Why PDB? PDB builds introductory materials to help beginners get started in the subject "", as in an entry level course as well as resources for extended learning. Toggle navigation PDB Educational portal of. Molecule of the Month. Diabetes is a worldwide health problem affecting hundreds of millions of people. Fortunately, with careful management of diet and medication, the many complications of diabetes can be reduced. Part of this treatment includes the monitoring of glucose levels in the blood, so that proper action may be taken if levels get too high.
The enzyme glucose oxidase has made glucose measurement fast, easy, and inexpensive. Glucose oxidase, shown here from PDB entry 1gpe , is a small, stable enzyme that oxidizes glucose into glucolactone, converting oxygen into hydrogen peroxide in the process. Its normal biological function appears to be centered on the peroxide that is formed: hydrogen peroxide is a toxic compound that can be used to kill bacteria. For instance, glucose oxidase is found on the surface of fungi, where it helps protect against bacterial infection, and it is also found in honey, where it acts as a natural preservative.
It is used as the heart of biosensors that measure the amount of glucose in blood. The trick to these biosensors is that the enzyme takes something that is difficult to measure--glucose--and creates something that is easy to measure--hydrogen peroxide. A typical laboratory glucose meter includes some of the enzyme trapped inside a membrane.
These studies illustrated the high potential for enhanced production of GOD in yeasts. Although reports have confirmed similar expression properties of recombinant enzymes to the native ones, their industrial applications still bear several limiting factors, such as their higher cost and deficient reconstitution. GOD secreted by yeast shows overexpression, which facilitates higher production and purification and helps in enzyme modification by random mutagenesis. A number of methods have been employed for the initial screening of GOD mutant lines using fungal cultures of the isolated clones Zhu et al.
Using an expression system of S. Enzyme engineering using currently available methods such as rational design, rational redesign, and directed evolution can lessen or remove the limitations to the improvement of GOD variants.
Rational redesign strategies were recently used to improve the catalytic function and stability of GOD. Holland et al. Fisher et al. Recently, Song et al. The results obtained were consistent with those of the computer-aided analysis, suggesting that this method may be useful for enzyme structure optimization. The directed protein evolution has recently come into use in the molecular modification of GOD, and it is also expected to improve the yield of GOD.
Zhu et al. Ostafe et al. Similarly, Prodanovic et al. Recently, Horaguchi et al. Their results revealed structural motifs of the protein are critical to its stability.
Although fungal sources offer a wide range of enzymatic properties, new features such as marked product inhibition, higher product yields, or secretion signal could be designed into specific GOD using innovative tools of state-of-the-art protein engineering. The high-level progress of GOD production through the engineering of strains was recently reviewed by Liu and Piwu Similarly, Suraniti et al.
Optimization of GOD can be achieved by employing modern molecular level tools and techniques in association with bio-process engineering technologies that may result in economically feasible enzyme production system. Application of efficient recombinant microbial technologies on new resources of GOD and protein engineering technology have proved significant efficacy to convert GOD into a relevant synthetic tool.
Thermostability of GOD may further enhance its industrial value in addition to its boosting functional food market. GOD is an efficient oxidant for the production of bread with improved quality and extended loaf volume in the baking industry Rasiah et al.
In addition, Vemulapalli and Hoseney reported the drying effects of GOD on the dough, which were mediated by the gel-forming ability of water-soluble pentosans, reduction in sulfhydryl content, and the increase of viscosity in the water-soluble dough. Also, GOD has been found to display protein cross-linking in the dough Rasiah et al. GOD improves the quality of bread and strengthening of wheat dough when used as an additive Bonet et al. GOD also enhances the viscoelasticity of dough Kouassi-Koffi et al.
Specifically, Kouassi-Koffi et al. However, enzymes must be added with care since undesirable effects can be caused by excessive enzymes. GOD, along with lipase also enhances the quality and shelf-life of pan bread El-Rashidy et al. Dagdelen and Gocmen studied the effects of GOD along with ascorbic acid and hemicellulase on bread quality and dough rheology and found that bread quality is mainly dependent on original wheat flour quality, while dough rheology depended on the amount of enzyme.
Kerman et al. Similarly, da Silva et al. Decamps et al. Aprodu and Banu studied the effects of Psyllium , pea fiber, oat bran, water, and GOD on rheology and baking properties of gluten-free bread made from maize, and it was suggested that GOD has significant ability to improve the specific bread volume for all types of fibers.
GOD plays a novel role in the manufacturing of beverages because it is used to diminish the low alcohol substances of wine by eliminating the residual glucose that would otherwise be converted into alcohol through anaerobic fermentative processes.
