Cellulose (2021) 28:8229–8253

https://doi.org/10.1007/s10570-021-04020-4 (0123456789().,-volV)
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89().,-volV)

REVIEW PAPER

A review of bacterial cellulose: sustainable production

from agricultural waste and applications in various fields
Leire Urbina . Marı́a Ángeles Corcuera . Nagore Gabilondo .
Arantxa Eceiza . Aloña Retegi

Received: 25 November 2020 / Accepted: 14 June 2021 / Published online: 10 July 2021
Ó The Author(s), under exclusive licence to Springer Nature B.V. 2021

Abstract Bacterial cellulose (BC) is a polymer with
interesting conformation and properties. BC can be
obtained in different shapes and is easily modified by
chemical and physical means, so its applications in the
production of new materials and nanocomposites for
different purposes have been in the focus of many
research projects. However, one of the major chal-
lenges to address in bacterium-derived polymer tech-
nology is to find suitable carbon sources as substrates
that are cheap and do not compete with food produc-
tion for achieving large scale industrial applications.
Agricultural wastes are defined as the residues from
the growing and processing of raw agricultural
products such as crops, fruits, vegetables and dairy
products. Their composition can vary depending on
the type of agricultural activity and harvesting condi-
tions, but these residues are suitable for the production
of BC. The aim of this review is to give insight into the
production of BC using agro-wastes and an overview
of the most interesting and novel applications of this
biopolymer in different areas i.e. environmental
applications, optoelectronic and conductive devices,
food ingredients and packaging, biomedicine, and 3D
printing technology.

L. Urbina (&)  M. Á. Corcuera  N. Gabilondo 

A. Eceiza  A. Retegi
Materials ? Technologies’ Group, Engineering School of
Gipuzkoa, Department of Chemical and Environmental
Engineering, University of the Basque Country (UPV/
EHU), Pza. Europa 1, 20018 Donostia - San Sebastián,
Spain
e-mail: leire.urbina@ehu.eus
M. Á. Corcuera
e-mail: marian.corcuera@ehu.eus
N. Gabilondo
e-mail: nagore.gabilondo@ehu.eus
A. Eceiza
e-mail: arantxa.eceiza@ehu.eus
A. Retegi
e-mail: alona.retegui@ehu.eus

