2013), and alter- three different bacteria, along with good biocompat-
ing porosity for skin and tissue regeneration (Khan ibility. Wahid et al. (2020), using PEI as the antimi-
et al. 2018; Khan et al. 2015a). Table 3 contains an crobial agent, also added epichlorohydrin (ECH) to
overview of studies on regenerated BC applications aid chemical cross-linking between PEI and BC. The

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Table 3  Overview of applications and additives for regenerated bacterial cellulose in the biomedical field
Application Solvent Additives References

Antimicrobial membrane for 1-butyl-3-methylimidazolium Copper nanoparticles Shao et al. (2016)

wound dressing chloride (BmimCl)
Antimicrobial sponge for wound 1-butyl-3-methylimidazolium Amoxicillin Ye et al. (2018)
dressing chloride (BmimCl)
Antimicrobial film for wound heal- NMMO Titanium dioxide nanoparticles Khan et al. (2015a, b)
ing and tissue regeneration
Controlled release of orally admin- NMMO Model drugs (famotidine or tiza- Badshah et al. (2020)
istered drugs nidine)
Antibacterial materials 1-butyl-3-methylimidazolium ε-polylysine and hyaluronic acid He et al. (2022)
chloride (BmimCl)
Antibacterial materials and bioel- NMMO Zinc oxide nanoparticles Ul-Islam et al. (2014)
ectroanalysis
Tissue regeneration scaffolds NMMO Sodium chloride crystals as Khan et al. (2015a)
porogens
Tissue regeneration scaffolds NMMO Sodium chloride crystals as poro- Khan et al. (2016)
gens and gelatin
Skin regeneration NMMO Gelatin microspheres as porogens Khan et al. (2018)
Incorporation in food for delivery trifluoroacetic acid (TFA) Polyvinyl alcohol (PVA) to allow Jayani et al. (2020)
of probiotics electrospinning
Underwater equipment, implant- LiOH/urea solution Epichlorohydrin (ECH) as chemi- Zhang et al. (2019)
able ionic devices, and tissue cal cross-linking agent
engineering scaffolds
Antibacterial hydrogel NaOH/urea solution Polyethyleneimine (PEI) and Wahid et al. (2020)
epichlorohydrin (ECH)
Bioaffinity and modulation of 1-ethyl-3-methylimidazolium Varied the anti-solvent Johns et al. (2017)
carbohydrate-binding modules acetate (EmimOAc)/dimethyl
(CBMs) sulfoxide (DMSO)
Protein immobilisation for various 1-ethyl-3-methylimidazolium Chitosan Kim et al. (2017b)
biomedical, environmental, and acetate (EmimOAc)
biocatalytic applications
Oral drug delivery NaOH/urea solution Acrylamide (AM) Pandey et al. (2013)

more permanent bonding of antibacterial agent here applications. In one study, regenerated BC scaffolds
allowed this material to overcome limitations of pre- with controlled porosity were fabricated with the
vious antibacterial native BC where leaching of active addition of NaCl crystals as a ‘porogen’ in the BC-
agents can reduce the long-term antibacterial proper- solution (Khan et al. 2015a). Using the dissolution-
ties of the material. A study where zinc oxide NPs regeneration route here, with NMMO as the solvent,
were used as the antibacterial additive used dissolu- allowed the authors to have better control over the
tion and regeneration to create BC films with differ- BC-to-porogen ratio and the overall shape of the
ent thicknesses (Ul-Islam et al. 2014). The addition of produced material. Figure 4 shows the difference in
nanoparticles here not only gave the films antibacte- shape uniformity and morphology between the regen-
rial properties, but also enhanced the tensile strength erated BC films with (3-D rBC) and without (Simple
and Young’s modulus. rBC) porogens. By characterising the cell adhesion,
proliferation and viability in the regenerated BC scaf-
Tissue engineering and skin regeneration folds, the study indicated this material was suitable
for tissue regeneration applications. In a second study
Another research group has focused on regener- by the same group, a similar fabrication method was
ated BC for tissue engineering and skin regeneration used wherein a gelatin-NMMO solution was mixed

