The Open Biotechnology Journal




ISSN: 1874-0707 ― Volume 15, 2021
REVIEW ARTICLE

Bacterial Cellulose from Food to Biomedical Products



Supajit Sraphet1, Bagher Javadi2, *
1 Institute of Molecular Biosciences, Mahidol University, Nakhon Pathom, Thailand
2 Department of Sciences, Faculty of Science and Technology, Suan Sunandha Rajabhat University, Bangkok, Thailand

Abstract

Cellulose production of aerobic bacteria with its very unique physiochemical properties attracted many researchers. The biosynthetic of Bacterial Cellulose (BC) was produced by low-cost media recently. BC has been used as biomaterials and food ingredient these days. Moreover, the capacity of BC composite gives the numerous application opportunities in other fields. Bacterial Cellulose (BC) development is differentiated from suspension planktonic culture by their Extracellular Polymeric Substances (EPS), down-regulation of growth rate and up-down the expression of genes. The attachment of microorganisms is highly dependent on their cell membrane structures and growth medium. This is a very complicated phenomenon that optimal conditions defined the specific architecture. This architecture is made of microbial cells and EPS. Cell growth and cell communication mechanisms effect biofilm development and detachment. Understandings of development and architecture mechanisms and control strategies have a great impact on the management of BC formation with beneficial microorganisms. This mini-review paper presents the overview of outstanding findings from isolating and characterizing the diversity of bacteria to BC's future application, from food to biosensor products. The review would help future researchers in the sustainable production of BC, applications advantages and opportunities in food industry, biomaterial and biomedicine.

Keywords: Bacterial cellulose, Molecular characterization, Biomedicine, Food products, Cellulose production, Forms of bacterial cellulose.


Article Information


Identifiers and Pagination:

Year: 2020
Volume: 14
First Page: 124
Last Page: 133
Publisher Id: TOBIOTJ-14-124
DOI: 10.2174/1874070702014010124

Article History:

Received Date: 27/6/2020
Revision Received Date: 16/9/2020
Acceptance Date: 7/10/2020
Electronic publication date: 22/12/2020
Collection year: 2020

© 2020 Sraphet & Javadi.

open-access license: This is an open access article distributed under the terms of the Creative Commons Attribution 4.0 International Public License (CC-BY 4.0), a copy of which is available at: https://creativecommons.org/licenses/by/4.0/legalcode. This license permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


* Address correspondence to this author at the Department of Sciences, Faculty of Science and Technology, Suan Sunandha Rajabhat University, Bangkok, Thailand; Tel: +66-02-160-1143-45; Fax: +66-02-160-1146; E-mail: Javadi.ba@ssru.ac.th





1. INTRODUCTION

Microorganisms are usually defined with their planktonic and suspension cell growth characterizations. Attachment of microorganisms on the surface exhibited the specific phenotypes. These phenotypic distinctions are due to gene expressions and growth rate. Therefore, biofilms showed the unique surface attachment-detachment, community structure and finally micro-ecosystems. Bacterial cellulose (BC) is a natural nanomaterial of exopolysaccharide biofilm from many bacterial genera such as Acetobacter, Sarcina and Agrobacterium and Komagataeibacter (former Gluconacetobacter) [1Rajwade JM, Paknikar KM, Kumbhar JV. Applications of bacterial cellulose and its composites in biomedicine. Appl Microbiol Biotechnol 2015; 99(6): 2491-511.
[http://dx.doi.org/10.1007/s00253-015-6426-3] [PMID: 25666681]
]. Komagataeibacter can produce BC while cultivated in a medium supplied with carbon and nitrogen [2Ruka DR, Simon GP, Dean KM. Altering the growth conditions of Gluconacetobacter xylinus to maximize the yield of bacterial cellulose. Carbohydr Polym 2012; 89(2): 613-22.
[http://dx.doi.org/10.1016/j.carbpol.2012.03.059] [PMID: 24750766]
]. The priority of BC production by this genus is due to their higher yield and purity. However, the species and strains biofilm structure and mechanical properties are different [3Chen S-Q, Lopez-Sanchez P, Wang D, Mikkelsen D, Gidley MJ. Mechanical properties of bacterial cellulose synthesised by diverse strains of the genus Komagataeibacter. Food Hydrocoll 2018; 81: 87-95.
[http://dx.doi.org/10.1016/j.foodhyd.2018.02.031]
].

Chinese reported BC firstly during their ancient production of fermented beverage kombucha tea. They observed the co-colony of acidic acid bacteria and yeast embedded on the beverage surface [4Marsh AJ, O’Sullivan O, Hill C, Ross RP, Cotter PD. Sequence-based analysis of the bacterial and fungal compositions of multiple kombucha (tea fungus) samples. Food Microbiol 2014; 38: 171-8.
[http://dx.doi.org/10.1016/j.fm.2013.09.003] [PMID: 24290641]
]. Generally, the microbial biofilm and in particular, BC production are the self-defense strategy towards sustainable acquiring and securing nutrients supply in harsh environments [5Reiniati I, Hrymak AN, Margaritis A. Recent developments in the production and applications of bacterial cellulose fibers and nanocrystals. Crit Rev Biotechnol 2017; 37(4): 510-24.
[http://dx.doi.org/10.1080/07388551.2016.1189871] [PMID: 27248159]
]. Many multi-step regulated mechanisms with the help of enzymes and catalytic complexes rigorously control the cell's BC synthesis reactions. Although, 1,4-β-glucan chains formation and their assembly to cellulose inside and outside of the bacterial cell are the main two steps of BC synthesis [5Reiniati I, Hrymak AN, Margaritis A. Recent developments in the production and applications of bacterial cellulose fibers and nanocrystals. Crit Rev Biotechnol 2017; 37(4): 510-24.
[http://dx.doi.org/10.1080/07388551.2016.1189871] [PMID: 27248159]
-7Lee KY, Buldum G, Mantalaris A, Bismarck A. More than meets the eye in bacterial cellulose: biosynthesis, bioprocessing, and applications in advanced fiber composites. Macromol Biosci 2014; 14(1): 10-32.
[http://dx.doi.org/10.1002/mabi.201300298] [PMID: 23897676]
] (Fig. 1).