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Graphic abstract
Agricultural waste as feedstock
Bacterial growth and polymer production
3D-nanoscale network of cellulose
Keywords Bacterial cellulose  Agro-wastes 
Environmental applications  Optoelectronic devices 
Food industry  Biomedical field  3D printing
Introduction
Cellulose is regarded as the most abundant biopolymer
in nature. It is a semicrystalline polysaccharide, a
linear homopolymer of D-glucopyranose residues
(C6H11O5) linked together by b-(1 ! 4)-glycosidic
bonds. Each glucose monomer of cellulose consists of
three hydroxyl groups, positioned at C6 for the primary
hydroxyl groups and at C2 and C3 for the secondary
hydroxyl groups that can form intra and intermolec-
ular hydrogen bonds. Therefore, due to this chemical
constitution and spatial conformation, cellulose often
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Cellulose (2021) 28:8229–8253
Biomedical applications
Wound-healing systems
Blood vessels
Drug delivery
Food ingredients and packaging
Stabilizing agent
Thickening agent
Multilayers
Nanoreinforcements
Environmental applications
Heavy metal retention
Agricultural remediation
Oil/water separation
Optoelectronic and conductive devices
Thin film transistors
Solar cells
Organic light-emitting diodes
3D printing
Bioinks
Specific geometries
presents a highly ordered structure. It is the main
structural component of the primary cell wall of most
plants where it is combined with other components,
hemicelluloses and lignin. Thus, for the isolation of
cellulose, the raw material must be subjected to
chemical, enzymatic or mechanical treatments
(Jawaid et al. 2017).
More interestingly, in 1886 Brown reported a type
of cellulose produced by bacteria, commonly known
as bacterial cellulose (BC) (Wang et al. 2019). The
kinds of bacteria that make cellulose are generally
Gram negative, strictly aerobic (require oxygen), non-
photosynthetic and can be found in vinegar, fruits and
vegetables or alcoholic beverages (Belgacem and
Gandini 2008; Castro et al. 2012). Apart from an
oxygen supply, these bacteria need organic substrates
i.e. carbon sources which can be obtained from
different raw materials such as fruit, vegetable or
Cellulose (2021) 28:8229–8253
lignocellulosic wastes and other nutrients such us
nitrogen, iron, zinc and vitamins (Hussain et al. 2019).
Among cellulose producing bacteria, a huge diversity
can be found along the cellulose biosynthesis path-
ways, as well as differences in the nanostructure of the
polymer or its chemical properties and crystallinity.
For example, Salmonella typhimurium and Escher-
ichia coli produce amorphous cellulose as a compo-
nent of their extracellular matrix while other bacterial
strains produce crystalline cellulose (Zogaj et al.
2001). Belonging to the latter group, the most
commonly studied genera of BC producers include
Acetobacter, Rhizobium, Sarcina, Agrobacterium,
Alcaligenes, etc. Komagataeibacter xylinus (previ-
ously named as Acetobacter xylinus, Acetobacter
xylinum or Gluconacetobacter xylinus) has been
considered as model microorganism for basic and
applied studies on cellulose (Mikkelsen et al. 2009;
Lavasani et al. 2017; Hernández-Arriaga et al. 2019).
Due to its ability to produce relatively high levels of
polymer, it is the most commonly studied source of
bacterial cellulose. In this case, BC is produced in the
air–liquid interface and it confers mechanical, chem-
ical and biological protection within the natural
habitat, so that the cells are trapped in the polymer
network, which keeps the bacterial population at the
air–liquid interface and thus allows the microorgan-
ism, a strict aerobic, to be close to the air (Fig. 1A, B).
With optimal culture conditions, cells grow while
BC is produced. To clarify this, a typical growth curve
(time-dependent increase in the microbial population)
for a bacterial population such Acetobacter xylinus (A.
xylinus) is shown in Fig. 1C. Different growth phases
can be observed within the growth curve: lag phase,
the exponential or log phase, the stationary phase, and
the death phase. The lag phase is an adaptation period
of the bacteria to the new environment, which in the
case of A. xylinus, may be influenced by the number of
inoculated cells, age and physiological state (Lestari
et al. 2014). Bacterial cellulose is produced during the
exponential phase (characterized by a period of the
exponential growth-the most rapid growth possible
under the conditions present in the batch system) and
the stationary phase (can be defined as a state of no net
growth). This was observed with A. xylinus in
pineapple juice and coconut water (Lestari et al.
2014). It was reported that the greatest bacterial
growth was after 6 days, although the maximum BC
8231
yield was achieved in 12 days when the use of sugar
on substrate was to be stationary.
In these kinds of bacteria, the cellulose synthesis
occurs in two main stages: the first is the conversion of
the carbon source in uridine diphosphoglucose
(UDPGlc), which is the substrate used to produce
cellulose, and the second stage is the formation of
cellulose and its secretion into the culture medium.
The process starts with the formation of UDPGlc,
followed by glucose polymerization into the b-1 ! 4
glucan chain and a nascent chain which forms ribbon-
like structure of cellulose chains formed by hundreds
or even thousands of individual cellulose chains.
These are extruded out through tiny pores present on
their cell envelope (Chawla et al. 2009; Lee et al.
2014). These chains combine and form elementary
fibers which further aggregate to form cellulose
ribbons (nanofibers) (Fig. 1D). The width/diameter
of the nanofibers can vary depending on the bacterial
strain and cultivation conditions, but the values range
from 5 to 100 nm, while the length is difficult to
determine due to the entanglement of the network, but
usually is micrometers in length (Urbina et al. 2017;
Halib et al. 2019). The ribbons generate a 3D network
with plenty of spaces between the nanofibers that lead
to a highly porous matrix with an expanded surface
area (Fig. 1E). The formed cellulose contains numer-
ous surface hydroxyl groups which form abundant
inter- and intra-fibrillar hydrogen bonds which explain
its mechanical strength, ability to be chemically
modified, and its hydrophilicity. The final BC is
subjected to a purification process to remove non-
cellulosic compounds, generally with NaOH or KOH,
and a smooth white gelatinous matrix with water
content about 99% is obtained (Fig. 1F).
Properties of bacterial cellulose
It is worth to note that during biosynthesis, the
packaging process leads to highly pure and crystalline
cellulose. BC has been reported to show higher
crystallinity than 80% while cellulose from plants
contains more amorphous regions and displays crys-
tallinity values ranging from 40–85% (Park et al.
2010). Figure 1G displays the arrangement of crys-
talline and amorphous regions in cellulose fibers. In
terms of crystallinity, cellulose displays six different
polymorphs, namely, I, II, IIII, IIIII, IVI and IVII
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Fig. 1 A Liquid culture medium with cellulose produced in the
air/liquid interface. B Microscopy image of Gluconacetobacter
medellinensis strain bacteria producing BC (bar 10 lm) and
schematic representation of bacterium extruding BC nanofibrils
through the pores of membrane. C Typical growth curve of
bacterial A. xylinus population. D BC biosynthesis process and
(Chand and Fahim 2008). Cellulose type I is the most
abundant form in nature and the one predominantly
produced by bacteria (they can also produce cellulose
II) (Van Zyl and Coburn 2019). The crystalline
structure of cellulose I is a mixture of two distinct
crystalline structure forms: cellulose Ia (triclinic) and
cellulose Ib (monoclinic). Depending on the source of
the cellulose, the relative abundance ratio of these two
allomorphs varies. Cellulose Ib is the main constituent
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Cellulose (2021) 28:8229–8253
molecular structure of BC. E Scanning Electron Microscopy
image of BC freeze-dried membrane. F BC wet purified
membrane. G Cellulose fibers arrangement with amorphous
and crystalline regions. H Typical X-ray diffractogram of BC
with crystallinity peaks indexed based on the cellulose Ia
of the cellulose from plants, while bacterial cellulose
is rich in the Ia allomorph (Castro et al. 2011). A
typical X-ray diffraction pattern of BC is displayed in
Fig. 1H, where three main peaks around 14.6°, 17°
and 22.8° with Miller indices of (100), (010) and (110)
are observed. The relative heights of these peaks differ
markedly from those on ideal diffraction patterns or
those from cellulose in plant cell walls because of
preferred orientation of the crystallites to the X-ray
Cellulose (2021) 28:8229–8253
beam (French 2014). Due to this high crystallinity,
presence of numerous strong hydrogen bonds and its
fiber conformation, BC shows excellent mechanical
performance even in wet state. For BC fibers, values
from 10–65 GPa and 80–800 MPa have been reported
for Young’s modulus and tensile strength, respectively
(Retegi et al. 2010; Wang et al. 2017). These values
usually vary depending on the bacterial strain, culti-
vation method, processing of BC and water content
(which has plastifying effect). In fact, BC has a strong
affinity with water, so in the swollen state values of
8 MPa and 1.5 MPa of Young’s modulus and tensile
strength, respectively, can be found (Urbina et al.
2018). Indeed, BC can be considered a hydrogel with a
high water holding capacity which can reach 100 times
its own weight. Moreover, BC is considered a
biodegradable material since the hydrophilicity and
water uptake capability are factors related to the
biodegradability. Hydrophilic materials are more
vulnerable to hydrolysis which is a process that
initiates the degradation of polymers, and allows the
material to be colonized by microorganisms and fungi,
which will have access to carbon as food (Torgbo and
Sukyai 2020). Due to this feature, the application of
BC for the production of biodegradable packaging
materials has been investigated (Urbina et al. 2016;
Azly et al. 2020). In addition to these interesting
properties and conformation, BC has proven to be
biocompatible in animal studies (de Lima et al. 2017;
Zhang et al. 2020). For successful biological implants,
the biomaterial must induce cellular expansion, inte-
gration and interactions with biological tissues, and it
has been demonstrated that biological cells can
anchor, scatter and integrate with bacterial cellulose
without toxic response, opening new applications in
the biomedical field such as tissue engineering and
regenerative medicine. Some of the applications
related to this area include development of artificial
blood vessels, artificial skin, biosensors, dental
implants, heat valves and bone regeneration (Silva
2018; Eslahi et al. 2020).
Moreover, this biopolymer shows a broad chemical
and physical modifiability. BC membranes possess
free hydroxyl groups on the surface, so they can be
easily modified with other polymers or additives to
obtain different surface characteristics such as lipo-
philic or hydrophilic, magnetic and optical properties
(Cichosz and Masek 2019; Sun et al. 2019). Besides,
BC has the possibility to be used as matrix or
8233
reinforcement of composites and hybrid materials
prepared through ex situ and in situ methods. Ex situ
methods consist on the combination of the purified BC
by the impregnation technique to fix, coat or infiltrate
the additive/polymer in the spaces between the
nanofibers of the BC membrane. Using in situ meth-
ods, BC-based composites can be obtained by the
addition of some additives or soluble polymers to the
culture medium so that the additive is incorporated
into the BC growing nanofiber network during its
biosynthesis. Both preparation methods lead to BC
with different properties and conformations (Khan-
delwal et al. 2016; Stumpf et al. 2018; Faisul Aris et al.
2019). Taking into account all these features, it is not
surprising that its production and combination or
modification with other materials for the development
of hybrid devices and composites have been exten-
sively studied in recent decades for many different
applications.
Cultivation methods and conditions
For the production of BC at lab scale, there are two
cultivation approaches that provide cellulose with
different microstructure and properties i.e. static and
dynamic/agitated cultivation. Static cultivation is a
simple and typical method for BC biosynthesis. In this
case, the BC membrane is produced in the air/liquid
interface of the liquid nutrient solution. Depending on
the flask or vessel used, variable shapes of the
membranes can be obtained, but in a film form as it
can be observed in Fig. 2A. The thickness of the
membrane depends on the incubation time which
usually does not exceed 14 days, because during long
fermentation times, there is an accumulation of
inhibitory metabolites such as glycolic and formic
acids and others (Esra Erbas Kiziltas et al. 2015). In
this cultivation method, cellulose is usually produced
in layers parallel to the film plane forming a very stiff
and strong structure, suitable for applications in
membrane technology, food packaging or scaffolds
in biomedicine (Urbina et al. 2016, 2018, 2019c;
Roman et al. 2019; Gromovykh et al. 2020). In
dynamic cultivation, BC is produced with agitation
and presents the form of spheres or pellets whose
shape and size depend on the agitation speed (that
controls the oxygen transfer) and cultivation time,
which usually is shorter than in static cultivation
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(A)
1 2
(B)
1 2
(C)
Static
(D)
Fig. 2 A BC produced in static cultivation in different shapes, 1
and 2 wet state, and 3 freeze-dried. B BC produced in dynamic
cultivation in sphere-like shape, 1 and 2 wet state, and 3 freeze-
dried. C SEM images of the cross-section of BC membrane
(Fig. 2B). In this case, due to the stress induced by the
movement, BC presents a more disordered structure,
lower crystallinity and higher water holding capacity
than the one obtained in static cultivation (Algar et al.
2014; Kosseva et al. 2020). Algar et al. 2014 observed
a well-organized structure formed by layers in BC
produced in static cultivation, while BC produced in
dynamic cultivation presented a disordered structure
(Fig. 2C). However, in the case of BC produced in
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Cellulose (2021) 28:8229–8253
3
3
Dynamic
produced in static cultivation (left) and dynamic cultivation
(right) (Algar et al. 2014). D BC produced in a plastic composite
support of a biofilm reactor (Cheng et al. 2009)
dynamic cultivation, the spherical conformation cou-
pled with the high water holding capacity open the
possibility to obtain new structures i.e. encapsulation
of active agents, especially interesting for drug
delivery systems (Hoshi et al. 2018; Urbina et al.
2020).
In terms of productivity, the observed tendency is
lower productivity in dynamic cultivation usually
related to the appearance of noncellulose-producing
Cellulose (2021) 28:8229–8253
mutants (Cel- mutants) which hinder or reduce BC
production (Wang et al. 2019). However, one of the
major challenges for implementation of bacterium-
derived polymers in current markets is to achieve
industrial scale production. For this, different biore-
actors have been developed such as oxygen enriched
bioreactors, biofilm reactors or rotating disk bioreac-
tors. Figure 2D shows BC produced in a biofilm
reactor (Cheng et al. 2009). In that case, BC was
produced attached to the plastic composite support
shaft due to the bioaccumulation of cells. Despite the
heterogeneous form, BC was produced in high yields
(7.05 g L-1) while maintaining the high crystallinity
(93%) and tensile strength and Young’s modulus of
34.2 MPa and 2,401 MPa, respectively. Moreover,
Lin et al. (2014) produced BC in a rotating disk
bioreactor. In that configuration, half of the area of the
plastic composite support disks was submerged in the
medium and the other half exposed to the atmosphere
and BC was produced attached to the disks. The
produced BC presented a crystallinity of 66.9% with
lower Youngs modulus (372.5 MPa) and strength
(15.9 MPa) compared to static BC (crystallinity of
88.7%, Young’s modulus of 3,955.6 MPa and strength
of 99.2 MPa).
Regarding culture conditions, these depend on the
bacterial strain in addition to the oxygen supply, pH,
temperature and the substrate used (carbon sources,
which generally are saccharides). The fermentation
process is normally carried out at around 28–30 °C
and pH 4–7. Most of the studies have reported that pH
below 4 is not convenient for bacterial growth (Zahan
et al. 2015), however, Castro et al. 2012 isolated a new
strain, Gluconacetobacter medellensis (G. medelli-
nensis), with the ability to produce high quantities of
BC with excellent properties at very acidic pH, 3.5.
Later, Urbina et al. (2017) reported the cell viability of
G. medellinensis measured by plating of serial dilu-
tions (from 10–1 to10-6) on -agar medium adjusted at
pH 4.0 or at pH 7.0. It was observed that cell viability
was favoured by acidic pH. This fact could be an
advantage in large production fermentation processes
since most microorganisms are not viable in such low
pH media so contamination by other bacteria, fungus
or yeasts could be avoided.
Until now, the most used culture medium for BC
production is the one developed by Hestrin and
Schramm, known as H–S medium or standard
medium, composed by D-glucose, peptone, yeast
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extract, sodium phosphate dibasic, and citric acid
(Hestrin and Schramm 1954). Following this compo-
sition, some modifications have been investigated that
vary the carbon source i.e. fructose, sucrose, glycerol,
mannitol, etc. (Mohammadkazemi et al. 2015). How-
ever, the use of commercial nutrient sources is not
economically viable since they result in high produc-
tion costs, which limit the production at the industrial
scale and thus applications of BC. For this reason, in
recent years the studies have been focused on the
development of cost-effective carbon feedstocks such
as by-products and residues from different industrial
sectors. In this way, some of the most interesting
research works involving agro-waste valorization as
feedstock for BC production are discussed.
Agro-wastes for BC production
The use of industrial waste for BC production is an
economically beneficial approach because it implies
the reduction of the costs associated first to the raw
materials in large scale industrial applications, and
second, to the disposal and management of these
residues which usually are associated with pollution
and accumulation problems. This means that the
employment of such feedstock leads to an environ-
mentally friendly process to obtain a value added
product. Agricultural wastes from agro-based indus-
tries, which can be in the form of liquids, slurries, or
solids, represent an important source of pollution.
Agro-wastes include residues from farms, crops,
horticulture and dairy products, such as straw, stem,
stalk, leaves, husk, shell, peel, pulp, stubble, etc.,
which come from cereals (rice, wheat, corn, sorghum,
barley), cotton, groundnut, jute, legumes (tomato,
bean, soy), coffee, cacao, tea, fruits (apple, grapes,
banana, mango, coco) and palm oil (Ezejiofor 2014;
Sadh et al. 2018). The composition of these wastes
depends on the system and type of agricultural
activities: some of them are composed by lignocellu-
losic biomass rich in carbohydrates, protein and
nitrogen and others mainly by cellulosic compounds
with essential oils and fatty acids. These features make
them very promising for BC production. In fact, in past
years several studies have been focused on the
utilization of agro-wastes for BC production; some
are summarized in Table 1.
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Table 1 Utilisation of different agro-wastes for bacterial cellulose production