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Fig. 4  Photographs (a)

and scanning electron
microscope images (b) of
scaffolds produced using
regenerated BC (simple
rBC) and regenerated BC
with porogens (3-D-rBC)
(Khan et al. 2015a) repro-
duced with permission from
Royal Society of Chemistry

with a solution of BC-NMMO along with the addi- as the residual gelatin remaining in the regenerated
tion of NaCl crystals as porogens (Khan et al. 2016). BC scaffold, resulted in scaffolds with high biocom-
The addition of gelatin helped to enhance the bio- patibility suitable for skin regeneration applications
compatibility and microporosity of the scaffolds for (Khan et al. 2018).
tissue regeneration applications. A third study created
a regular porous structure in regenerated BC scaffolds Drug delivery
through the addition of gelatin microspheres (GMS).
The GMS were fabricated using a water-in-oil emul- Researchers have also investigated the use of regen-
sion method and were sieved to obtain particles with erated BC in drug delivery systems. Badshah et al.
a consistent size range. The GMS were added to a (2020) dissolved BC in NMMO and loaded the
solution of BC-NMMO, with brief stirring (1 min) solution with model drugs (famotidine and tiza-
before the solution was regenerated using a film cast- nidine) to study drug release from regenerated
ing method. The uniform porous structure, as well BC-drug matrices (Badshah et al. 2020). Here, the

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dissolution-regeneration route was used to more eas- the NaOH in the solvent system weakening the cross-
ily control the thickness of matrices compared to as- linking points. This weakening of cross-links was
produced, native BC. Further, the amorphous nature proposed as the reason behind the lower strength and
of regenerated BC is thought to have been responsible higher swelling ratio of these hydrogels compared to
for enhanced biodegradation, a desired characteris- their non-solubilised hydrogels. The high porosity
tic of oral drug delivery materials. The drug release and swelling behaviour of the regenerated BC-AM
kinetics indicated these matrices were suitable for hydrogels led to a higher drug loading capacity and
immediate release drug delivery applications. In higher drug release percentage, indicating their supe-
another study targeting drug delivery applications, riority in drug release applications compared to non-
Pandey and colleagues fabricated superabsorbent solubilised BC.
hydrogels using graft polymerisation of acrylamide
(AM) on BC in the solution state (Pandey et al. 2013). Alternative fabrication methods
Interestingly, the authors also prepared a non-solu-
bilised BC hydrogel following the same fabrication Many of the materials for biomedical applications
technique, replacing BC dissolution with dispersion discussed so far have involved film or mould casting
in water. The regenerated BC-AM hydrogels were of cellulose solutions. Alternative fabrication meth-
determined to have a lower degree of cross-linking ods have involved electrospinning, drop-wise regen-
(characterised by % gel fraction) than the non-solubi- eration/coagulation, and partial cellulose dissolution.
lised hydrogels. The authors believed this was due to Figure 5 contains a schematic comparing the common

Fig. 5  Alternative fabrication methods for regenerated BC in biomedical applications including drop-wise coagulation and electro-
spinning compared with the commonly used film casting method. Created with Chemix (https://​chemix.​org)

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 10176 Cellulose (2024) 31:10165–10190

film casting technique with electrospinning and drop- used as a chemical cross-linking agent whilst physical
wise bead production. An alternative production cross-linking occurred between hydroxyl groups on
method was employed by Jayani and colleagues who cellulose chains. The authors took advantage of the
dissolved BC in Trifluoroacetic acid (TFA) and cre- difficulty of dissolving BC through partial dissolu-
ated nanofibrous mats via electrospinning (Jayani tion, the extent of which was controlled by regulat-
et al. 2020). The BC solution was mixed with a poly- ing the concentration of LiOH and urea used. Inter-
vinyl alcohol (PVA)-TFA solution to allow electro- estingly, Zhang and colleagues showed that hydrogels
spinning. As this product was targeted at food-based synthesised using cotton as the cellulose source in an
pharmaceutical applications care was taken to confirm identical process had inferior mechanical properties
that all solvent had been removed from the resulting compared to the regenerated BC hydrogels. The BC
nanofibers. A probiotic, which was incorporated onto hydrogels had tensile strength twenty times higher
the regenerated BC nanofibers via an adsorption- and a compressive strength 4.5 times higher than the
incubation method, showed a survival rate of 71.1% regenerated cotton hydrogels. The authors attributed
after 24 days in storage. Another interesting material these enhanced properties to the partially dissolved
form made available through BC dissolution is hydro- cellulose matrix supported by undissolved bacterial
gel beads, such as the regenerated BC-chitosan beads cellulose fibres. The strength of these hydrogels sug-
prepared by Kim et al. (2017b) for application as gests their application in underwater equipment as
enzyme supports. The study focused on BC due to the well as biomedical applications such as implantable
desirable high degree of polymerisation and purity of ionic devices and tissue engineering scaffolds.
this material and utilised the dissolution-regeneration
route to fabricate uniform shapes that are difficult to Composites and nanocomposites
obtain from native BC. Chitosan was first dissolved in
an ionic liquid (EmimOAc) before the addition of BC, Similar to the previously mentioned study by Zhang
followed by drop-wise regeneration in distilled water. and colleagues, Soykeabkaew et al. (2009) also used
Regenerated microcrystalline cellulose (MCC)-chi- partial dissolution to produce all-cellulose nanocom-
tosan beads were also prepared using a similar tech- posites, aiming to utilise the high modulus of BC
nique for comparison. The BC-chitosan beads showed nanofibers as reinforcement. The BC pellicles were
higher adsorbed protein content and better thermal immersed in a mixture of distilled water, acetone and
stability of the lipase protein than the beads produced N,N-dimethylacetamide (DMAc) before being par-
using MCC. The study suggested this was due to an tially dissolved in the wet state using lithium chloride
increased number of interactions between BC and the (LiCl)/DMAc. Figure 6 demonstrates the produc-
lipase, due to the BC’s higher degree of polymerisa- tion process schematically. The authors showed that
tion. However, it should be noted that due to limita- the mechanical properties of the composites could
tions caused by the high viscosity of the BC-chitosan be controlled by the immersion time in the solvent.
solution, the BC-chitosan beads had a higher ratio of BC immersed for 10 min had higher tensile strength
chitosan to cellulose compared to the MCC-chitosan and elongation compared to an undissolved BC sheet,
beads. Partial dissolution of BC in LiOH/urea alka- whilst immersion times of 40 and 60 min led to sig-
line solvent was utilised by Zhang et al. (2019) who nificantly lower strength but far greater elongation
fabricated self-reinforced double-crosslinked bacte- at break, with strain values as high as 30%. These
rial cellulose hydrogels with a nanofibre-network- composites provide an interesting example of mono
self-reinforced structure. Epichlorohydrin (ECH) was materiality combined with a sustainable raw material.