The main advantage of BC is the purity of cellulose. Impurities of lignin, hemicelluloses, and pectin in plant derived cellulose enforced the application of harsh chemicals in purification process (environmental safety); however BC purification process needs low energy [8Huang Y, Zhu C, Yang J, Nie Y, Chen C, Sun D. Recent advances in bacterial cellulose. Cellulose 2014; 21(1): 1-30.
[http://dx.doi.org/10.1007/s10570-013-0088-z]
, 9Samfira I, Butnariu M, Rodino S, Butu M. Structural investigation of mistletoe plants from various hosts exhibiting diverse lignin phenotypes. Dig J Nanomater Biostruct 2013; 8: 1679-86.]. Unique properties of BC include crystallinity indexes up to 85% [10Siró I, Plackett D. Microfibrillated cellulose and new nanocomposite materials: a review. Cellulose 2010; 17(3): 459-94.
[http://dx.doi.org/10.1007/s10570-010-9405-y]
], a higher degree of polymerization, significant tensile characterization [11Iguchi M. Review bacterial celullose a masterpiece of natures arts. J Mater Sci 2000; 35: 1-10.
[http://dx.doi.org/10.1023/A:1004775229149]
-13Paximada P, Tsouko E, Kopsahelis N, Koutinas AA, Mandala I. Bacterial cellulose as stabilizer of o/w emulsions. Food Hydrocoll 2016; 53: 225-32.
[http://dx.doi.org/10.1016/j.foodhyd.2014.12.003]
], a specific area in BC fibers, higher water holding capacity, and longer drying time [14Sulaeva I, Henniges U, Rosenau T, Potthast A. Bacterial cellulose as a material for wound treatment: Properties and modifications. A review. Biotechnol Adv 2015; 33(8): 1547-71.
[http://dx.doi.org/10.1016/j.biotechadv.2015.07.009] [PMID: 26253857]
-16Shi Z, Zhang Y, Phillips GO, Yang G. Utilization of bacterial cellulose in food. Food Hydrocoll 2014; 35: 539-45.
[http://dx.doi.org/10.1016/j.foodhyd.2013.07.012]
]. These unique features are related to the high aspect ratio of fibrils, which made the increased surface area hold higher water capacity and tightly bound them to hydroxyl groups. BC generally is very flexible and easy to modify based on its many reactive groups. High porosity and surface area make BC appropriate for physical and chemical interactions with many compounds, including antimicrobials [10Siró I, Plackett D. Microfibrillated cellulose and new nanocomposite materials: a review. Cellulose 2010; 17(3): 459-94.
[http://dx.doi.org/10.1007/s10570-010-9405-y]
, 17Gelin K, Bodin A, Gatenholm P, Mihranyan A, Edwards K, Strømme M. Characterization of water in bacterial cellulose using dielectric spectroscopy and electron microscopy. Polymer (Guildf) 2007; 48(26): 7623-31.
[http://dx.doi.org/10.1016/j.polymer.2007.10.039]
, 18Shah N, Ul-Islam M, Khattak WA, Park JK. Overview of bacterial cellulose composites: a multipurpose advanced material. Carbohydr Polym 2013; 98(2): 1585-98.
[http://dx.doi.org/10.1016/j.carbpol.2013.08.018] [PMID: 24053844]
] (Fig. 2).

Many factors control and structure the BC yield and properties. Bacterial strains, medium composition and cultivation process mainly determine the morphology, properties and eventually range of possible application of BC. Several BC applications were recently developed, for example, biomedical applications of BC mostly focused on materials for tissue engineering, wound dressing, artificial skin, blood vessels, and carriers for drug delivery [1Rajwade JM, Paknikar KM, Kumbhar JV. Applications of bacterial cellulose and its composites in biomedicine. Appl Microbiol Biotechnol 2015; 99(6): 2491-511.
[http://dx.doi.org/10.1007/s00253-015-6426-3] [PMID: 25666681]
, 12Tsouko E, Kourmentza C, Ladakis D, et al. Bacterial cellulose production from industrial waste and by-product streams. Int J Mol Sci 2015; 16(7): 14832-49.
[http://dx.doi.org/10.3390/ijms160714832] [PMID: 26140376]
] (Fig. 3).

Optimization of the production line from strain selection to final modification needs much effort in research. Achieving these goals attracted many researchers to explore better BC producing species/ strains and low-cost media. It is noteworthy to mention that replacing the expensive media Hestrin and Schramm (HS) has been done by many researchers [19Jozala AF, Pértile RAN, dos Santos CA, et al. Bacterial cellulose production by Gluconacetobacter xylinus by employing alternative culture media. Appl Microbiol Biotechnol 2015; 99(3): 1181-90.
[http://dx.doi.org/10.1007/s00253-014-6232-3] [PMID: 25472434]
-23Pacheco G, Nogueira CR, Meneguin AB, Trovatti E, Silva MC, Machado RT, et al. Development and characterization of bacterial cellulose produced by cashew tree residues as alternative carbon source. Ind Crops Prod 2017; 107: 13-9.
[http://dx.doi.org/10.1016/j.indcrop.2017.05.026]
]. However, the BC produced purity was in less degree as those required for some applications such as biomedical and industrial applications.

The production of cellulosic bacteria generally includes the two main steps, bacterial strain and bioprocess production. Static and stirred cultivation, besides semi-continuous or continuous fermentation methods, are the major bioprocess productions. The shapes and properties of final cellulose products are in significant dependence to the strain of bacteria, static, agitated, batch or feed batch production processes [24Lin S-P, Calvar IL, Catchmark JM, Liu J-R, Demirci A, Cheng K-C. Biosynthesis, production and applications of bacterial cellulose. Cellulose 2013; 20(5): 2191-219.
[http://dx.doi.org/10.1007/s10570-013-9994-3]
]. This review paper presents an overview of outstanding findings in BC production and applications.

Fig. (1)
Bacterial cells and 1,4-β-glucan chains (microfibrils) exopolysaccharides.


Fig. (2)
The main advantage of the Bacterial Cellulose (BC).


Fig. (3)
Factors control and structure the BC yield and properties, applications of BC.