Agro-waste Pre-treatment Extra nutrient Cultivation Bacteria BC
production
(g L-1)

Rice bark (Goelzer Acid Glucose Static/dynamic Acetobacter xylinum 2.42/1.57

et al. 2009) hydrolysis ? enzymatic ATCC 23769
hydrolysis
Oat hulls (Skiba et al. Nitric acid Black tea Static Medusomyces 2.20
2020) hydrolysis ? NaOH extract gisevii Sa-12
treatment ? enzymatic
hydrolysis
Bagasse (Qi et al. 2017) Acid hydrolysis/enzymatic No Static Gluconacetobacter 1.09/0.42
hydrolisis xylinus CH001
Wheat straw (Chen Ionic liquid ? enzymatic Yeast extract Static Acetobacter xylinus 8.3
et al. 2013) hydrolysis Peptone ATCC 23770
Tobacco waste (Ye Boiling in water and steam No Static Acetobacter 5.2
et al. 2019) distillation for nicotine xylinum ATCC
removal 23767
Corn stalk (Cheng et al. Acetic acid Baco-peptone Static Acetobacter xylinum 2.86
2017) hydrolysis ? detoxification Yeast extract ATCC 23767
d-Mannitol
Magnesium
sulphate
Ethanol
Coffee husk (Rani and Boiling in water No Static Gluconacetobacter 5.6
Appaiah 2013) hansenii UAC09
Orange peel (Kuo et al. Homogenization with Yeast extract Static Gluconacetobacter 6.13 ± 0.22
2019) water ? filtration Peptone xylinus
Acetate buffer
Citrus peel and pomace Enzymatic hydrolysis Yeast extract Static Komagataeibacter 5.70 ± 0.70
(Fan et al. 2016) Ethanol xylinus CICC
No.10529
Peptone
Pecan nutshell Acid hydrolysis Yeast extract Static Gluconacetobacter 2.81 ± 0.04
(Dórame-Miranda Peptone entanii
et al. 2019)
Ethanol
Salts
Pineapple peel (Algar Homogenization with Sugar cane Static/dynamic Gluconacetobacter 3.97 ± 0.36/
et al. 2014) water ? filtration medellinensis 0.82 ± 0.26
Rotten banana/rotten Homogenization with water No Static Komagataeibacter 4.81/1.95
mango (Molina- medellinensis
Ramı́rez et al. 2018)
Apple pomace (Urbina Homogenization with Sugar cane Static Gluconacetobacter 2.5 ± 0.3
et al. 2017) water ? filtration medellinensis
Pineapple peel/ Homogenization with Sucrose Static Komagataeibacter *125/*100
watermelon peel water ? sieving Ammonium hansenii MCM
(Kumbhar et al. 2015) sulphate B-967
Cycloheximide
Ripe dates (Lotfiman Ultrasonication with water Yeast extract Static Acetobacter xylinum 5.8
et al. 2018) Peptone 0416
Na2HPO4
Citric acid

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Cellulose (2021) 28:8229–8253 8237

Table 1 continued
Agro-waste Pre-treatment Extra nutrient Cultivation Bacteria BC
production
(g L-1)