Fig. 6  Schematic of the

selective dissolution of cel-
lulosic fibres (Soykeabkaew
et al. 2009) reproduced with
permission from Springer
Nature

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Other single-material composites, such as all-poly- materials, these values are promising especially given
propylene composites (Cabrera et al. 2004), achieve the flexibility, biocompatibility, and biodegradabil-
composite-like properties without introducing the ity of BC. Two different conductive regenerated BC
common separation problems associated with recy- composites were found in literature, both focusing
cling composite materials. However, single-material on applications in electrical field-enhanced wound
composites made form synthetic polymers are still healing. The studies used NaOH/urea solution as
hindered by sustainability problems, as they are pro- a solvent and incorporated electrically conductive
duced from non-renewable sources. Achieving a additives to the BC in the solution state, along with
variation in properties for films produced entirely of the use of ECH as a cross-linker. Wang et al. (2020)
cellulose prove the concept of a recyclable compos- introduced carbon nanotubes (CNT) to a solution or
ite that is also biodegradable and renewable. Other BC and polypyrrole (PPy) and increased the conduc-
authors have demonstrated the selective dissolution of tivity from 3.47 × ­10–10 S/cm for neat regenerated BC
BC in ionic liquid (BmimCl), however the study was to 1.67 × ­10–3 S/cm for the BC/PPy/CNT material.
preliminary and no concrete characterisation was pro- Alternatively, Mao et al. (2020) were able to achieve a
vided (Suryanto et al. 2020). maximum conductivity of 7.04 × ­10–4 S/cm at a 2wt%
loading of MXene (­Ti3C2Tx). The incorporation of
Electronics and conductive regenerated BC MXene in regenerated BC led to significant increases
in tensile strength, compressive strength, and Young’s
Demonstrative of the diverse application areas that modulus, though the compressive and tensile strains
have been investigated for regenerated BC is the decreased.
application field of electronics and sensors. A review
by Pan et al. (2023) explored literature around bac- Filtration and separation
terial cellulose utilisation in hydrogel sensors. Some
cases where BC was dispersed or homogenised in In possibly one of the earliest works involving regen-
water and cast or shaped were referred to as ‘regen- erated BC, Phisalaphong and colleagues produced a
erated’ BC. It is important to note that, in terms of membrane from BC dissolved in a NaOH/urea solu-
cellulose, regenerated cellulose refers to cellulose that tion (Phisalaphong et al. 2008a, b). Citing the films
has been dissolved and regenerated, with the crystal- chemical stability and biocompatibility, the authors
line structure of the regenerated material varying sig- suggested these membranes might be applied in nano-
nificantly from that of native cellulose. The few stud- separation for medical or chemical processes. In a
ies that have focused on the use of regenerated BC in following study by the same group, regenerated BC/
electronics and sensors include its use as a compo- alginate films were produced by mixing solutions
nent in a triboelectric nanogenerator (TENG) (H.-J. of BC in NaOH/urea solution and alginate in water,
Kim et al. 2017a), as well as composite hydrogels for before casting and coagulating the membranes (Phis-
use in electrical field-enhanced wound healing (Mao alaphong et al. 2008a, b). Through the addition of alg-
et al. 2020; Wang et al. 2020). inate, the water vapor permeability (WVP) and degree
Kim et al. (2017a) dissolved BC in ethyl acetate, of swelling of the membranes were improved, whilst
made possible by earlier nitration of the BC material. the alginate had the effect of reducing the mechanical
The top electrode of the TENG consisted of the regen- properties. Due to the nano-porosity of the film, the
erated BC film cast atop a copper foil, with the oppos- authors suggested its use in separation processes and
ing side of the foil being covered by a polypropylene indicated that tests for application in pervaporation
film. The bottom electrode consisted of another fric- were underway. Other authors have aimed to broaden
tion part, comprising copper foil, with the side oppo- the potential application areas of BC by creating
site to the BC covered with a polyoxymethylene plate. separation membranes, with a focus on determining
The requirement for dissolution of BC here comes the solubility of BC in a common cellulose solvent,
from the need to produce a film with uniform proper- LiCl/DMAc (Shen et al. 2010). Excessive process-
ties. Under a small force the TENG reached an accu- ing was required, in the form of grinding to a pow-
mulative charge of 8.1 µC/m2 and a peak-power den- der and a pre-treatment/activation step, to dissolve 3
sity of 4.8 mW/m2. While lower than other synthetic wt% BC. The type of cellulose used (BC or PC), as