2. STRAIN SELECTION

Naturally, the bacterial species of Achromobacter [25Rangaswamy B. K P V, Hungund B. Microbial cellulose production from bacteria isolated from rotten fruit. Int J Polym Sci 2015; 2015: 1-8.
[http://dx.doi.org/10.1155/2015/280784]
, 26Hareesh E. abdul faisal P, Benjamin S. Optimization of parameters for the production of cellulase from Achromobacter xylosoxidans BSS4 by solid-state fermentation. Electro J Bio 2016; 4: 443-8.], Alcaligenes [27Gorgieva S, Trček J. Bacterial Cellulose: Production, modification and perspectives in biomedical applications. Nanomaterials (Basel) 2019; 9(10)E1352
[http://dx.doi.org/10.3390/nano9101352] [PMID: 31547134]
], Aerobacter [28Sunagawa N, Tajima K, Hosoda M, et al. Cellulose production by Enterobacter sp. CJF-002 and identification of genes for cellulose biosynthesis. Cellulose 2012; 19(6): 1989-2001.
[http://dx.doi.org/10.1007/s10570-012-9777-2]
, 29Zhao H, Xia J, Wang J, et al. Production of bacterial cellulose using polysaccharide fermentation wastewater as inexpensive nutrient sources. Biotechnol Biotechnol Equip 2018; 32(2): 350-6.
[http://dx.doi.org/10.1080/13102818.2017.1418673]
], Agrobacterium [30Matthysse AG, Thomas DL, White AR. Mechanism of cellulose synthesis in Agrobacterium tumefaciens. J Bacteriol 1995; 177(4): 1076-81.
[http://dx.doi.org/10.1128/JB.177.4.1076-1081.1995] [PMID: 7860586]
-33Matthysse AG. Attachment of Agrobacterium to plant surfaces. Front Plant Sci 2014; 5: 252.
[http://dx.doi.org/10.3389/fpls.2014.00252] [PMID: 24926300]
], Azotobacter [6Chawla P, Bajaj I, Survase S, Singhal R. Microbial cellulose: fermentative production and applications. Food Technol Biotechnol 2009; 47: 107-24.], Gluconacetobacter [19Jozala AF, Pértile RAN, dos Santos CA, et al. Bacterial cellulose production by Gluconacetobacter xylinus by employing alternative culture media. Appl Microbiol Biotechnol 2015; 99(3): 1181-90.
[http://dx.doi.org/10.1007/s00253-014-6232-3] [PMID: 25472434]
, 34Costa AFS, Almeida FCG, Vinhas GM, Sarubbo LA. Production of bacterial cellulose by Gluconacetobacter hansenii using corn steep liquor as nutrient sources. Front Microbiol 2017; 8: 2027.
[http://dx.doi.org/10.3389/fmicb.2017.02027] [PMID: 29089941]
, 35Du R, Zhao F, Peng Q, Zhou Z, Han Y. Production and characterization of bacterial cellulose produced by Gluconacetobacter xylinus isolated from Chinese persimmon vinegar. Carbohydr Polym 2018; 194: 200-7.
[http://dx.doi.org/10.1016/j.carbpol.2018.04.041] [PMID: 29801830]
], Pseudomonas [36Raheem Kazim A. Production, optimization, and characterization of cellulose produced from Pseudomonas spp. World Journal of Experimental Biosciences 2015; 3(2): 89-93.], Rhizobium [37Ahmed S, Raheem Kazim A, Mahmood H. Increasing Cellulose Production from Rhizobium leguminosarum bv. viciae. Journal of Al-Nahrain University- Science 2017; 20(1): 120-5., 38Robledo M, Rivera L, Jiménez-Zurdo JI, et al. Role of Rhizobium endoglucanase CelC2 in cellulose biosynthesis and biofilm formation on plant roots and abiotic surfaces. Microb Cell Fact 2012; 11: 125.
[http://dx.doi.org/10.1186/1475-2859-11-125] [PMID: 22970813]
], Sarcina [39Moniri M, Boroumand Moghaddam A, Azizi S, et al. Production and status of bacterial cellulose in biomedical engineering. Nanomaterials (Basel) 2017; 7(9): 257.
[http://dx.doi.org/10.3390/nano7090257] [PMID: 32962322]
] and Dickeya and Rhodobacter [40Ji K, Wang W, Zeng B, et al. Bacterial cellulose synthesis mechanism of facultative anaerobe Enterobacter sp. FY-07. Sci Rep 2016; 6: 21863.
[http://dx.doi.org/10.1038/srep21863] [PMID: 26911736]
] can produce cellulose.

However, the Gluconacetobacter genus is the primary group of BC producers. This fact is due to the wide range of carbon and nitrogen sources used by the Gluconacetobacter genus [7Lee KY, Buldum G, Mantalaris A, Bismarck A. More than meets the eye in bacterial cellulose: biosynthesis, bioprocessing, and applications in advanced fiber composites. Macromol Biosci 2014; 14(1): 10-32.
[http://dx.doi.org/10.1002/mabi.201300298] [PMID: 23897676]
, 41Ul-Islam M, Khan S, Ullah MW, Park JK. Bacterial cellulose composites: Synthetic strategies and multiple applications in bio-medical and electro-conductive fields. Biotechnol J 2015; 10(12): 1847-61.
[http://dx.doi.org/10.1002/biot.201500106] [PMID: 26395011]
]. In recent days with biotechnology, the engineered bacteria with low consuming nutrients and high yield were produced [41Ul-Islam M, Khan S, Ullah MW, Park JK. Bacterial cellulose composites: Synthetic strategies and multiple applications in bio-medical and electro-conductive fields. Biotechnol J 2015; 10(12): 1847-61.
[http://dx.doi.org/10.1002/biot.201500106] [PMID: 26395011]
, 42Liu M, Li S, Xie Y, et al. Enhanced bacterial cellulose production by Gluconacetobacter xylinus via expression of Vitreoscilla hemoglobin and oxygen tension regulation. Appl Microbiol Biotechnol 2018; 102(3): 1155-65.
[http://dx.doi.org/10.1007/s00253-017-8680-z] [PMID: 29199354]
]. For example, Komagataeibacter rhaeticus was engineered and fully sequenced with low nitrogen and high yield. It is noteworthy to mention that genetic tool kit was developed for this bacterium to engineer the desired bacteria recently [43Florea M, Hagemann H, Santosa G, et al. Engineering control of bacterial cellulose production using a genetic toolkit and a new cellulose-producing strain. Proc Natl Acad Sci USA 2016; 113(24): E3431-40.
[http://dx.doi.org/10.1073/pnas.1522985113] [PMID: 27247386]
].

Fig. (4)
Genetic mechanisms of cellulose production. Cellulose synthase (4p02.pdb).


3. GENETIC MECHANISMS OF CELLULOSE PRODUCTION

Uridine diphosphoglucose (UDPGIc) synthesis and polymerization of glucose are the two main cellulose production steps in bacteria. The polymerization is achieved by cellulose synthase, which formed an unbranched chain of 1,4-β-glucan. Naturally, carbon compound of hexoses, glycerol, dihydroxyacetone, pyruvate, and dicarboxylic acids entering the Krebs cycle, gluconeogenesis or the pentose phosphate cycle, followed by phosphorylation and isomerization and finally UDPGIc pyrophosphorylase converts the compounds into UDPGIc, a precursor for the cellulose production. The efficiency of this production is different in different bacteria, for example, for A. xylinum is 50% [27Gorgieva S, Trček J. Bacterial Cellulose: Production, modification and perspectives in biomedical applications. Nanomaterials (Basel) 2019; 9(10)E1352
[http://dx.doi.org/10.3390/nano9101352] [PMID: 31547134]
] (Fig. 4). Remarkable genetic engineering approach was presented recently. In this approach, the most efficient microbial BC producer, Komagataeibacter rhaeticus iGEM, was isolated, and its full genome was sequenced. This bacterium was then engineered for functionalization of cellulose production. The functionalization engineering results will help other researchers apply a similar method for producing the more specific BC with unique pattern for biomaterial applications [43Florea M, Hagemann H, Santosa G, et al. Engineering control of bacterial cellulose production using a genetic toolkit and a new cellulose-producing strain. Proc Natl Acad Sci USA 2016; 113(24): E3431-40.
[http://dx.doi.org/10.1073/pnas.1522985113] [PMID: 27247386]
].