Grape pomace (Cerrutti Homogenization with Corn steep Static Acetobacter xylinum 6.56
et al. 2016) water ? filtration liquor NRRL
B-42
Potato peel (Abdelraof Nitric acid hydrolysis No Static Gluconacetobacter 2.61
et al. 2019) xylinum ATCC
10245
Candied jujube- Acid hydrolysis No Static Gluconacetobacter 2.25
processing waste xylinum CGMCC
water (Li et al. 2015) No.2955
Soybean oil refinery No Corn steep Static Gluconacetobacter 2.20
effluent (Qiao et al. powder xylinus
2019)
Sugarcane molasse No No Static Komagataeibacter 1.90
(Machado et al. 2018) rhaeticus
Cheese or curd whey No No Dynamic Gluconacetobacter 5.45 ± 0.09
(Revin et al. 2018) sucrofermentans B-
11267
Cotton waste textiles Ionic liquid ? enzymatic Peptone Static Acetobacter xylinus 10.8
(Hong et al. 2012) hydrolysis Yeast extract ATCC 23770
*
Referred to wet weight

BC production from crop residue
Agricultural crop wastes are usually produced in the
form of lignocellulosic solids such as bark, hulls,
bagasse, stalk, straw, stems and leaves. As a starting
point, lignocellulosic materials can be processed and
given some pre-treatments to obtain carbohydrate rich
preparations for microbial growth. Glucose is usually
the most preferred sugar for bacteria, as it has been
observed to be the most consumed sugar, although
fructose and sucrose can be also metabolized simul-
taneously, but to a lesser extent (Urbina et al. 2017;
Kuo et al. 2019). The most commonly method used is
the chemical hydrolysis, which consists on the dilution
of the raw materials in acid or alkali aqueous media to
solubilize and break down polysaccharides- cellulose
and hemicelluloses- obtaining hydrolysates rich in
fermentable sugars (Vojvodić et al. 2016). Addition-
ally, enzymatic hydrolysis or enzymolysis can be
found, which is usually performed with cellulase
enzymes. This group of enzymes is produced by fungi,
bacteria, protozoans, plants and animals and catalyze
the decomposition of cellulose (Jayasekara and Rat-
nayake 2019). In some cases both pre-treatments,
chemical and enzymatic, are combined (Hsu et al.
2010). Goelzer et al. (2009) processed rice bark by
both hydrolysis treatments to obtain 0.78 g 100 g-1 of
reducing sugars for bacterial cellulose production,
although they supplemented the medium with glucose
to obtain a BC static production of 2.42 g L-1. Oat, a
cereal grain, is another crop which generates residues
i.e. oat hulls, and it has been used for BC production.
Skiba et al. (2020) conducted oat hulls to successive
pre-treatments to produce BC under pilot-plant con-
ditions. Firstly, oat hulls were subjected to nitric acid
hydrolysis, and then to a NaOH treatment in order to
eliminate lignin, which is an inhibitor for BC produc-
ing bacteria. Finally, enzymatic hydrolysis was per-
formed and 2.20 g L-1 of BC were obtained.
Although acid hydrolysis has been widely used for
this purpose, this method implies high energy con-
sumption due to the high temperatures and erosion of

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equipments, so, in order to invest towards green
chemistry processes, other pre-treatments have been
employed to treat agro-wastes for microbial feed-
stocks. In this way, ionic liquids followed by enzy-
molysis have been used to treat wheat straw (Chen
et al. 2013). A high BC production of 8.3 g L-1 was
obtained with the addition of extra nutrients rich in
nitrogen, i.e., yeast extracts and peptone. Tobacco is
another crop cultivated worldwide that produces
wastes in the form of leaves, stems and scraps. Ye
et al. (2019) observed that these wastes contain sugars
which can be used to produce BC. Firstly, tobacco
waste extract was pre-treated with boiling water. In
this case, it was observed that nicotine was an inhibitor
of BC production, so steam distillation was used for its
removal. In that study, changes of pH during fermen-
tation were the determining factor for BC production
and not the addition of extra nutrients. At the middle of
the fermentation a drop of the initial pH was observed
from 6.50 to 2.65, so a two-stage fermentation process
was followed by the re-adjusting of the pH at 7 days of
fermentation, which led to a BC production of 5.2 g
L-1. As it can be observed, when crop wastes are used
for BC production, pre-treatment methods are very
important. In the case of corn stalk it was observed that
inhibitors contained in prehydrolysate (furfural and
lignin) were adverse to the production of bacterial
cellulose, so these were removed through detoxifica-
tion treatments (Cheng et al. 2017). Corn stalk was
firstly pre-treated by acetic acid and subsequently
detoxified by the combination of activated carbon
and ion exchange resin treatment to remove micro-
bial growth inhibitors in order to obtain a BC
production of 2.86 g L-1.
BC production from industrial processing
Other types of agro-waste widely used for BC
production are derived mainly from industrial food
processing. In this group, fresh and processed fruits
and vegetables and fluid dairy products or brewery
products can be included, and the residues are
generated in the form of wastewater, slurry or solids
(husk, peels, pomace). In some cases, simple pre-
treatments have been used such us boiling water to
obtain slurries, for example when using coffee cherry
husks for BC static production (Rani and Appaiah
2013). With total sugar of 7.2% a BC production of
5.6 g L-1 was obtained without the addition of other
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Cellulose (2021) 28:8229–8253
nutrients. However, with the addition of extra sources
like urea, corn steep liquor, ethanol, or acetic acid,
higher BC production was observed (up to 8.2 g L-1).
The use of fruit waste has resulted in a very promising
option for BC production due to the high sugar
contents and simple pre-treatments, usually homoge-
nization with water and filtration of solids, which
could be an advantage in industrial applications due to
the reduction of the costs. In counterpart, most fruits
are naturally acidic, such as apples, citrus fruits,
pineapples, strawberries, etc., so a pH control is
necessary for their use in bacterial fermentation. Kuo
et al. (2019) compared BC production in media
prepared with orange peel hydrolyzate and with no
pre-treatment (orange peel fluid). A higher BC
production was observed in the non-hydrolyzed
medium (6.13 g L-1) than in the hydrolyzed one
(4.11 g L-1). In addition, the determining factors for
higher BC production were pH effect (controlled by
acetate buffer) and addition of nitrogen sources to the
orange peel fluid. However, as commented previously,
bacterial strains capable to produce BC at very acidic
pH can be found. This is the case of the work
developed by Algar et al. (2014) and Urbina et al.
(2017) where pineapple peel at pH 3.5 and apple
pomace from cider production at pH 4.0 were used,
respectively, for BC production using Gluconaceto-
bacter medellinensis. Interestingly, in the last project,
a microbiological study was performed. The relative
quantification of the expression of the genes involved
in the production of BC revealed a higher expression
in the case of the medium prepared with apple pomace
than in the commercial H–S medium, which also could
explain the higher BC production (2.5 g L-1 vs
1.8 g L-1, respectively). Apart from fruits, veg-
etable wastes have also been suitable for BC produc-
tion such as peels of potatos, one of the most
consumed vegetables worldwide (Abdelraof et al.
2019). Moreover, by-products of dairy industry, i.e.
cheese and milk whey, have also been analyzed as
source of nutrients for BC production. Both, without
additives (Revin et al. 2018) or in combination with
other wastes such as rotten fruit (Jozala et al. 2015),
led to promising BC production. In addition to food
processing wastes, there are research on wastes from
the textile industry that have been used for BC
production, such as Hong et al. 2012, which used
cotton-based waste textiles. Old undyed 100% cotton
T-shirts were treated with an ionic liquid and then
Cellulose (2021) 28:8229–8253
hydrolyzed enzymatically. A high BC production of
10.8 g L-1 was obtained with extra nutrients (yeast
extracts and peptone).
In view of the research involving agro-wastes for
BC production it can be concluded that each raw
material requires a processing method and the process
conditions must be adjusted depending on the bacterial
strain to reach an optimum production of BC. In some
cases, the determining factor is the pH change, in
others the lack of some nutrients such as nitrogen, and
even the fermentation time due to the generation of
some inhibitors. Nevertheless, the results reported in
recent years reveal the feasibility of using these wastes
for BC biosynthesis to obtain materials with interest-
ing properties and suggest a very promising produc-
tion of bacterium-derived polymers at industrial scale.
Properties of BC produced from agro-waste
Generally, research work in which non-conventional
carbon sources are used for BC production are mainly
focused on aspects related to production performance:
the main goal is to improve the production perfor-
mance of the process compared to commercial carbon
sources. In addition, the properties of the obtained BC
are also important to analyze, since these will later
condition its applications. BC properties and confor-
mation, i.e. water holding capacity or swelling capac-
ity (WHC), porosity, degree of polymerization (DP),
crystallinity index (CI), mean crystallite size, average
fiber diameter (D) or mechanical performance
(Young’s modulus (E) and tensile strength (r)),
among others, are affected by different factors
including the carbon source used for BC-growing
bacteria. Singhsa et al. (2018) analyzed the morphol-
ogy, crystallinity and crystallite size of the BC
produced by five strains of Komagataeibacter xylinus
in static and agitated fermentation conditions using six
different carbon sources: glucose, fructose, lactose,
maltitol, sucralose, and xylitol. They observed that the
variation of the carbon sources did not impact the
morphology of BC produced by each strain and the
crystallite size and Ia allomorph content data did not
differ greatly between carbon sources among each
bacterial strain. However, the crystallinity of the
cellulose produced in the different carbon sources was
apparently different. As can be observed in Table 2,
which gathers some of the properties analyzed in some
research work using agro-wastes for BC production, is
8239
strongly affected by the carbon source. However, apart
from crystallinity, changes in morphology, arrange-
ment, and length of the ribbons have been observed
when using agro-wastes as feedstock and this has been
attributed to the presence of different polysaccharides.
Kumbhar et al. (2015) used pineapple and watermelon
peels for BC production, and compared the morphol-
ogy of the obtained BC with the one grown in H–S
medium. They observed that the ribbons of the BC
obtained in H–S medium were longer and more
uniform and concluded that changing medium com-
ponents (mixture of sugars) led to different assembling
patterns. This was also observed by Urbina et al.
(2017) that used apple residues from the cider
production mixed with sugarcane for BC biosynthesis,
so that the culture media contained a mixture of sugars
(glucose, fructose and sucrose). They observed that the
nanofibrils of BC from H–S medium were slightly
thicker (63 ± 6 nm) than those from the apple waste
medium (55 ± 6 nm). In the literature, a great variety
of fiber average diameter data can be found when
using agro-wastes for BC production. For example,
when candied jujube processing wastewater (Li et al.
2015) was used for BC production, very small
diameters (5.9 nm) were observed. In contrast, in the
case of using pecan nutshell (Dórame-Miranda et al.
2019) the diameter was 165 ± 43.03 nm. In the last
case, it is worth mentioning that the nanofiber
diameter before purification was smaller
(120.29 ± 30.01 nm) than after treatment with aque-
ous sodium hydroxide (165 ± 43.03 nm). This result
indicated that the process of mercerization increased
the nanofiber volume. This fact is something that
should be taken into account, since although the
commercial dissemination of BC obtained from agro-
wastes is not exhaustively exploited yet, BC must be
purified to make it suitable for its potential
applications.
BC application and potential applications of BC
from agro-waste
As described above, BC is a biopolymer that presents
unique conformation, excellent physical and mechan-
ical properties and offers the possibility to be modified
using different methods to produce a new generation
of materials and nanocomposites with improved
characteristics. In fact, it has proven to be suitable for
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8240 Cellulose (2021) 28:8229–8253