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well as the type of anti-solvent/coagulation medium, crystallised salt particles when not subjected to light.
should be selected based on the targeted application. The evaporation properties of the composite biofoam
Regenerated cellulose hydrogels produced using low showed stability over twenty 8-h tests and had better
DP α-cellulose coagulated in methanol were shown performance than other BC-based solar evaporators.
to have a better carbohydrate-binding module (CBM) This work demonstrates how the properties of BC can
migration rate than a high DP regenerated BC coun- be tailored to specific advanced applications using the
terpart, whilst BC hydrogels regenerated in water had dissolution-regeneration route.
a higher accessible surface area than both BC regen-
erated in methanol, and α-cellulose regenerated in Packaging
water or methanol (Johns et al. 2017).
By using a novel fabrication technique, composite The motivation behind the investigation of renewable
foams have been created that contain regenerated BC, and sustainable materials for packaging applications
as well as native BC mixed with an active component. is clear–plastic packaging is synonymous with pollu-
A composite biofoam was created for application in tion. Development of BC packaging meets criteria for
solar-powered water treatment, wastewater purifica- renewability as well as biodegradability. The biocom-
tion and solar desalination (Zhang et al. 2021). This patibility of BC may have further advantages in terms
bi-layered material consisted of one layer comprising of food packaging. It is understood that developing
a vacuum-filtered dispersion of native BC and CuS as and improving BC dissolution methods may open up
the photothermal layer and another layer comprising opportunities of BC application in areas such as pack-
ECH cross-linked regenerated BC as the water trans- aging, food, and pharmaceuticals (Tilak et al. 2019).
porting, thermally insulating layer. The production Both TFA and DMSO have been trialled as BC sol-
process for this material is described in Fig. 7 below. vents, with findings indicating that DMSO could not
The biofoam exhibited self-cleaning and self-floating dissolve BC under conventional heating, microwave
abilities whilst also achieving a high evaporation effi- heating or cold treatment. TFA-BC solutions were
ciency. The self-floating characteristic was attributed successfully prepared using the above methods, with
to the regenerated BC foam, whilst the self-cleaning microwave heating resulting in the fastest dissolution
was characterised by the foam’s ability to dissolve across all cellulose concentrations tested (2–5 wt%)

Fig. 7  Fabrication process for the regenerated BC biofoam for solar desalination (Zhang et al. 2021) reproduced with permission
from Elsevier

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Cellulose (2024) 31:10165–10190 10179