Moreover, the H 2 O 2 produced during chemical processes imparts a bactericidal impact on acidic corrosive and lactic corrosive microbes amid the fermentative procedures. This process must be conducted by adding GOD prior to fermentation as GOD utilizes a portion of the glucose presents, making it inaccessible for liquor aging, bringing about wine with decreased alcohols and simultaneous generation of H 2 O 2 that reduces the growth of fermentative microorganisms.
The H 2 O 2 produced could easily be removed from the system using CAT, which breaks it into oxygen and water. The bactericidal effect of H 2 O 2 reduces the addition of other chemical preservatives in the wine Malherbe et al. GOD has shown significant efficacy on determining glucose content in body fluid and effectively removes oxygen and residual glucose from beverages Yildiz et al.
Lopes et al. Further, Mason et al. GOD has been used effectively to remove remaining glucose and oxygen from foods to extend their shelf-life Zia et al. The reaction of protein amino group and reducing sugars is known as non-enzymatic Maillard browning, which results in the formation of unwanted flavor and undesirable browning in dried egg powder, suggesting prior removal of glucose content from the liquid egg before its drying Sisak et al.
Removal of glucose provides dried egg powder a prolonged shelf-life and increased microbial tolerance. Also, production of H 2 O 2 by Maillard reaction helps to destroy unwanted microbes normally found in liquid egg Dobbenie et al. Increasing browning mediated by the Maillard reaction is also a noteworthy aspect causing harmful effects to eggs and potato products.
Application of GOD may provide sustainable results to reduce unwanted browning in the same manner Low et al. One of the major applications of the GOD catalyzed reaction is the production of GA and its derivative salts. GA was found to play extensive roles in different sectors of food industries and utilized as a causticity controller, raising specialist, color stabilizer, an antioxidant and chelating operator in bread, feeds, beverages, and so on Brookes et al. In dairy industries, GA is used for the cheese curd formation, improvement of heat stability of milk, prevention of milk stone, and cleaning of aluminum cans.
However, GA is most widely used as acidulant, sequestrant as well as potential anti-oxidant in various industries Golikova et al. In the pharmaceutical based industries, the metal derived Na, Ca, Zn, and Fe salts of GA is widely used in the synthesis of important drugs including sodium, calcium or ferrum gluconates, and glucono-delta-lactone which have diverse industrial applications Ramachandran et al.
Sodium gluconate has great potential to chelate metal ions and can be used to remove bitterness from food stuff Costa et al. Pharmaceutical application of calcium gluconate has been confirmed for the treatment of calcium associated deficiencies Khurshid et al. GA can be produced through biochemical, electrochemical, bioelectrochemical, and fermentative processes, although fermentative processes are preferred for GA production as other approaches are expensive and less productive Wong et al.
It has been reported that GA production through GOD catalyzed reaction is highly dependent on the substrates used, oxygen concentration and temperature Ramachandran et al. Moreover, the catalytic efficiency for the conversion of glucose to GA is highly dependent on the stability of GOD.
Several studies have been conducted to optimize the production of GA. For example, Purane et al. Ping et al. Further, Ramezani et al. It was found that the increased oxygen gas velocity resulted in the increasing rate of glucose oxidation reaction because of the higher transformation of oxygen from a gas to a liquid state.
Optimization of mass transfer characteristics and operating condition during GA production with immobilized GOD, it was found that a bubble-column reactor had better mass transfer properties because it provided higher GA production under low GOD activity relative to other reactors. Recently, many investigations have investigated GA production using substrates through multi-enzymatic steps. Mafra et al. Similarly, Silva et al. GA production through fermentative processes also depends on morphological parameters of the fermenting organism used.
Indeed, it was reported that the dispersed pattern of the mycelial morphology of A. GA production through fermentative processes is the most widely accepted technique and the most common challenge to these methods is downstream processing separation and purification.
However, these hurdles can be resolved by membrane-based separation Pal et al. Many food products contain oxygen, which promotes bacterial growth. Karimi et al. Moreover, GOD could be utilized for the removal of oxygen from the top of bottled beverages such as wine and beer to maintain the taste and flavor Labuza and Breene, ; Wong et al.
The overall GOD catalytic reaction consumes two glucose particles and an oxygen molecule, resulting in the production of two GA molecules. During the reaction, the consumption of oxygen allows GOD to be used as a strong antioxidant and scavenger of oxygen, thus facilitating its application as a food preservative due to stabilizing effect.