Table 2 Properties of BC produced from agro-wastes

Agro-waste BC properties
WHC CIXRD (%) D (nm) E (MPa) r (MPa)

Pecan nutshell (Dórame-Miranda et al. 90.1 165.5 ± 43.03

2019)
Cotton waste textiles (Hong et al. 2012) 98.8 ± 0.3 (%) 0.070 ± 0.01
Pineapple peel (Algar et al. 2014) * 1400 (%) 84
Rotten banana/rotten mango/cheese 9.4 ± 1.2/ 280.6 ± 31.3/
whey (Molina-Ramı́rez et al. 2018) 1.6 ± 0.4/ 28.1 ± 6.4/
0.8 ± 0.2 17.7 ± 1.8
Apple pomace (Urbina et al. 2017) 8194 ± 1226 (%) 89 55 ± 6 10,410 ± 1190 89 ± 30
Pineapple peel/watermelon peel 119.9 ± 16.18 61.4/56
(Kumbhar et al. 2015) (%)/
170.64 ± 4.16
(%)
Ripe dates (Lotfiman et al. 2018) 142 (%) 74–76 90–105
Grape pomace (Cerrutti et al. 2016) 68–85 32.3–29.1
Citrus peel and pomace (Fan et al. 63 50
2016)
Cucumber/melon/kiwifruit/tomato/ 600–900 (%) 80.27–92.96 47.64–61.11
quince/apple peels (Güzel and Akp
2020)
Candied jujube-processing waste water 64.9–80.9 5.9
(Li et al. 2015)
Sugarcane molasse (Machado et al. * 160 (g/g) 86 119 600
2018)
Soybean oil refinery effluent (Qiao 111 ± 9 (g/g) 64
et al. 2019)
Cheese or curd whey (Revin et al. 50.2 100–180
2018)

several applications in different fields. Therefore, in
this section some of the newest and challenging
applications of BC in environmental applications,
optoelectronic devices, food industry, biomedical field
and 3D printing technology are discussed (Fig. 3).
It is worth mentioning that to date, most of the studies
are carried out from cellulose obtained in standard
media, because the applications of bacterial cellulose
obtained from agro-wastes are usually limited. This
fact is not because the properties of the BC obtained in
alternative cultures have not proven to be similar or
better than the ones obtained with commercial
cultures, but because the cellulose would have to
undergo purification processes and a proper standard-
ization is needed. However, some research work has
proven the suitability of BC obtained from agro-waste
for several applications, so these will be mentioned at
the end of this section.
Environmental applications
Nowadays, due to the new environmental regulations,
there is a growing tendency to search for green
materials and bio-based products to invest towards
sustainability, industrial ecology and green processes.
Bacterial cellulose, as a biopolymer produced in
fermentation processes with bacteria, meets the
requirements to produce a new generation of
biodegradable and high specific materials for envi-
ronmental applications. Due to its interesting confor-
mation and strong 3D nanofibrillar network BC
provides a promising alternative for wastewater
purification and reuse technologies. Until now, it has

123
Cellulose (2021) 28:8229–8253 8241

Environmental applications

Heavy metals removal

Catalysis of organic pollutants
Absorption of organic solvents
BC membrane Oil/water separation

Optoelectronic and conductive devices

Dielectric materials
Thin film transistors
Solar cells
Organic light-emitting diodes
Food ingredients and packaging

Stabilizing agent
Thickening agent
Multilayers
Nanoreinforcements
Biomedical applications