(Tilak et al. 2019). An important characteristic of dissolving pulp, and BC) has been dissolved in ionic
food packaging, or packaging wrap in general, is a liquid (EmimOAc) for use as a 3D-printable ink
good mechanical strength combined with suitably (Markstedt et al. 2014). The dissolution of cellulose
high elongation. Regenerated BC films for food pack- here allows for broader manufacturing techniques
aging have been fabricated with a range of mechani- for controlling the structure and property of cellu-
cal properties, ranging from high strength/low elon- losic materials. Owing to BCs high degree of poly-
gation (Yu et al. 2023), moderate strength/moderate merisation, the solution formed from EmimOAc-
elongation (Shanshan et al. 2012), and low strength/ BC had favourable viscoelastic properties for use
high elongation (Wang et al. 2021). This shows the as an ink in 3D bioprinting. Solutions containing
versatility of regenerated BC when different func- 2 and 4 wt% BC and 4wt% dissolving pulp showed
tional additives are included. Similar to other applica- the best shear-thinning properties for 3D printing,
tion areas, additives can be included in the solution allowing the solution to thin during extrusion under
state (Wang et al. 2021; Yu et al. 2023), or via immer- shear before regaining high viscosity and maintain-
sion of the regenerated BC (Shanshan et al. 2012). ing its shape as a gel-like material until coagulation.
A regenerated BC film cross-linked using an envi- An alternative technique was employed to produce
ronmentally friendly cross-linking agent (citric acid) a 3D printable, UV curable dispersion of regener-
was able to preserve banana for longer than a com- ated BC mixed with acrylic acid (AA) (Smirnov
mercial PVC plastic wrap (Yu et al. 2023). Using a et al. 2021). The polymerizable component (AA)
food additive as the cross-linking agent allowed for an was dissolved in either the ionic liquid BmimCl or
edible film and expedited the degradation of the film the deep eutectic solvent (DES) choline chloride
in soil, being fully degraded after 11 days. Packaging (ChCl) and BC was mixed into these solutions. The
produced from regenerated BC with additives such as mixture of AA and partially dissolved (indicated by
citric acid, gelatin and MgO or glycerin had suitable XRD and AFM) BC were regenerated as a disper-
properties for food preservation owing to the films’ sion in water before cross-linking and photo initiat-
antibacterial properties (Yu et al. 2023), hydrophobic- ing agents were added, and this dispersion of regen-
ity (Wang et al. 2021), low water vapor permeability erated AA-BC was used as the 3D printing ink.
(Shanshan et al. 2012; Wang et al. 2021), and dense Figure 8 shows a schematic of the production and
and uniform structure (Shanshan et al. 2012). characterisation process used. The choline-treated
3D printable dispersion performed better in terms
3D printing of rheological properties and mechanical properties.
The enhanced mechanical properties of the choline-
The high viscosity of solutions containing high DP based material were thought to be caused by better
BC has attracted research into 3D printing ink appli- grafting of poly(acrylic acid) onto the surface of BC
cations. Cellulose from different sources (MCC, during UV curing.

Fig. 8  Schematic of BC processing and characterisation for UV-curable 3D printed BC-AA samples reproduced from Smirnov et al.
2021 under CC BY licence (Smirnov et al. 2021)

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Limitations in production, processing, BC (Quijano et al. 2024). Agitation is commonly