Additionally, GOD has been effectively used in the food system as a strong stabilizer due to its oxygen removing ability and prevents color and flavor loss in a variety of beverages including canned fish, beer, soft and energetic drinks Crueger and Crueger, ; Bhat et al.
GOD can also be used instead of potassium bromate as an oxidizing agent in bread making Moore and Chen, Streptococcus mutans , which inhabits the oral cavity and causes tooth-decay, is carried by almost every human being. The ability of GOD to kill S. Hill et al. Senol et al. GOD has antimicrobial activity against different foodborne pathogens. GOD has shown enormous potential to inhibit the growth of various foodborne pathogens, including Clostridium perfringens, Campylobacter jejuni, Salmonella infantis, Staphylococcus aureus , and Listeria monocytogenes Tiina and Sandholm, ; Kapat et al.
Further, GOD covalently immobilized on biorientated polypropylene films was found to inhibit the growth of E. Murray et al. Malherbe et al. Zia et al. GOD produced by P. Application of edible antimicrobial films has been approved to enhance the shelf-life of food products by releasing enough amount of antimicrobial substances on the surface of food products.
The polyamide and ionomer films with immobilized GOD inhibited the growth of bacteria such as E. Recently, a new photo-dynamic glucose-based antimicrobial system encapsulating GOD, HRP, and BRET bioluminescence resonance energy transfer was developed for the inactivation of various bacterial and fungal pathogens through the network of organic and inorganic materials Yuan et al.
Biosensors are widely used in the food industry, monitoring of environmental hazards, and clinical applications. GOD has been widely employed in glucose-based biosensors because of its high selectivity for glucose and functionality under extreme temperature, pH, and ionic resistance. Glucose biosensors for diabetic blood monitoring are very convenient, reliable, rapid, and accurate. Many studies have been conducted to develop sophisticated advanced technologies, and better alternatives such as point sample tests, and the continuous glucose monitor CGM are being developed Wang and Lee, ; Sode et al.
The CGM sensor has shown a significant role in diabetes as an alternative means while measuring blood glucose level and provides an alarming node on events associated with blood glucose metabolism Wang and Lee, Further, in recent years, the drawbacks and limitations associated with glucose biosensors have been nullified using advanced approaches such as electrodes, membranes, enzyme immobilization, and nano-composite film modified electrodes.
Amperometric glucose biosensors can be divided into three generations based on their operative principles. The chemical reaction leading to oxidation of glucose causes depletion of oxygen or production of H 2 O 2. The involvement of other redox species was the major problem associated with this generation. In second generation biosensors, mediators are involved in the backward and forward flux of electrons between the enzyme and electrodes, but this generation of biosensors had a low turnover rate and reduced proximity between the electrodes.
Hence, these biosensors suffered from redox interferences mostly oxygen. The direct electron transfer process could be achieved through several modifications and the biosensors developed based on such modified electrodes have been found to have good reproducibility, selectivity, and stability for glucose oxidations.
Velmurugan et al. The method of immobilization of GOD is considered a pivotal factor for the development of highly stable glucose biosensors with long-term operational life.
The various methods by which GOD is incorporated into a biosensor include absorption, covalent attachment, cross-linking and micro-encapsulation inert membrane entrapping enzyme into the transducer surface.
Moreover, such immobilization could be achieved through one or in combination with others. Hong et al. One of the most important aspects of such GOD based immobilization is the greater loading of enzymes for the efficient functioning which makes the overall system stable and selective in the form of cross-linking precipitated GOD aggregates.
It has been reported that the problem of enzyme absorption on GOD based biosensors could be improved by using silicalite modified electrodes. Recently, the uses of carbon nano-chips CNCs have been used to modify the glassy carbon electrode for immobilizing GOD with the help of chitosan. The properties of the immobilized enzyme in biosensors depend on both the enzyme and the supportive material involved Ang et al. The two major limitations that restrict the immobilization of GOD on solid electrodes include inadequate electrical communications between the active sites of GOD and the surface of electrodes including enzyme leaching.
A huge range of electrode substrates has been used recently to overcome these problems which include metal-based nano-particles, carbon nano-tubes CNTs , mesoporous silica, polymers, and sol-gels Table 3.
Apart from these methods, the use of CNTs is considered a notable advancement in biosensing for the construction of glucose sensors due to their ability to promote the reactions of electron transfer of biologically significant biomolecules. Low-site-density based nano-electrodes aligning CNTs have been used for the detection of glucose Lin et al. The use of nano-materials in biosensors and bioelectronic devices has provided a new platform for efficient glucose monitoring due to their improved response time, high sensitivities, low detection limits, wide range linearity, and low power requirements.