Wound-healing systems
Scaffolds
Blood vessels
Drug delivery
3D printing

Bioinks
Specific geometries

Fig. 3 Recent applications of BC membranes in different fields discussed in this section

been widely used for supporting different nanoparti-
cles, biopolymers and additives for applications in the
membrane technology such as heavy metals removal,
catalysis of organic pollutants, absorption of organic
solvents, oil/water separation processes.
The presence of heavy metals in environment is
associated with industrial activities in metal-mechan-
ics industries such as batteries, tanneries or fertilizers.
One of the problems associated with the appearance of
heavy metals is the potential for accumulation causing
more exposure of these metals to an organism than
could be found alone in the environment, so the
elimination of these contaminants form wastewaters is
an important issue. For these applications, BC mem-
branes are used as templates, usually combined with
chelating agents or absorbents, such as chitosan,
ethylenediaminetetraacetic acid (EDTA), magnetic
nanoparticles, graphene oxide or subjected to chem-
ical modifications such as oxidation with 2,2,6,6-
tetramethylpy-peridine-1-oxy radical (TEMPO-oxi-
dation). BC has proven to be an efficient template
for the elimination of copper (Cu), strontium (Sr),
antimony (Sb), lead (Pb), iron (Fe), chromium (Cr)
and arsenic (As), among others, reaching efficiencies
up to 90% (Stoica-Guzun et al. 2016; Hassan et al.
2019; Mensah et al. 2019; Meng et al. 2019; Cheng
et al. 2019). Organic dyes are another group of water
pollutants that are difficult to treat by traditional or
biochemical methods. A green chemistry approach
that is currently widely used is photocatalysis, which
involves nanostructured semiconductors and light to
trigger the corresponding chemical reactions. In this
context, BC/polydopamine/TiO2 nanoparticle
nanocomposites for photocatalytic degradation of
dyes have been produced (Yang et al. 2020a). The
use of BC structure modified with polydopamine
greatly improved surface area and active sites for the
adsorption. The nanocomposites resulted to have
excellent photo-degradation performance, stability
and reusability after five cyclic tests. 4-Nitrophenol
is a hazardous pollutant usually present in wastewater
which originates from dye, paper and pharmaceutical
industries. The typical strategy to eliminate and take
advantage of this contaminant is the reduction to
4-aminophenol, which is used for the synthesis of
peptides, pharmaceuticals and corrosion protection

123
8242
compounds (Ibrahim et al. 2019). Metal nanoparticles
are widely investigated for the catalysis of these
reactions and the employment of an adequate support
to avoid the aggregation and not impair their catalytic
performance is essential. Song et al. (2020) reported
that BC skeleton was a very convenient support for the
dispersion of metal nanoparticles (Cu or Ni) improv-
ing the interactions and mass transfer for the reduction
of 4-nitrophenol. An optimized catalyst (BC/Cu-0.5)
was developed with the ability to perform the reduc-
tion process in 8 min and even could be reused.
Moreover, oil spills and oily wastewaters generated
from the industrial activities are a major source of
contamination, so the use of green absorbents and
filters based on BC for such applications has increased.
BC membranes functionalized with silica derived
compounds to obtain BC/SiO2 network aerogels have
proven to have high separation efficiencies in oil/water
and water/oil emulsions and even reached oil recov-
eries up to 88% (He et al. 2018; Hou et al. 2019).
Additionally, the feasibility of the neat BC membrane
without modification as a filter for oil removal
(Galdino et al. 2020) has also been proven. BC
membranes showed 100% of oil removal in emulsions
with concentrations of 10, 150 and 230 ppm. In the
context of water purification, apart from the membrane
shape, BC has also been used as a carbon precursor for
the preparation of materials for the desalination of
water via the capacitive method. Belaustegui et al.
(2020) prepared graphene enriched BC-based elec-
trode materials by carbonisation, which exhibited
unprecedented desalination capacities for nanocar-
bons due to the hierarchical porosity, density, lattice
defects and high content of surface oxygenated
species.
Optoelectronic and electronic devices
The field of electronics is another area in which
cellulose, and thus BC, is gaining special attention.
The tendency is to invest towards new substrates with
proper flexibility, transparency, thermal stability in
addition to sustainability and recyclability to develop
electronic devices. These features can be obtained
with cellulose based nanocomposites. The excellent
mechanical performance, thermal stability and surface
morphology of BC are characteristics which make it
interesting for applications in electronic and optoelec-
tronic devices, which include dielectric materials for
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Cellulose (2021) 28:8229–8253
transistors, capacitors and supercapacitors, solar cells
and organic light-emitting diodes (OLEDs) (Fortunato
et al. 2016; Vicente et al. 2017).
A dielectric material is an insulator material whose
interface is in contact with conductor/semiconductor
materials. As it has been reported, the addition or
utilization of BC improves the insulator properties of
traditional paper, as neat BC possess an intrinsic
insulating behaviour which is very convenient in such
applications (Zhuravleva et al. 2018, 2019). More
specifically, BC has also been studied as a fractional
order material for the realization of all-polymeric BC-
based fractional order electronic devices for circuits.
Caponetto et al. (2019) reported for the first time a
study of the fractional order nature of BC as a base
material in a real Fractional-Order Element (FOE)
device. These are used for the realization or approx-
imation of fractional differential and integral equa-
tions which are used to model the behaviour of
electrical current to the voltage in circuits composed
by capacitors and inductors. Another recent approach
for the use of BC in electronics has been the
development of triboelectric nanogenerators (TENG).
These are based on tribo-electrification and electro-
static effects induced by contact (like friction) which
can harvest mechanical energy from the environment
and biological systems. Shao et al. 2019 combined the
surface structure of BC film with high dielectric
particles BaTiO3 to improve dielectric constant and
nanostructure for the development of an environmen-
tally friendly TENG. This device showed excellent
stability and had the capacity to harvest mechanical
energy by human motion such as movements from
arm/finger of human body to generate micro-current.
On the other hand, the insulator/conductive properties
of BC can be tailored or tuned by the addition and
modification of the BC structure with conductive
additives opening a wide range of applications in
semiconductor devices. Hosseini et al. (2019) fabri-
cated a highly flexible ternary system with BC as a
template for the incorporation of Ag nanoparticles and
polyaniline, and reported a high energy density as an
electrode for a supercapacitor. Currently, for the
production of BC-based materials with high electrical
conductivity, carbonaceous materials such as fullere-
nes, carbon nanotubes and graphene are gaining
special attention. Graphene is a carbon material with
large surface area, good mechanical strength, inher-
ently high conductivity and charge transport
Cellulose (2021) 28:8229–8253
behaviour and in combination with the 3D structure of
BC interesting nanocomposites can be obtained for
multifunctional electronics (Guan et al. 2019; Sheng
et al. 2019). As commented above, BC has also been
studied in the technology of solar thermal energy and
storage. MXene materials are regarded a new family
of multifunctional 2D materials with many attractive
and tunable properties (with the chemical formula
Mn?1XnTx, where M stands for an early transition
metal, X represents carbon and/or nitrogen and Tx
refers to surface functional groups). Tang et al. (2019)
combined the BC structure with MXene as the
photothermal material and polyethylene glycol to
produce a nanocomposite with great shape stability,
photothermal conversion ability and high energy
storage capacity. Moreover, BC also presents inter-
esting features for the production of OLEDs. These
devices should be flexible and transparent as they have
applications in televisions, monitors or portable device
screens. BC nanofibers present diameters less than
one-tenth of the visible light wavelength, being a free
light scattering medium, so the ultrafine network of
BC has been used to produce transparent paper
materials. For this purpose, BC has been combined
with other materials such as poly(2-methoxy-5(2-
ethylhexyloxy)-1,4-phenylenevinylene)-MEH:PPV (a
semiconductor and luminescent polymer) or with
organic–inorganic sol–gel materials, composed of
boehmite nanoparticles and epoxi modified siloxane
(Aleshin et al. 2017; Legnani et al. 2019).
Food packaging and ingredients
The use of BC as raw material has a long history in
Asian countries such as Philippines, where a tradi-
tional dessert known as Nata is produced, called ‘‘Nata
de coco’’ or ‘‘Nata de pina’’ depending on the flavour
(Shi et al. 2014). Wet BC membranes present a smooth
feel and gelatinous consistency which makes it
interesting for food applications. It has been reported
that the gelatinous membrane of BC became edible
after processing it with sugar alcohol or with alginate
and calcium chloride, and that it presented the
consistency of fruit or molluscs (Okiyama et al.
1992). It is worth noting that BC can be considered a
type of dietary fiber, which was classified as safe by
the USA Food and Drug Administration in 1992, so its
use as raw material, food ingredient or additive is
commonly accepted (Ullah et al. 2016). Until now, BC
8243
has been used in food-related applications as a
thickening, gelling and water-binding agent and it
has proven to improve the rheology of food mixtures
(Paximada et al. 2016a; b). Concurrently, it has proven
to be a very convenient stabilizing additive in
Pickering emulsions. They are emulsions stabilized
by particles that are absorbed at the interface, forming
a layer around the surface of the dispersed phase
droplets, and prevent coagulation and coalescence
phenomena (Zhai et al. 2018; Martins et al. 2020).
From the dietetic point of view, there are several
studies that prove the safety and beneficial effects of
bacterial cellulose including fat substitution or choles-
terol lowering (Dourado et al. 2017). Guo et al. 2018
used BC to produce reduced-fat ice cream and they
observed improvements in the stability, emulsifying
and rheological properties and higher resistance to
melting with sensory attributes similar or superior to
the control ice creams. Akoğlu et al. 2018 investigated
the effect of the addition of BC to mayonnaise as fat
replacer on the rheological behaviour and sensory
characteristics with satisfactory results.
In addition, due to its strong network and barrier
properties BC has been evaluated as a packaging
material in many research sstudies. In the field of food
packaging, paper-based materials are regarded as a
suitable option as paper is completely renewable and
recyclable. Moreover, in the production of packages
and bags, paper is an important material because it can
be considered a suitable alternative to replace syn-
thetic plastics. The application of nanotechnology in
the papermaking science offers the possibility to
produce paper with improved properties and perfor-
mance in terms of hydrophobicity, strength, barrier
properties and processing. In fact, nanopapers fabri-
cated from cellulose nanofibers show interesting
features such as high strength, thermal stability,
optical transparency and good oxygen barrier proper-
ties due to the strongly reduced penetration of oxygen
molecules through the highly entangled fibrillar
structure. These enhanced properties would not be
achieved using the traditional microsized pulp papers
(Samyn et al. 2018). Indeed, BC can be considered a
nanopaper itself which presents high mechanical
performance due to its 3D network-like structure
formed by highly crystalline cellulose nanofibers. Due
to its interesting properties, the use of bacterial
cellulose in packaging related applications has also
been investigated. In fact, BC has been previously
123
8244
used to reinforce papers and cellulosic surfaces
(Santos et al. 2017; Campano et al. 2018) and bacterial
cellulose nanocrystals (BCNCs) have been used as
reinforcing fillers for hybrid nanocomposites with
potential applications in eco-friendly food packaging
(Urbina et al. 2019b). Moreover, the application of BC
in ‘‘active packaging’’ has been extensively investi-
gated in order to design packaging materials with
components that release or absorb substances from or
into the packaged food or the surrounding environ-
ment in order to extend the shelf-life and maintain or
improve the condition of packaged food. Shafipour
Yordshahi et al. (2020) developed an antimicrobial
ground meat wrapping nanopaper by the impregnation
of bacterial cellulose with lyophilized postbiotics of
Lactobacillus plantarum. The as prepared films were
optimized and their antimicrobial activity against
Listeria monocytogenes was proved. Xie et al. (2020b)
observed an improvement of the tensile strength and
reduced water vapor and oxygen permeability and
moisture content with the addition of BC to potato peel
based films for food packaging applications. In
addition, curcumin (a polyphenolic compound iso-
lated from the rhizomes of the plant Curcuma longa)
was added to provide antioxidant properties to the
films. Apart from antioxidant properties, curcumin has
resulted to be a suitable additive to detect pH changes
and boric acid traces, factors than can affect food
quality and human health. Yang et al., (2020a, b)
produced in situ hybrid BC and chitin nanofiber films
with curcumin nanoparticles for the development of
smart packages. These films exhibited antioxidant and
antimicrobial properties and also changed their colour
with variations on pH and in presence of boric acid.
Biomedical applications
Nowadays, one of the most attractive applications of
BC is in the biomedical field. The first and most
important aspect in this type of applications is to prove
the biocompatibility of the materials for the cells to
maintain their tissue-specific functions and support. In
this way, the biocompatibility of this biopolymer has
been demonstrated in several works, both in vitro and
in vivo. de Lima et al. (2017) tested the biocompat-
ibility of BC membranes for dural defect on rats during
120 days and they observed good mechanical stabil-
ity, similar properties to local tissues and absence of
inflammatory or neurotoxicity reactions. In fact, BC
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Cellulose (2021) 28:8229–8253
nanofibers are microscopically similar to collagen
fibers and collagen is the main component of human
connective tissue, so this feature makes it attractive for
applications in tissue engineering (Torgbo and Sukyai
2018). Until now it has been used for wound healing
(Pourali et al. 2018; Sajjad et al. 2019), artificial skin
(Keskin et al. 2017), scaffold materials (Li et al. 2016;
Kumbhar et al. 2017), dental implants (Voicu et al.
2017) or artificial blood vessels (Lee and Park 2017).
For this last application, it is interesting to comment on
the work developed by some researchers where
bacterial cellulose gels were produced in tubular form
(Fink et al. 2007; Putra et al. 2008; Zang et al. 2015).
With the employment of the correct material mold and
oxygen transfer, BC tubes can be obtained with
desired length, inner diameter, and thickness. In
addition, antimicrobial activity is an important prop-
erty in such applications, but BC itself does not present
this feature, so the modification of BC membranes to
induce antimicrobial activity is a common trend. For
this, biopolymers such as chitosan, metal oxide
structures or anti-inflammatory drugs have been used
for grafting/anchoring into BC membranes to provide
antimicrobial activity (Sun et al. 2019; Chantereau
et al. 2019; Xie et al. 2020a; Ju et al. 2020). Another
blooming application of BC is the development of
drug delivery systems. In these applications hydrogels
are gaining special attention. Hydrogels are charac-
terized by a 3D network which has the capacity to
swell and retain a large fraction of water within the
structure. Due to its 3D nanofibrillar network structure
and high water holding or swelling capacity, BC can
be considered an hydrogel itself, so it has been used to
produce systems capable to get the right amount of the
active agent such as anti-inflammatories or antiseptics,
and then release them in the right place and with the
correct dose in the organism (Luz et al. 2018; Weyell
et al. 2019; Beekmann et al. 2020; Inoue et al. 2020).
Finally, BC has also proven to be a suitable material
for the development of shape memory materials in
biomedical applications. These materials have the
ability to recover their original shape after being
deformed in a temporary shape under the application
of external stimulus, so this was the approach followed
to develop biomedical devices based on BC and
polyurethane triggered by human body liquids in order
to minimize invasive surgeries (Urbina et al. 2019a).
Cellulose (2021) 28:8229–8253 8245