and dissolution of bacterial cellulose achieved by shaking the fermentation vessel or stir-
ring the fermentation media (Siti et al. 2014; Tanskul
Production et al. 2013). Whilst static fermentation results in a
continuous BC film, agitated fermentation leads to
Production of native bacterial cellulose involves inoc- the formation of irregular BC pellets with properties
ulating a nutrient-containing substrate with an appro- that may differ considerably from those of the stati-
priate strain of bacteria. The previously mentioned cally produced BC film (Lahiri et al. 2021). Alterna-
dessert food, nata de coco, is bacterial cellulose com- tive fermentation methods have been investigated to
mercially produced from the fermentation of coconut combine the benefits of static and agitated fermenta-
water, which contains all the appropriate nutrients. tion including rotating disk reactors (Jagtap and Dast-
At the laboratory scale the nutrient requirements of ager 2024; Pa’e et al. 2011), bubble column reactors
the substrate are commonly satisfied using the Hes- (Song et al. 2009) or shaking followed by static fer-
trin Schramm (HS) media, comprising glucose, yeast mentation (Andritsou et al. 2018).
extract, peptone, disodium phosphate, and citric
acid (Quijano et al. 2024). The cost of HS media is Drying and hornification
a potential barrier to commercialisation, which has
led to the search for alternatives. A recent review by Regardless of the bacterial strain, carbon and nutrient
Quijano et al. (2024) revealed that almost 300 differ- source, or fermentation method used for BC produc-
ent ‘ingredients’ from more than 12 different indus- tion, most cases of BC dissolution require the mate-
tries have been studied as substrates for BC produc- rial to be dried first. The drying of cellulose often
tion. Additional to alternative substrates, alternative leads to some extent of hornification. Hornification
strains of bacteria are also being investigated with the is characterised by the mechanical stiffening of cel-
aim of increasing the cellulose yield per litre of fer- lulosic material and a decrease in the water retention
mentation media. For example, Thorat and Dastager value, caused by cyclic wetting and drying. Three
(2018) identified a new strain of high-yield cellulose- common mechanisms exist for explaining hornifica-
producing bacteria by studying bacterial isolates from tion; physical cross-linking of fibrils via hydrogen
a range of rotten fruits and fermented beverages. bonding and van der Waals interactions (Aghajanza-
During BC growth, the aerobic bacteria are located deh et al. 2023; Ballesteros et al. 2017; Sharma et al.
at the liquid–air interface of the fermentation media 2014), co-crystallisation of neighbouring crystallites
(Maneerung et al. 2008; Siti et al. 2014; Zhong 2020). (Idström et al. 2013; Newman 2004), and chemical
The polymerisation of crystalline cellulose fibrils bonding of fibrils via lactone bridges (Aghajanzadeh
acts as a motive force for bacterial cells and this et al. 2023; Fernandez Diniz et al. 2004). Figure 9
force, coupled with gravity, is thought to facilitate demonstrates how hornification might cause irrevers-
the downward movement of the cellulosic film dur- ible bonds between cellulose fibrils, inhibiting the
ing fermentation (Ruan et al. 2016). This film exists rewetting of cellulose. Whilst the exact mechanisms
as a hydrated pellicle containing over 99% water by of hornification remain to be concluded, this phenom-
mass. The high surface area and hydrophilicity of the enon is common to the pulp and paper industry and
cellulose nanofibrils cause the space between fibrils has been extensively studied. For cellulose from plant
to become filled with water from the surrounding sources such as wood, the extent of hornification is
aqueous fermentation media (Manoukian et al. 2019). related to the level of hemicelluloses and lignin pre-
BC is most commonly produced under static condi- sent. High yield pulps (with higher hemicellulose and
tions (Quijano et al. 2024), where the fermentation lignin content) tend to experience less hornification
media is not disturbed for the duration of BC growth. than the purer cellulose pulps (Spence et al. 2010). As
However, agitated fermentation is another production hornification arises from the close contact of adjacent
method that has become common. Agitating the fer- cellulose fibrils, the hemicelluloses and lignin are
mentation media during BC growth aims to ensure thought to act as a barrier to direct bonding of cel-
homogeneous distribution and supply of oxygen (Siti lulose fibrils (Palme et al. 2016; Sellman et al. 2023).
et al. 2014) and reduce the time required to produce For bacterial cellulose, the lack of hemicelluloses and

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Fig. 9  Schematic of the

hornification of cellulose
fibrils when cycled from a
wet state (left) to a dry state
(middle) and after rewetting
(right)

lignin to function as spacers between fibrils coupled adjacent fibrils and subsequent cross-linking (Martins
with the higher water content is a possible cause of et al. 2020). A common additive to nanocelluloses
higher susceptibility to hornification. to prevent hornification is carboxymethyl cellulose
The stiffening and lower water retention values (CMC). This has also been used for bacterial cellu-
caused by hornification are problematic for many lose (Martins et al. 2020), as well as additives such
applications, and the aggregation of cellulose fibrils as sorbitol (Rossi et al. 2023) and glycerol (Ulmer
may affect the solubility of cellulose by decreasing 2020). Müller et al. (2014) investigated bacterial cel-
the reactivity (Ferreira et al. 2020). Ferreira and co- lulose for drug delivery applications. In their study,
authors concluded that, in some circumstances, the freeze-drying was determined to be unsuitable for the
changes in physical structure of cellulose caused by given application. To enhance rehydration of the BC
hornification may have more impact on reactivity than they instead successfully incorporated additives such
the viscosity-average molecular weight of cellulose as ­MgCl2 to inhibit physical cross-linking during dry-
(proportionate to the degree of polymerisation) (Fer- ing (Müller et al. 2014).
reira et al. 2020). Another study on the influence of
dried and never-dried cellulose on solubility in aque- Dissolution of bacterial cellulose
ous solvents showed that while drying cellulose may
be beneficial to dissolution in some solvents, it was Cellulose from any origin is known to be poorly solu-
inhibitive to dissolution in other solvents (Spinu et al. ble in water or common organic solvents (Rabideau
2011). For cellulose in general, methods to reduce or and Ismail 2015). It is often stated that the insolu-
avoid hornification include alternative drying tech- bility of cellulose is caused by its strong, highly
niques such as freeze-drying, solvent exchange, the branched hydrogen bonding network (Acharya et al.
addition of capping agents prior to drying (Aghajan- 2021; Heinze 2016; Makarov et al. 2019; Pinkert
zadeh et al. 2023). Studies into drying and dissolu- et al. 2009; Visakh and Thomas 2010) whilst others
tion of BC often focus on freeze-drying (Andree et al. argue that, in aqueous environments, cellulose insolu-
2021; Illa et al. 2019; Stanisławska et al. 2020; Zhang bility is more dependent on the hydrophobic interac-
et al. 2011). Research into the effect of drying meth- tions (Lindman et al. 2010; Medronho and Lindman
ods on the properties of bacterial cellulose by Andree 2015; Wohlert et al. 2022). Other factors thought to
et al. (2021) found that, compared to oven drying at contribute to cellulose insolubility are the high crys-
100 °C and drying in a climate chamber at 23 °C, only tallinity (Acharya et al. 2021; Ahmed et al. 2020;
freeze-drying could markedly reduce stiffening on Makarov et al. 2019; Pinkert et al. 2009), high molec-
drying. Additives can also be used to create a steric ular weight (or degree of polymerisation) (Aghmih
barrier during drying which prevents the contact of et al. 2023; Lindman et al. 2010), and chain stiffness