A list of major nano-materials, CNTs, and carbon nano-fibers CNFs used as electrical connectors between the electrode and the redox center, are given in Table 3. Wang X. GOD also plays an important role in the induction of defense responses in plants. Involvement of H 2 O 2 during the plant resistance to bacterial disease agent was also revealed in Arabidopsis plant challenged by transconjugants of Pseudomonas syringae pv. The application of genetic engineering tools has provided sustainable results in GOD expression in plants with increasing resistance to plants from bacterial infections Wu et al.
Further, Maruthasalam et al. Endogenous H 2 O 2 levels of tobacco plants Nicotiana tobaccum L. Similarly, the GOD gene from A. Kachroo et al. Moreover, GOD isolated from A. Hence, A. Bleaching provides decolorization of natural pigments with a pure white appearance of the fibers. GOD has proven to be effective in the production of H 2 O 2 for bleaching in the textile industry, the most effective bleaching agent of industrial significance Bankar et al.
Moreover, the GOD application to the bleaching of textiles during upstream resizing and bio-scouring processes has shown promising results with the additional release of glucose Buschle-Diller et al. The immobilization of GOD enzyme for the generation of H 2 O 2 and its optimization has significantly affected the processing of bleaching in textiles. Tzanov et al. The stability of GOD has been enhanced by a few immobilization procedures on different backings Quinto et al. Recently, Aber et al.
Farooq et al. Furthermore, H 2 O 2 produced by GOD was shown to be a significant alternative to the most extensively used commercial H 2 O 2 , in the textile processing industries. Moreover, Tzanov et al. However, the initial high concentration of glucose may cause discoloration of fabrics due to the presence of residual glucose. This problem can be overcome by using an excessive amount of GOD with an increased incubation time Saravanan et al.
Other important aspects of enzymatic processing in textiles are that the H 2 O 2 generated during bleaching produces a comparable effect to scoured woven cotton fabric, while the GA produced acts as a chelator for metal ions, removing the need for use of an additional stabilizing agent Tzanov et al.
Additionally, the simultaneous application of GOD with peroxidases in the decoloration process improves bleaching of natural fibers Opwis et al. Biofuel cells BFCs use either enzymes or whole cell organisms as a biocatalyst to generate power directly from fuel substrates glucose and ethanol.
These enzyme-based systems are considered better alternatives for the development of future implantable devices Sode et al. The design for manufacturing BFC is adjusted in such a way so that one electrode consisting of electro-conductive material is modified by a biocatalyst enzymes for specialized oxidation and reduction reactions.
More recently, efforts have been put forward to improve the catalytic efficiency of this system several folds by co-immobilization of GOD with other enzymes such as CAT. Christwardana et al. Figure 1. Generalized diagram of typical enzymatic biofuel cell EBC with associated components. Most of the output voltage and current signals in a typical BFC depends on the concentration of fuel, hence, enzyme-based BFCs could serve as an alternative means of an enzyme sensor system Katz et al. Recently, there has been a great deal of efforts toward the development of bio-electrochemical devices based on unique enzymes.
The presence of electron transfer sub-units or domain in these enzymes imparts specificity to directly transfer electrons to the electrodes during a bio-catalytic reaction Tsugawa et al. The sensitivity and conversion efficiency of BFCs are highly determined by efficient electron transfer occurring at the enzyme active center and electrode interface. However, these critical factors of sensitivity and conversion efficiency lead to difficulty in GOD catalysis based BFCs because the redox center inside this enzyme is buried inside the structure, a long way from any feasible electrode binding site Sode et al.
This problem can be resolved by using artificial electron acceptors and mediators or by precisely modifying the electrode surface with nano-scale conductive materials. These mediators can be polymerized specifically onto the surface of electrodes or co-immobilized with GOD to facilitate the rate of electron transfer by several-fold.
Recently, a novel method entrapping cross-linked aggregates of GOD within a graphitized mesoporous carbon GMC network has been reported for the production of GOD nano-composites with an ability to provide the maximum rate of electron transfer and high electrical conductivity Garcia-Perez et al.
Nano-carbon functionalization has shown perfect compatibility with other biological and chemical approaches with enhanced enzymatic functionality in implanted BFCs.
Babadi et al. Some recent approaches to the development of enzyme-based BFCs are listed in Table 2. Figure 2. Generalized diagram of an enzymatic biofuel cell EBC with a mediator bioanode and direct electron transfer DET based biocathode mediators are involved in fluxing electron flow between enzyme and electrode.