4. CULTURE MEDIUM

BC medium is costly, as it needs a high demand of glucose and other nutrients [44Hungund B, Prabhu S, Shetty C, Acharya S, Prabhu V, Gupta S. Production of bacterial cellulose from Gluconacetobacter persimmonis GH-2 using dual and cheaper carbon sources. J Microb Biochem Technol 2013; 5(2): 31-3.
[http://dx.doi.org/10.4172/1948-5948.1000095]
]. Hestrin and Schramm (HS) medium (glucose, peptone, and yeast extract) is used for BC production; however, the uses of the wastes, foods, and fruits as a medium have been reported recently [44Hungund B, Prabhu S, Shetty C, Acharya S, Prabhu V, Gupta S. Production of bacterial cellulose from Gluconacetobacter persimmonis GH-2 using dual and cheaper carbon sources. J Microb Biochem Technol 2013; 5(2): 31-3.
[http://dx.doi.org/10.4172/1948-5948.1000095]
-50Ul-Islam M, Khan T, Park JK. Water holding and release properties of bacterial cellulose obtained by in situ and ex situ modification. Carbohydr Polym 2012; 88(2): 596-603. a
[http://dx.doi.org/10.1016/j.carbpol.2012.01.006]
]. For example, fruit juices such as orange, pineapple, apple, Japanese pear, and grape showed a higher yield in Acetobacter xylinum BC production [46Kurosumi A, Sasaki C, Yamashita Y, Nakamura Y. Utilization of various fruit juices as carbon source for production of bacterial cellulose by Acetobacter xylinum NBRC 13693. Carbohydr Polym 2009; 76(2): 333-5.
[http://dx.doi.org/10.1016/j.carbpol.2008.11.009]
]. Furthermore, other fruits such as pineapple, pomegranate, muskmelon, watermelon, tomato, orange, and also molasses, starch hydrolysate, sugarcane juice, coconut water, coconut milk were used as carbon sources [44Hungund B, Prabhu S, Shetty C, Acharya S, Prabhu V, Gupta S. Production of bacterial cellulose from Gluconacetobacter persimmonis GH-2 using dual and cheaper carbon sources. J Microb Biochem Technol 2013; 5(2): 31-3.
[http://dx.doi.org/10.4172/1948-5948.1000095]
]. The same properties of BC from these sources were reported with more expensive media [48Tyagi N, Suresh S. Production of cellulose from sugarcane molasses using Gluconacetobacter intermedius SNT-1: optimization & characterization. J Clean Prod 2016; 112: 71-80.
[http://dx.doi.org/10.1016/j.jclepro.2015.07.054]
]. Organic acid, carbohydrates, ethanol, and acetic acid, have been used as additives in BC production media [51Ul-Islam M, Khan T, Park JK. Nanoreinforced bacterial cellulose-montmorillonite composites for biomedical applications. Carbohydr Polym 2012; 89(4): 1189-97. b
[http://dx.doi.org/10.1016/j.carbpol.2012.03.093] [PMID: 24750931]
, 52Ahmed S, Kanchi S, Kumar G. Handbook of biopolymers: Advances and multifaceted applications 2018.
[http://dx.doi.org/10.1201/9780429024757]
] (Fig. 5).

Optimization of the media during the process is the key to succeed in producing sustainable BC. Many factors such as pH, oxygen supply, and temperature should be controlled and optimized during the process [7Lee KY, Buldum G, Mantalaris A, Bismarck A. More than meets the eye in bacterial cellulose: biosynthesis, bioprocessing, and applications in advanced fiber composites. Macromol Biosci 2014; 14(1): 10-32.
[http://dx.doi.org/10.1002/mabi.201300298] [PMID: 23897676]
, 41Ul-Islam M, Khan S, Ullah MW, Park JK. Bacterial cellulose composites: Synthetic strategies and multiple applications in bio-medical and electro-conductive fields. Biotechnol J 2015; 10(12): 1847-61.
[http://dx.doi.org/10.1002/biot.201500106] [PMID: 26395011]
]. All of these factors affect the yield and properties of the final products. The pH and temperature are highly dependent to the species and strain of bacteria. Generally, pH could be between 4.0- 7.0. On the other hand, the pH of the culture can directly control the accumulation time and the secondary metabolite. This can affect nutrition supply in the process. For example, the dried weight of BC in Acetobacter xylinum 0416 was 60% higher in the control pH system compared to the uncontrolled. Furthermore, this bacterium's growth rate was 30% higher in lower pH conditions [53Zahan KA. 'Pa’e N, Muhamad II. Monitoring the effect of ph on bacterial cellulose production and Acetobacter xylinum 0416 Growth in a Rotary Discs Reactor. Arab J Sci Eng 2015; 40(7): 1881-5.
[http://dx.doi.org/10.1007/s13369-015-1712-z]
]. Also, aeration and oxygen optimization can play a critical role in BC production. Insufficient oxygen supply inhibits bacterial growth, and high oxygen supply favor gluconic acid production [7Lee KY, Buldum G, Mantalaris A, Bismarck A. More than meets the eye in bacterial cellulose: biosynthesis, bioprocessing, and applications in advanced fiber composites. Macromol Biosci 2014; 14(1): 10-32.
[http://dx.doi.org/10.1002/mabi.201300298] [PMID: 23897676]
, 51Ul-Islam M, Khan T, Park JK. Nanoreinforced bacterial cellulose-montmorillonite composites for biomedical applications. Carbohydr Polym 2012; 89(4): 1189-97. b
[http://dx.doi.org/10.1016/j.carbpol.2012.03.093] [PMID: 24750931]
, 54Wu SC, Li MH. Production of bacterial cellulose membranes in a modified airlift bioreactor by Gluconacetobacter xylinus. J Biosci Bioeng 2015; 120(4): 444-9.
[http://dx.doi.org/10.1016/j.jbiosc.2015.02.018] [PMID: 25823854]
]. Temperature as an essential factor in BC production should optimize carefully. For example, optimal Komagataeibacter xylinus B-12068 temperature growth is preferably 28-30 °C [52Ahmed S, Kanchi S, Kumar G. Handbook of biopolymers: Advances and multifaceted applications 2018.
[http://dx.doi.org/10.1201/9780429024757]
] (Fig. 6).

Fig. (5)
Cellulose production by different strains of A. xylinum.


Fig. (6)
Cellulose production of different species.