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 10182 Cellulose (2024) 31:10165–10190

hindering conformational changes in the amphiphilic such as wood pulp (El Seoud et al. 2007; Swatloski
polymer (Glasser et al. 2012; Lindman et al. 2010). et al. 2002), waste cotton (Haslinger et al. 2019),
Some argue that insolubility is not due to thermo- waste paper and cardboard (Ma et al. 2016) and dif-
dynamic or kinetic limitations, but instead is related ferent plant celluloses (Lethesh et al. 2020). Due to
to the hierarchical nature of cellulose (Singh et al. the benign processing of BC, the DP remains high.
2015). The structure and organisation of cellulose in Studies that reported dissolution of bacterial cel-
solution has also been debated. Authors have pro- lulose showed lower concentrations of cellulose in
posed that solutions of cellulose contain dispersions solution, however most of these studies included bac-
of nano-scale crystalline agglomerations of undis- terial cellulose with very high DP (1500–6500). The
solved cellulose molecules (Sashina 2018; Singh et al. study which obtained the highest value for solubility
2015; Zhou et al. 2022), whilst others argue that cel- for BC (12.2 wt%) involved depolymerising the BC
lulose chains are completely separated and molecular to a DP of 634. Researchers suggested that the lower
dispersion is achieved (Sayyed et al. 2019). A study solubility of BC was due high purity (Raghuwanshi
by Zhou et al. (Zhou et al. 2022) showed that the type et al. 2021), high degree of polymerisation (Makarov
of solvent used determined if a molecularly dispersed et al. 2020; Schlufter et al. 2006), high crystallin-
solution was achieved. Examples of possible cellulose ity (Makarov et al. 2019; Raghuwanshi et al. 2021),
arrangements in solution discussed in the above lit- unique hierarchical fibrillar structure (Raghuwanshi
erature are shown conceptually in Fig. 10 below. et al. 2021), extremely fine fibrils (Schlufter et al.
At the time of writing, there were no studies where 2006), lack of other biopolymers (Schlufter et al.
dissolution of BC exceeded 12.2 wt% (Heinze et al. 2006), and the ordered structure of macromolecules
2008; Soares Silva et al. 2023), whereas cellulose (Makarov et al. 2019).
from sources such as plants and MCC is commonly
dissolved at higher concentrations in solvents such
as ionic liquid, and can reach as high as 40–45 wt%
(Lethesh et al. 2020). High concentrations of cel-
lulose have been dissolved using cellulose sources

Fig. 10  Conceptual schematic of cellulose organisation and structure before and after dissolution demonstrating molecular disper-
sion, a combination of molecular dispersion and aggregation of crystalline regions, and a dispersion of crystalline agglomerates

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Cellulose (2024) 31:10165–10190 10183