The use of immobilization facilitates retention of biomass in reactor geometry, enabling their economic reuse and development of the continuous process. The technique also improves stability and prevents product contamination, paving the way for use of crude enzyme preparations such as whole cells in the bioprocessing D'Souza, Therefore, bio-capacitors have been developed using charge pumps connected to fuel cells.
This novel approach has generated high voltage with sufficient temporary currents to operate an electric device without changing the design and construction of the EFC Sode et al. The limited life expectancy of BFCs could be enhanced using technologies that favor enzyme stability, whereas power supply could be resolved by improving catalytic efficiency using directed evolution or other protein engineering methods.
In addition to the above-mentioned benefits, GOD could be used in some miscellaneous applications. For example, GOD has been used in several immunoassays and staining techniques and shown to have an ability on the removal of an excess level of glucose Megazyme, In medical sectors, GOD is widely employed in the monitoring of diabetic patients to measure their blood glucose activities using finger-prick blood samples.
In addition, GOD induces several apoptosis characteristics viz, mitochondrial dysfunction, accumulation of Bax and release of cytochrome C in mitochondria, accompanied by activation of caspase-9 and caspase-3 Rost et al. Moreover, GOD was also reported to induce cellular senescence in immortal renal cells through integrin-linked kinase ILK by downregulating Klotho gene expression, an aging- related kidney-secreted hormone with antioxidant properties Yamamoto et al.
Apart from these applications, it has been reported that the enzymatic activity of the white rot fungi P. Wang C. The visual CL bio-platform showed outstanding performance with high selectivity, a wide linear range and a low detection limit for sensing trace glucose. Recently, Nascimento et al. These nano-pipettes functioned as specifically developed nano-sensors to measure the glucose level in single cells with temporal and spatial resolutions.
The covalently immobilized GOD at the sensor tip interacted with glucose forming GA, which was measured as the change in impedance due to the drop in the pH. Bandodkar et al. Diets supplemented with GOD have been reported to enhance growth performance, increase the contents of growth and development-related hormones and improve the fecal microflora of growing piglets Tang et al.
Using a similar approach, Tang et al. However, more research is needed to determine the various effects of dietary GOD supplementation on other parameters of intestinal health.
Moreover, during localization microscopic studies, GOD was used in single molecule localization microscopy SMLM buffers to decrease the solution oxygenation as well as to prevent fluorophore photobleaching Szczurek et al. The above review explains the potential applications of GOD in various industries with increasing demand in the food and flavoring, pharmaceutical, biotechnology and bioelectronic sectors. With a predicted annual growth rate of 7.
Although a plethora of microbial resources are currently available for efficient production of this enzyme, only a small fraction of microbial entities, particularly some selected strains of fungi and yeast, is currently exploited for the production of the enzyme.
Further, huge attention should be paid on finding new sources of GOD and to develop cost effective fermentative processes for the efficient production and commercial exploitation of the enzyme.
However, we currently do not have sufficient information available regarding the commercial production of GOD through fermentative processes and its further subsequent uses in different industries.
Recently, enzyme immobilization processes and other technological innovative approaches have drawn scientific attention because of the potential for developing novel methods for the GOD production at large scale. However, problems such as diffusional constraints and decreased enzyme activity after immobilization have further limited the uses of immobilized GOD and therefore need to be improved to achieve greater benefits. The recent advancements in the field of bioinformatics can also revolutionized the field of GOD immobilization through docking and molecular dynamics techniques which can provide detailed information about the enzyme and ligands interaction.
Large scale and efficient uses of GOD in various industrial sectors could be achieved through the use of modern biotechnological approaches such as concoction adjustment of existing chemicals through protein designing, site-directed mutagenesis, and recombinant expression of GOD genes in other potential microbes that could be further used at large scale to meet future demands.
Fortunately, the different recent trends as mentioned throughout this review suggest that we are on the path of establishing a worldwide bio-based economy, and GOD may have a great contribution in this context. MD provided the general concept, and drafted part of the manuscript.
All authors revised and approved it for publication. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Abbasi, M. Improvement of the stability and activity of immobilized glucose oxidase on modified iron oxide magnetic nanoparticles. Aber, S. Immobilization of glucose oxidase on Fe 3 O 4 magnetic nanoparticles and its application in the removal of acid yellow Water Air Soil Pollut.
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Sugar and Tooth Decay. Virtual ChemBook. Blood Glucose Test With a couple of million people in the U. Click for larger image. Oxidation of Glucose: Blood glucose levels are now measured by a procedure based upon the enzyme glucose oxidase. Color Producing Reaction: Several methods of detection are possible, but in most cases, the hydrogen peroxide reacts with a second color producing chemical.
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