5. CULTIVATION METHODS

The cultivation mode of the BC can be static or agitated. Static modes of cultivation usually take 5-20 days due to the bacterial strain and nutrients supply. The production is generally on the area of air/ liquid interface [24Lin S-P, Calvar IL, Catchmark JM, Liu J-R, Demirci A, Cheng K-C. Biosynthesis, production and applications of bacterial cellulose. Cellulose 2013; 20(5): 2191-219.
[http://dx.doi.org/10.1007/s10570-013-9994-3]
]. The final BC membrane shape depends on the material used for growing the BC in this method. This method used for the predefined shape of BC required. Disadvantages of this mode of action include low yield and more time consumption. [55Lin D, Lopez-Sanchez P, Li R, Li Z. Production of bacterial cellulose by Gluconacetobacter hansenii CGMCC 3917 using only waste beer yeast as nutrient source. Bioresour Technol 2014; 151: 113-9.
[http://dx.doi.org/10.1016/j.biortech.2013.10.052] [PMID: 24212131]
, 56Esa F, Tasirin SM, Rahman NA. Overview of bacterial cellulose production and application. Agric Agric Sci Procedia 2014; 2: 113-9.
[http://dx.doi.org/10.1016/j.aaspro.2014.11.017]
]. Fed-batch cultivation could help to overcome these problems [56Esa F, Tasirin SM, Rahman NA. Overview of bacterial cellulose production and application. Agric Agric Sci Procedia 2014; 2: 113-9.
[http://dx.doi.org/10.1016/j.aaspro.2014.11.017]
]. Researchers showed two to three times' higher yields with fed-batch culture compared to batch cultivation [49Shezad O, Khan S, Khan T, Park JK. Production of bacterial cellulose in static conditions by a simple fed-batch cultivation strategy. Korean J Chem Eng 2009; 26(6): 1689-92.
[http://dx.doi.org/10.1007/s11814-009-0232-5]
]. Specific bioreactors were produced to use in the static mode culture of BC recently [57Kralisch D, Hessler N, Klemm D, Erdmann R, Schmidt W. White biotechnology for cellulose manufacturing--the HoLiR concept. Biotechnol Bioeng 2010; 105(4): 740-7.
[PMID: 19816981]
, 58Kim D, Ku S. Beneficial effects of Monascus sp. KCCM 10093 pigments and derivatives: a mini review. Molecules 2018; 23(1): 98.
[http://dx.doi.org/10.3390/molecules23010098] [PMID: 29301350]
]. Another mode of cultivation is agitated, which has a more oxygen supply therefore, the yield is higher than the static mode [59Hornung M, Ludwig M, Schmauder H. Optimizing the production of bacterial cellulose in surface culture: a novel aerosol bioreactor working on a fed batch principle (Part 3). Eng Life Sci 2007; 7(1): 35-41.
[http://dx.doi.org/10.1002/elsc.200620164]
]. The shape of BC final product depends on the agitation speed. The drawback of the agitate mode of cultivation is the production of cellulose-negative mutants population. This mutant can produce BC with different properties as a subpopulation. To overcome this problem, the use of ethanol as a supplement nutrient is recommended [50Ul-Islam M, Khan T, Park JK. Water holding and release properties of bacterial cellulose obtained by in situ and ex situ modification. Carbohydr Polym 2012; 88(2): 596-603. a
[http://dx.doi.org/10.1016/j.carbpol.2012.01.006]
, 58Kim D, Ku S. Beneficial effects of Monascus sp. KCCM 10093 pigments and derivatives: a mini review. Molecules 2018; 23(1): 98.
[http://dx.doi.org/10.3390/molecules23010098] [PMID: 29301350]
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[http://dx.doi.org/10.1042/BA20000065] [PMID: 11171030]
]. Stirred tank bioreactor can be used in the agitated mode for BC production. Shear stress is a great drawback of this BC production mode [62Shoda M, Sugano Y. Recent advances in bacterial cellulose production. Biotechnol Bioprocess Eng; BBE 2005; 10(1): 1.
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]. Airlift bioreactor is another bioreactor with better efficient results [54Wu SC, Li MH. Production of bacterial cellulose membranes in a modified airlift bioreactor by Gluconacetobacter xylinus. J Biosci Bioeng 2015; 120(4): 444-9.
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]. In this bioreactor the oxygen supply is provided from the bottom of the tank, therefore preparing better BC production conditions [64Chao Y-p, Sugano Y, Kouda T, Yoshinaga F, Shoda M. Production of bacterial cellulose by Acetobacter xylinumwith an airlift reactor. Biotechnol Tech 1997; 11(11): 829-32.
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6. SCRUTINIZING THE BC FORMATION: FROM PLANKTONIC CELL TO BIOFILM

Many researchers observed the biofilm in general with a simple microscope [65Heukelekian H, Heller A. Relation between food concentration and surface for bacterial growth. J Bacteriol 1940; 40(4): 547-58.
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, 66Zobell CE. The Effect of solid surfaces upon bacterial activity. J Bacteriol 1943; 46(1): 39-56.
[http://dx.doi.org/10.1128/JB.46.1.39-56.1943] [PMID: 16560677]
]. However, the high-resolution electron microscope allowed us to examine the specific detailed characterizations of microbial biofilms [67Jones HC, Roth IL, Sanders WM III. Electron microscopic study of a slime layer. J Bacteriol 1969; 99(1): 316-25.
[http://dx.doi.org/10.1128/JB.99.1.316-325.1969] [PMID: 5802613]
]. It is noteworthy to mention that Ruthenium red (that stains the polysaccharide) facilitated the 'researcher's effort to show the matrix (structure) of biofilm based on the polysaccharide. Furthermore, the gene regulation studies and utilization of laser scanning microscope had a significant impact on understanding biofilm formation and characterization [67Jones HC, Roth IL, Sanders WM III. Electron microscopic study of a slime layer. J Bacteriol 1969; 99(1): 316-25.
[http://dx.doi.org/10.1128/JB.99.1.316-325.1969] [PMID: 5802613]
-69Costerton JW, Geesey GG, Cheng KJ. How bacteria stick. Sci Am 1978; 238(1): 86-95.
[http://dx.doi.org/10.1038/scientificamerican0178-86] [PMID: 635520]
]. Traditionally biofilm is defined as the assemblage of 'microorganism's cells on the surface. This structure is irreversibly associated with themselves and the surrounding matrix. Biofilm matrix has a unique structure base on the cell and non-cellular materials of the medium. The main feature of biofilm is attachments that can be defined as unique substratum interactions, culture medium, hydrodynamic of aqueous environment and cell surface characteristics. These features are very important for scrutinizing the BC formation specifically and bacterial biofilm formation in general. Substratum is considered as the solid surfaces such as tissue, indwelling medical devices, industrial devices (piping), and natural aquatic systems. The physiochemical features of the surface play an important role in attachment; these features can effect on rate and extent of biofilm formation. Rough hydrophobic nonpolar surfaces are more favorable for rapid biofilm formation [70Fletcher M, Loeb GI. Influence of substratum characteristics on the attachment of a marine pseudomonad to solid surfaces. Appl Environ Microbiol 1979; 37(1): 67-72.
[http://dx.doi.org/10.1128/AEM.37.1.67-72.1979] [PMID: 16345338]
-74Bales PM, Renke EM, May SL, Shen Y, Nelson DC. Purification and characterization of biofilm-associated EPS Exopolysaccharides from ESKAPE organisms and other pathogens. PLoS One 2013; 8(6)e67950
[http://dx.doi.org/10.1371/journal.pone.0067950] [PMID: 23805330]
]. It is important to note that the surface of materials in an aqueous environment could facilitate and increase biofilm formation speed [75Loeb GI, Neihof RA. Marine conditioning films Applied Chemistry at Protein Interfaces Advances in Chemistry 145 1975; 319-35.
[http://dx.doi.org/10.1021/ba-1975-0145.ch016]
, 76Colvin KM, Irie Y, Tart CS, et al. The Pel and Psl polysaccharides provide Pseudomonas aeruginosa structural redundancy within the biofilm matrix. Environ Microbiol 2012; 14(8): 1913-28.
[http://dx.doi.org/10.1111/j.1462-2920.2011.02657.x] [PMID: 22176658]
]. The hydrodynamic of the environment with flow velocity could control biofilm formation (the higher velocity made thinner biofilm boundary layer). Furthermore, the settle down of cell suspension in the aqueous medium can control flow velocity [77Rijnaarts HH, Norde W, Bouwer EJ, Lyklema J, Zehnder AJ. Bacterial adhesion under static and dynamic conditions. Appl Environ Microbiol 1993; 59(10): 3255-65.
[http://dx.doi.org/10.1128/AEM.59.10.3255-3265.1993] [PMID: 16349063]
-79Mika F, Hengge R. Small RNAs in the control of RpoS, CsgD, and biofilm architecture of Escherichia coli. RNA Biol 2014; 11(5): 494-507.
[http://dx.doi.org/10.4161/rna.28867] [PMID: 25028968]
].