Future perspectives costs and cellulose solvent costs is essential. Regard-

ing properties of regenerated BC materials, develop-
The environmental impacts surrounding the use of ing sustainable methods to functionalise regenerated
fossil-based synthetic polymers has become an inter- bacterial cellulose will help to broaden the applica-
esting and concerning problem not only for academia tion areas. For example, functionalising regenerated
and industry, but also for the general public and gov- BC films for hydrophobicity could lead to a material
ernments around the world. It is apparent now more that is a viable alternative to plastic wrap/packaging
than ever that solutions are required that will enable produced from hydrophobic synthetic polymers. The
the transition from synthetic polymers to sustainable incorporation of nanoparticles into regenerated BC
biopolymers and renewable materials. Cellulose has materials may also play an important role in creating
been a front-runner in the search for a strong, bio- functional and advanced materials from BC.
degradable polymer of the future. Wood is the main
source of cellulose for many investigative and com- Acknowledgments The authors acknowledge the sup-
port from the Australian Research Council (ARC) Hub for
mercial applications. However, as production of cel- Future Fibres, which is a collaboration between the Austral-
lulose-based materials increases, there will be excess ian Research Council (ARC), Deakin University, and Industry
demand on the World’s forests. With climate change Partner Organisations to create high impact solutions using
an ever-pressing issue, excessive deforestation would the world class research teams and facilities at the Institute for
Frontier Materials (IFM). CRediT: Idea for article: Lucas Ros-
be detrimental to controlling greenhouse gas levels son and Nolene Byrne; literature search and analysis: Lucas
and thus global warming. Bacterial cellulose offers Rosson and Boon Tan; draft writing: Lucas Rosson; revision
an alternative source to produce cellulosic materials, and editing: Lucas Rosson, Nolene Byrne, Boon Tan and
with potentially less resource-intensive requirements Wayne Best.
compared to plant-based cellulose. Capable of being
Author’s contribution LR and NB had the idea for the arti-
produced from waste with a smaller land footprint cle LR wrote the draft manuscript. LR, WB and BT acquired/
than plant cellulose with little-to-no associated defor- produced the figures for the manuscript. LR and BT conducted
estation, bacterial cellulose may prove to be a truly the literature search and analysis. All authors reviewed and
edited the manuscript.
sustainable feedstock for future applications of cel-
lulosic materials. Coupled with the reduced resource Funding Open Access funding enabled and organized by
requirements of bacterial cellulose synthesis, the high CAUL and its Member Institutions. The work performed by
purity of BC holds the advantage of requiring only a Lucas Rosson and Nolene Byrne was funded by the Australian
benign washing process to obtain pure cellulose. Research Council (ARC) Hub for Future Fibres also known as
the ARC Research Hub for Functional and Sustainable Fibres
To enhance the utilisation of BC as an alternative (Grant number IH210100023). The work performed by Boon
to plant cellulose, work needs to be done to discover Tan and Wayne Best was funded by Nanollose Ltd.
more about this material. Further research into the
mechanisms of bacterial cellulose dissolution and Data availability No datasets were generated or analysed
during the current study.
how the structure of the material affects dissolution
is required. The development of unique solvents or Declarations
processes to allow the dissolution of higher concen-
trations of high DP bacterial cellulose may lead to a Conflict of interest The funding recieved from the Australian
Research Council was for the salary of Lucas Rosson. Wayne
more wide-spread interest in this material. It may also Best (CEO) and Boon Tan (Operations and Qaulity Manager)
be beneficial to utilise cellulose depolymerisation are employees and shareholders of Nanollose Ltd. Nanollose is
techniques that are environmentally benign to aid in a biotechnology company that works with regenerated bacterial
dissolution, as we have seen the solubility of cellu- cellulose. There may be a percieved conflict of interest in this
publication reviewing regenerated bacterial cellulose. However,
lose depends strongly on the DP. Further to this, any the content of the review does not contain biased views for prod-
solvent used for the processing and dissolution of BC ucts or processes manufactured or owned by Nanollose.
should aim to be highly recyclable to ensure the envi-
ronmental benefits of using this material are not out- Ethical approval Not Applicable.
weighed by environmental impacts of the dissolution
process. In terms of developing materials and pro- Open Access This article is licensed under a Creative Com-
cesses commercially, reducing both BC production mons Attribution 4.0 International License, which permits

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 10184 Cellulose (2024) 31:10165–10190

use, sharing, adaptation, distribution and reproduction in any Andritsou V, de Melo EM, Tsouko E, Ladakis D, Maragk-
medium or format, as long as you give appropriate credit to the oudaki S, Koutinas AA, Matharu AS (2018) Synthesis
original author(s) and the source, provide a link to the Crea- and characterization of bacterial cellulose from citrus-
tive Commons licence, and indicate if changes were made. The based sustainable resources. ACS Omega 3(8):10365–
images or other third party material in this article are included 10373. https://​doi.​org/​10.​1021/​acsom​ega.​8b013​15
in the article’s Creative Commons licence, unless indicated Avcioglu NH (2022) Bacterial cellulose: recent progress in
otherwise in a credit line to the material. If material is not production and industrial applications. World J Micro-
included in the article’s Creative Commons licence and your biol Biotechnol 38(5):86. https://​doi.​org/​10.​1007/​
intended use is not permitted by statutory regulation or exceeds s11274-​022-​03271-y
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from the copyright holder. To view a copy of this licence, visit M, Khan T (2020) Development and evaluation of drug
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