3D printing technology

Traditionally, BC membranes have been mainly

formed in a non-immobilized state, in fluid culture
media containing bacteria deposited on the desired
substrates, surfaces or recipients. One of the most
challenging aspects of BC is to obtain a matrix with
controlled 3D geometry and complex shape. In this
sense, the research in 3D-printing technology, called
‘‘additive manufacturing’’, is growing rapidly because
it offers the possibility to produce complex structures
and personalized products with multiple functions. In
this field, ‘‘bioprinting’’ is an emerging approach for
the fabrication of complex structures containing living
cells. For this purpose, the investigations are focused
on the development of living inks or ‘‘bioinks’’ with
the physical and rheological features for the printing
process, but also the capability to immobilize bacteria
while maintaining their metabolic activity and satis-
fying their biological needs. The idea is to harvest BC
producing bacteria in an appropriate viscoelastic
matrix such as hydrogels, in a one step approach,
while the BC is produced free-formed into intricate
geometry. Schaffner et al. (2017) developed a
‘‘bioink’’ based on hyaluronic acid, k-carrageenan
and fumed silica for the immobilization of Gluconace-
tobacter xylinus which they called called ‘‘Flink’’, to
obtain BC geometries for biomedical applications.
The composition of the ‘‘bioink’’ was optimized to
keep adequate rheological properties and extrudability
with no alteration by the presence of bacteria. In this
research a doll face was scanned and the ‘‘bioink’’ was
3D printed in that complex shape to produce a ‘‘face
shape BC’’ in view of developing a skin transplant
(Fig. 4A). Moreover, in order to enhance the oxygen Fig. 4 A A doll face was scanned, and a 4.5 wt % Flink
containing A. xylinum was deposited onto the face using a
supply for the embedded bacteria and improve the
custom-built 3D printer. In situ cellulose growth led to the
dimensional stability of the ‘‘bioinks’’, solid matrix- formation of a cellulose-reinforced hydrogel that, after removal
assisted 3D printing (SMAP) has been reported. In this of all biological residues, can serve as a skin transplant
case, matrices such as viscous liquids or hydrophobic (Schaffner et al. 2017). B Bacteria containing ink were printed
inside of solid matrix. Oxygen was supplied through the solid
solid particles are used as a support for extruded
particles to the surface of the cellulose nanofiber ink containing
hydrogel or ‘‘bioink’’ printing structures, providing a Gluconacetobacter xylinus, allowing the bacteria to metabolize
gap space between the particles through which the (Shin et al. 2019). C Coil, connected ring, tetrahedron, stacked
oxygen required for biosynthesis of cellulose can be lattice, and sandglass structure of CNF/BC after incubation as
printed according to the CAD designs (Shin et al. 2019). D Left:
delivered. Shin et al. (2019) reported that the use of an
Silicone mold used to guide the bioprinter during the bacterial
appropriate matrix allows creating complex BC culture to reproduce the large-scale features of the outer ear and
structures such as spheres, hollow tubes, coils and right: 3D BNC implant prototype produced in the shape of the
connected rings (Fig. 4B, C). In that work, the whole outer ear (Nimeskern et al. 2013)
material for the solid matrix was a spherical polyte-
trafluoroethylene (PTFE) microparticle due to its