The medium characteristics have a great impact on biofilm formation. pH, nutrient level, and temperature can play a substantial role in forming biofilm [80Donlan RM, Pipes WO, Yohe TL. Biofilm formation on cast iron substrata in water distribution systems. Water Res 1994; 28(6): 1497-503.
[http://dx.doi.org/10.1016/0043-1354(94)90318-2]
-84Bartowsky EJ, Henschke PA. Acetic acid bacteria spoilage of bottled red wine -- a review. Int J Food Microbiol 2008; 125(1): 60-70.
[http://dx.doi.org/10.1016/j.ijfoodmicro.2007.10.016] [PMID: 18237809]
]. It is important to recall here that bacterial cell surface properties such as hydrophobicity and flagella influence the attachment and rate of biofilm formation. Furthermore, the production of EPS has a great impact on biofilm development [85Rosenberg M, Kjelleberg S. Hydrophobic Interactions: Role in Bacterial Adhesion.Advances in Microbial Ecology Boston, MA 1986; 353-93.
[http://dx.doi.org/10.1007/978-1-4757-0611-6_8]
-88Bashan Y, Levanony H. Active attachment of Azospirillum brasilense Cd to Quartz Sand and to a Light-textured Soil by Protein Bridging. Microbiology 1988; 134(8): 2269-79.
[http://dx.doi.org/10.1099/00221287-134-8-2269]
]. Genes regulations have a countless effect on biofilm formation; for example, in Pseudomonas aeruginosa algC gene up regulation was observed during the bacterial attachments [89Davies DG, Geesey GG. Regulation of the alginate biosynthesis gene algC in Pseudomonas aeruginosa during biofilm development in continuous culture. Appl Environ Microbiol 1995; 61(3): 860-7.
[http://dx.doi.org/10.1128/AEM.61.3.860-867.1995] [PMID: 7793920]
].

Moreover, polyphosphokinase synthesis genes in Pseudomonas aeruginosa were up-regulated [90Prigent-Combaret C, Vidal O, Dorel C, Lejeune P. Abiotic surface sensing and biofilm-dependent regulation of gene expression in Escherichia coli. J Bacteriol 1999; 181(19): 5993-6002.
[http://dx.doi.org/10.1128/JB.181.19.5993-6002.1999] [PMID: 10498711]
]. The up-regulated genes in Staphylococcus aureus genes involved in the glycolysis pathway was reported previously [91Becker P, Hufnagle W, Peters G, Herrmann M. Detection of differential gene expression in biofilm-forming versus planktonic populations of Staphylococcus aureus using micro-representational-difference analysis. Appl Environ Microbiol 2001; 67(7): 2958-65.
[http://dx.doi.org/10.1128/AEM.67.7.2958-2965.2001] [PMID: 11425708]
]. The detailed information on gene regulation of biofilm formation was reviewed earlier [91Becker P, Hufnagle W, Peters G, Herrmann M. Detection of differential gene expression in biofilm-forming versus planktonic populations of Staphylococcus aureus using micro-representational-difference analysis. Appl Environ Microbiol 2001; 67(7): 2958-65.
[http://dx.doi.org/10.1128/AEM.67.7.2958-2965.2001] [PMID: 11425708]
].

7. DOWNSTREAM PROCESSING

These parts of the production consist of separation of BC from media and purification of biopolymers. The BC can remove easily from static mode cultivation by separation. In agitated mode BC removes from the media by centrifugation or filtration. The alkali treatment will do the purification of cellulose with NaOH or KOH [5Reiniati I, Hrymak AN, Margaritis A. Recent developments in the production and applications of bacterial cellulose fibers and nanocrystals. Crit Rev Biotechnol 2017; 37(4): 510-24.
[http://dx.doi.org/10.1080/07388551.2016.1189871] [PMID: 27248159]
, 6Chawla P, Bajaj I, Survase S, Singhal R. Microbial cellulose: fermentative production and applications. Food Technol Biotechnol 2009; 47: 107-24.]. The level of purification is depending to the final application, for example we need more purify cellulose for biomedical applications compare to the food application. Sometimes drying step adds to the downstream process. Three kinds of room, oven, freeze and supercritical drying methods were used in this step. Drying can change BC's characteristic; therefore, it should be chosen very carefully [92Zeng M, Laromaine A, Roig A. Bacterial cellulose films: influence of bacterial strain and drying route on film properties. Cellulose 2014; 21(6): 4455-69.
[http://dx.doi.org/10.1007/s10570-014-0408-y]
].