123
8246
hydrophobicity and bioinertness, and the developed
‘‘bioink’’ for the growth of Gluconacetobacter xylinus
was composed by mannitol, peptone, yeast extract and
different cellulose nanofiber contents as rheology
modifiers. Interestingly, Nimeskern et al. (2013)
developed a bioprinter which dispensed controlled
volumes of bacterial culture media, and a precision
motion system for controlling the dispensing location.
With this system, a specific ear-shaped bacterial
nanocellulose implant prototype was biofabricated
using Gluconacetobacter xylinus strain. For this, a
silicone mold was used to inoculate the bacteria
suspension, the dispersing valve was programmed to
spray 500 lL of sterile culture media into the mold
every 6 h and the motion system was programmed to
move along the helix of the mold during dispensing.
After 18 days a BC ear-shape implant was obtained
(Fig. 4D).
Potential applications of BC from agro-wastes
It has been reported that BC obtained by some agro-
wastes can be colored and may absorb undesirable
substances, so a correct purification is mandatory.
These facts can be the reason to limit the use of BC in
areas where regulatory requirements are highly
restrictive, i.e. biomedicine, pharmaceutics, cosmetics
or food industry (Velásquez-Riaño and Bojacá 2017).
However, in food packaging applications it has been
reported that for some products, colored films can be
beneficial for both the consumer attraction and
protection of the food against negative effects of light
(Menzel et al. 2019). In those cases, the color of BC
films would not be such an inconvenience. Until now,
there is some research work in which BC obtained
from agro-waste has been used for potential applica-
tions in food packaging (Urbina et al. 2016, 2019b, c).
Urbina et al. (2019c) produced stiff bacterial cellulose
BC nanopapers (obtained from apple residues) with a
polyhydroxyalkanoate (PHA)/apple extract coating
for potential food packaging. In other work, the
textural properties of BC obtained from agro-waste,
i.e. watermelon and pineapple peels, were measured to
set up appropriate quality standards for food ingredi-
ents, and the results confirmed the suitability of the
produced BC for applications in food industry (Kumb-
har et al. 2015). In addition, there is work in which the
different agro-wastes used resulted in BC with differ-
ent properties. Therefore, according to those
123
Cellulose (2021) 28:8229–8253
properties applications were predicted, i.e. high-
strength composites or food stabilizers (Molina-
Ramı́rez et al. 2018). On the other hand, BC obtained
from agro-waste has proven to be an excellent porous
support for the attachment of other polymers. BC-
based environmentally friendly membranes have been
developed by two different routes, in situ and ex situ,
with chitosan as a functional entity for the elimination
of copper (Cu) in wastewater (Urbina et al. 2018).
Dhar et al. (2019) developed BC (obtained from
sugarcane straw) based films with highly conductive
properties. BC nanofibers formed highly integrated
and entangled networks with reduced graphene oxide
that resulted in nanocomposites with high electrical
conductivity, with potential applications in flexible
electronic devices. Finally, even though in the
biomedical field the applications of BC obtained from
agro-waste are more limited, there is some research
work in which interesting results have been obtained.
High performance water-activated shape memory
nanocomposites based on BC membranes (obtained
from apple residues) and a waterborne polyurethane
have been developed (Urbina et al. 2019a). Cell
viability, proliferation and adhesion were analyzed
and biocompatibility was observed for both neat BC
and nanocomposites. The presence of BC not only
improved the shape fixity ability of the nanocompos-
ites, but it induced a much faster and almost complete
shape recovery, so the use of these nanocomposites in
biomedical devices was proposed. In Table 3 some of
the potential applications of BC obtained from agro-
waste can be found. From everything mentioned
above, it can be concluded that the applications of
BC obtained from agro-waste are still under develop-
ment and although in the future a proper standardiza-
tion of them will be required to put them in practice,
much progress is being made regarding this issue.
Conclusions and future prospects
Bacterial cellulose is a bacterium-produced biopoly-
mer that has been extensively studied in the last
decades due to its remarkable properties and confor-
mation. Currently, efforts are being made to produce
BC from cheaper, readily available, and renewable
media. The sustainable production of this biopolymer
using agro-wastes is a clearly beneficial approach not
only from the environmental point of view, since it
Cellulose (2021) 28:8229–8253 8247

Table 3 Potential applications of BC obtained from agro-wastes

Agro-waste Application of BC References

Apple pomace Antioxidant films for food packaging Urbina et al. (2019c, b)
Pineapple/watermelon peel Food ingredients Kumbhar et al. (2015)
Forest fruits Antimicrobial food packaging Mocanu et al. (2019)
Molasses Antimicrobial food packaging Salari et al. (2018)
Rotten banana/rotten mango/cheese whey Composites /food suspensions stabilizer Molina-Ramı́rez et al. (2018)
Apple pomace Membranes for water purification Urbina et al. (2018)
Sugarcane straw Electronic devices (conductive films) Dhar et al. (2019)
Apple pomace Shape memory nanocomposites for biomedicine Urbina et al. (2019a)
Coconut water Wound dressing Lin et al. (2013)
Mature coconut water Capsules for pharmaceutical industry Kamarudin et al. (2018)

leads to the disposal and management of these wastes
that can generate pollution problems, but also to boost
the industrial production of the polymer due to
reduction of the costs associated to the raw materials.
The pre-treatments used for each agro-waste can vary
depending on the source, but all have the purpose to
prepare carbohydrate-rich preparations for maximum
BC production. In addition, it has been proven that the
characteristics of cellulose obtained from these agri-
cultural wastes are usually very similar or even better
than those obtained from the standard media, therefore
potential applications are coming to light, although are
still in development. Finally, this review comments on
the potential of BC for the production of a new
generation of multifunctional, value-added materials
with advanced applications in five areas of great
importance. With these prospects, the production and
applications of BC seem to increase at industrial scale.
Acknowledgments The authors thank for the financial
support from the Spanish Ministry of Economy and
Competitiveness (MAT2020-PID2019-105090RB-I00) and to
the University of the Basque Country for the GIU18/216.
L. Urbina wishes to acknowledge the University of the Basque
Country (UPV/EHU) for the postdoctoral DOKBERRI grant.
Declarations
Conflict of interest The authors have no conflicts of interest
to declare that are relevant to the content of this article.
Ethical approval This article does not contain any studies
with human participants or animals performed by any of the
authors. In this experiment, we did not collect any samples of
humans and animals.
Informed consent Informed consent was obtained from all
individual participants included in the study.
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