8. FORMS OF BACTERIAL CELLULOSE

Intact membrane, disassembled BC and BC nanocrystals are three BC forms [41Ul-Islam M, Khan S, Ullah MW, Park JK. Bacterial cellulose composites: Synthetic strategies and multiple applications in bio-medical and electro-conductive fields. Biotechnol J 2015; 10(12): 1847-61.
[http://dx.doi.org/10.1002/biot.201500106] [PMID: 26395011]
, 93Zhu H, Jia S, Yang H, Tang W, Jia Y, Tan Z. Characterization of bacteriostatic sausage casing: a composite of bacterial cellulose embedded with ɛ-polylysine. Food Sci Biotechnol 2010; 19(6): 1479-84.
[http://dx.doi.org/10.1007/s10068-010-0211-y]
, 94Chang S-T, Chen L-C, Lin S-B, Chen H-H. Nano-biomaterials application: Morphology and physical properties of bacterial cellulose/gelatin composites via crosslinking. Food Hydrocoll 2012; 27(1): 137-44.
[http://dx.doi.org/10.1016/j.foodhyd.2011.08.004]
]. An intact membrane can be immersed in dispersion with other materials. This has the advantage of simpler disintegration from other materials but cannot change the fermentation process [50Ul-Islam M, Khan T, Park JK. Water holding and release properties of bacterial cellulose obtained by in situ and ex situ modification. Carbohydr Polym 2012; 88(2): 596-603. a
[http://dx.doi.org/10.1016/j.carbpol.2012.01.006]
, 93Zhu H, Jia S, Yang H, Tang W, Jia Y, Tan Z. Characterization of bacteriostatic sausage casing: a composite of bacterial cellulose embedded with ɛ-polylysine. Food Sci Biotechnol 2010; 19(6): 1479-84.
[http://dx.doi.org/10.1007/s10068-010-0211-y]
, 94Chang S-T, Chen L-C, Lin S-B, Chen H-H. Nano-biomaterials application: Morphology and physical properties of bacterial cellulose/gelatin composites via crosslinking. Food Hydrocoll 2012; 27(1): 137-44.
[http://dx.doi.org/10.1016/j.foodhyd.2011.08.004]
]. BC membrane on the other hand, can be disassembled, and therefore they are easier to integrate with other materials and even become better powder and film [95Guo Y, Zhang X, Hao W, et al. Nano-bacterial cellulose/soy protein isolate complex gel as fat substitutes in ice cream model. Carbohydr Polym 2018; 198: 620-30.
[http://dx.doi.org/10.1016/j.carbpol.2018.06.078] [PMID: 30093042]
-97Viana RM, Sá NMSM, Barros MO, Borges MF, Azeredo HMC. Nanofibrillated bacterial cellulose and pectin edible films added with fruit purees. Carbohydr Polym 2018; 196: 27-32.
[http://dx.doi.org/10.1016/j.carbpol.2018.05.017] [PMID: 29891296]
]. BC nanocrystal has been produced by acid hydrolysis, which removes the amorphous of the cellulose [98'O’Connor B, Berry R, Goguen R Commercialization of cellulose nanocrystal (NCC™) production: A business case focusing on the importance of proactive EHS management Nanotechnology environmental health and safety 2014; 225-46.].

9. STRUCTURE OF BACTERIAL CELLULOSE

Cellulose indeed is polysaccharide consisting of carbon, hydrogen, and oxygen. This carbohydrate polymeric structure composes of glucose units. The role of cellulose in the bacterial film is stability towards different chemical or temperature environments. High mechanical strength, crystallinity, and ultra-fine and pure fiber network are very dependent on highly insoluble and inelastic cellulose fibrils. Pore and tunnels within the thin layer (pellicle) of bacterial cellulose can retain water 16 times higher than plant cellulose [99dos Santos MA, Grenha A. Polysaccharide nanoparticles for protein and Peptide delivery: exploring less-known materials. Adv Protein Chem Struct Biol 2015; 98: 223-61.
[http://dx.doi.org/10.1016/bs.apcsb.2014.11.003] [PMID: 25819281]
].

The cellulose pellicle was formed on the upper surface of air/ media, which shows the importance of oxygen for this process. The pellicle of bacterial cellulose with 15 GPa (Young modulus) [100Vigentini I, Fabrizio V, Dellacà F, Rossi S, Azario I, Mondin C, et al. Set-up of bacterial cellulose production from the genus Komagataeibacter and its use in a gluten-free bakery product as a case study. Front Microbiol 1953; 2019: 10.
[PMID: 31551945]
] is considered as a very tough polymeric film. This unique structure is related to fibrils conformation, which is bound tightly by hydrogen bonds. The supermolecular structures of aligned glucan chains with inter and intrachain hydrogen bonds formed the microfibrils (Fig. 7). These microfibrils randomly assemble the fibrils, which literally construct the bacterial cellulose pellicle. It is noteworthy that bacterial cellulose belongs crystallographically to Cellulose I, which means the crystalline fibrils are in parallel arrangement [101Jacek P, Dourado F, Gama M, Bielecki S. Molecular aspects of bacterial nanocellulose biosynthesis. Microb Biotechnol 2019; 12(4): 633-49.
[http://dx.doi.org/10.1111/1751-7915.13386] [PMID: 30883026]
]. Various types of irregularities of fibrils structures of cellulose (kinks or twists of the microfibrils and capillaries) provide the structural heterogeneity with much greater surface area compared to smooth fiber (Fig. 7).

10. MAIN APPLICATIONS OF BC

BC is recognized by the US food and drug administration as generally safe (GRAS) food [16Shi Z, Zhang Y, Phillips GO, Yang G. Utilization of bacterial cellulose in food. Food Hydrocoll 2014; 35: 539-45.
[http://dx.doi.org/10.1016/j.foodhyd.2013.07.012]
]. As a fiber, BC is considered good indigestible food and prescribed in humans as dietetic food [102Mohite BV, Patil SV. A novel biomaterial: bacterial cellulose and its new era applications. Biotechnol Appl Biochem 2014; 61(2): 101-10.
[http://dx.doi.org/10.1002/bab.1148] [PMID: 24033726]
, 103Fontana JD, Koop HS, Tiboni M, Grzybowski A, Pereira A, Kruger CD, et al. New insights on bacterial cellulose Food biosynthesis 2017; 213-49.]. One of the most famous BC is Nata-de-coco, which is the BC grown from coconut water with carbohydrates and amino acid. This cubic BC is immersed in sugar syrup [104Zhang J, Yang Y, Deng J, Wang Y, Hu Q, Li C, et al. Dynamic profile of the microbiota during coconut water pre-fermentation for nata de coco production. Lebensm Wiss Technol 2017; 81: 87-93.
[http://dx.doi.org/10.1016/j.lwt.2017.03.036]
].

Fig. (7)
Structure of Bacterial cellulose.


The product export reached 6.6 billion USD in 2011 [105Piadozo MES. Nata de coco industry in the Philippines Bacterial nanocellulose 2016; 215-29.
[http://dx.doi.org/10.1016/B978-0-444-63458-0.00013-5]
]. This kind of BC can be considered low calorie desserts / snacks and plays a great role as a novel foamy and crunchy product. On the other hand, the fats in the food are always associated with several health problems such as obesity, diabetes, high blood cholesterol levels, and heart diseases, therefore, replacing the fat with BC can help to improve the food industry and human health [106Rios RV, Garzón R, Lannes SCS, Rosell CM. Use of succinyl chitosan as fat replacer on cake formulations. LWT 2018; 96: 260-5.
[http://dx.doi.org/10.1016/j.lwt.2018.05.041]
-108Aydinol P, Ozcan T. Production of reduced‐fat Labneh cheese with inulin and β‐glucan fibre‐based fat replacer. Int J Dairy Technol 2018; 71(2): 362-71.
[http://dx.doi.org/10.1111/1471-0307.12456]
]. BC is already used in meatballs upto 20% or 10% [42Liu M, Li S, Xie Y, et al. Enhanced bacterial cellulose production by Gluconacetobacter xylinus via expression of Vitreoscilla hemoglobin and oxygen tension regulation. Appl Microbiol Biotechnol 2018; 102(3): 1155-65.
[http://dx.doi.org/10.1007/s00253-017-8680-z] [PMID: 29199354]
, 55Lin D, Lopez-Sanchez P, Li R, Li Z. Production of bacterial cellulose by Gluconacetobacter hansenii CGMCC 3917 using only waste beer yeast as nutrient source. Bioresour Technol 2014; 151: 113-9.
[http://dx.doi.org/10.1016/j.biortech.2013.10.052] [PMID: 24212131]
].

Meat is analogous with BC and the mold Monascus purpureus was prepared and introduced with many advantages such as antihypercholesterole. This Monascus-BC complex can be a good substitute for consumers with special dietary restrictions [58Kim D, Ku S. Beneficial effects of Monascus sp. KCCM 10093 pigments and derivatives: a mini review. Molecules 2018; 23(1): 98.
[http://dx.doi.org/10.3390/molecules23010098] [PMID: 29301350]
, 109Sheu F, Wang CL, Shyu YT. Fermentation of Monascus purpureus on bacterial cellulose-nata and the color stability of monascus-nata complex. J Food Sci 2000; 65(2): 342-5.
[http://dx.doi.org/10.1111/j.1365-2621.2000.tb16004.x]
, 110Ullah H, Santos HA, Khan T. Applications of bacterial cellulose in food, cosmetics and drug delivery. Cellulose 2016; 23(4): 2291-314.
[http://dx.doi.org/10.1007/s10570-016-0986-y]
]. Furthermore, BC was used as a thicker food product to increase the strength in gelling products and water binding. Also, it was applied as a stabilizer of Pickering emulsions [13Paximada P, Tsouko E, Kopsahelis N, Koutinas AA, Mandala I. Bacterial cellulose as stabilizer of o/w emulsions. Food Hydrocoll 2016; 53: 225-32.
[http://dx.doi.org/10.1016/j.foodhyd.2014.12.003]
, 111Dankovich TA, Gray DG. Contact angle measurements on smooth nanocrystalline cellulose (I) thin films. J Adhes Sci Technol 2011; 25(6-7): 699-708.
[http://dx.doi.org/10.1163/016942410X525885]
]. As an immobilizing agent of probiotics and enzyme, BC also has great attention for many researchers [112Chávarri M, Marañón I, Villarán MC. Encapsulation technology to protect probiotic bacteria. Probiotics: IntechOpen 2012.
[http://dx.doi.org/10.5772/50046]
, 113Fijałkowski K, Peitler D, Rakoczy R, Żywicka A. Survival of probiotic lactic acid bacteria immobilized in different forms of bacterial cellulose in simulated gastric juices and bile salt solution. Lebensm Wiss Technol 2016; 68: 322-8.
[http://dx.doi.org/10.1016/j.lwt.2015.12.038]
]. It is noteworthy to mention that lipase, laccase and lysozyme immobilized by BC were reported by researchers [114Wu S-C, Wu S-M, Su F-M. Novel process for immobilizing an enzyme on a bacterial cellulose membrane through repeated absorption. J Chem Technol Biotechnol 2017; 92(1): 109-14.
[http://dx.doi.org/10.1002/jctb.4994]
-116Bayazidi P, Almasi H, Asl AK. Immobilization of lysozyme on bacterial cellulose nanofibers: Characteristics, antimicrobial activity and morphological properties. Int J Biol Macromol 2018; 107(Pt B): 2544-51.
[http://dx.doi.org/10.1016/j.ijbiomac.2017.10.137] [PMID: 29079438]
].

BC applications in food packaging as a film and coating were also reported by many researchers [117Otoni CG, Avena‐Bustillos RJ, Azeredo HM, Lorevice MV, Moura MR, Mattoso LH, et al. Recent advances on edible films based on fruits and vegetables—a review. Compr Rev Food Sci Food Saf 2017; 16(5): 1151-69.
[http://dx.doi.org/10.1111/1541-4337.12281]
]. Impregnating BC with other polymers could bring many advantages to the composites. BC-chitosan produced by researchers recently showed the antimicrobial activity against gram-positive and gram-negative bacteria beside the better elastic property [118Ross P, Mayer R, Benziman M. Cellulose biosynthesis and function in bacteria. Microbiol Rev 1991; 55(1): 35-58.
[http://dx.doi.org/10.1128/MMBR.55.1.35-58.1991] [PMID: 2030672]
].

In the biomedical application of BC, several BC-composites were developed by researchers. The wound healing, skin and tissue regeneration, healing under infectious environment, development of artificial organs, blood vessels, skin substitutes, and many devices such as conducting devices, displays, optoelectronics, sensors and biosensors have been reported recently [18Shah N, Ul-Islam M, Khattak WA, Park JK. Overview of bacterial cellulose composites: a multipurpose advanced material. Carbohydr Polym 2013; 98(2): 1585-98.
[http://dx.doi.org/10.1016/j.carbpol.2013.08.018] [PMID: 24053844]
, 102Mohite BV, Patil SV. A novel biomaterial: bacterial cellulose and its new era applications. Biotechnol Appl Biochem 2014; 61(2): 101-10.
[http://dx.doi.org/10.1002/bab.1148] [PMID: 24033726]
, 119Meng N, Zhou N-L, Zhang S-Q, Shen J. Synthesis and antimicrobial activities of polymer/montmorillonite–chlorhexidine acetate nanocomposite films. Appl Clay Sci 2009; 42(3): 667-70.
[http://dx.doi.org/10.1016/j.clay.2008.06.016]
, 120Czaja WK, Young DJ, Kawecki M, Brown RM Jr. The future prospects of microbial cellulose in biomedical applications. Biomacromolecules 2007; 8(1): 1-12.
[http://dx.doi.org/10.1021/bm060620d] [PMID: 17206781]
].

CONCLUSION

Many challenges need to be addressed in BC research, such as introducing low-cost medium and efficient producing system with low capital investment. Besides many applications of BC and BC-composites, the need to study bacterial diversity is necessary. Furthermore, the study of 3-D printing of BC for food and food packages with specific geometry can provide more information in this field. Additionally, BC producers' diversity and molecular characterization can give great insight on low cost production of BC, especially for food and biomedical products. Last but not least, engineering the BC producers towards specific functionalization can answer many industrial and biomedical needs. Tunable control on BC production and novel structural pattern can be important for BC producer bacteria's future engineering.

CONSENT FOR PUBLICATION

Not applicable.

FUNDING

This work was financially supported by the Institute For Research and Development, Suan Sunandha Rajabhat University (SSRU) (Grant number 441/2563).

CONFLICT OF INTEREST

The authors declare no conflict of interest, financial or otherwise.

ACKNOWLEDGEMENTS

The authors would like to thank Dr. Anat Thapinta, Dean of the Faculty of Science and Technology, SSRU and Dr. Suwaree Yordchim, Director of the Institute of Research and Development, SSRU for their outstanding support. Wirongrong Thamyo is acknowledged for excellent secretarial assistance.

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(National Central University, Taiwan)


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