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Biotechnological implications of hydrolytic enzymes from marine microbes
Chapter 7
Biotechnological implications of hydrolytic enzymes from marine microbes Poonam Vashist1, R. Kanchana2, Vazhakatt Lilly Anne Devasia1, #, Priyanka V. Shirodkar1, Usha D. Muraleedharan1, * 1
Department of Biotechnology, Goa University, Goa, India; 2Department of Biotechnology, Parvatibai Chowgule College of Arts and Science -Autonomous, Margao, Goa, India; # Present address: Department of Biotechnology, Hindustan College of Arts and Science, Padur, Kelambakkam, Chennai, India * Corresponding author: [email protected].
7.1 Introduction Biocatalyst and biotechnology are intervolved terms that have camouflaged almost every socioenvironmental activity of our living style in the 21st century. To date, almost 4000 enzymes are known, out of which approximately 5% of microbial original types are used commercially [1]. The marine microbial population, while still largely unexplored because of its diversities and extreme environment, is being screened with the help of high-throughput technologies such as shotgun sequencing or pyrosequencing for deeper exploration [2], resulting in the reporting of approximately 20,000 species per liter of marine water samples [3]. Microbial enzymes have proved superior over chemical catalysts and enzymes derived from plants or animals, by virtue of their versatility, stability, enantioselectivity, reduced process time, intake of low-energy input, and costeffectiveness, besides their nontoxic and eco-friendly characteristics [4]. Competition amongst microorganisms for space and nutrients in the marine environment is a powerful selective force that has led to the generation of multifarious enzyme systems to adapt to the complicated environments. Many of them are thus endowed with desirable features, from a general biotechnological perspective. However, these features of microbes as well as enzymes can be enhanced with the help of techno-studies such as recombinant DNA/ protein engineering [5].
Advances in Biological Science Research. https://doi.org/10.1016/B978-0-12-817497-5.00007-0 Copyright © 2019 Elsevier Inc. All rights reserved.
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Marine microorganisms, whose genetic and biochemical diversity studies are not quite out of their infancy, are of significant interest as a potential source for novel applications [6,7]. Researchers the world over have successfully isolated and characterized a variety of enzymes with novel activities from marine bacteria, actinomycetes, fungi, and other marine microorganisms, of which a few products have already found large-scale industrial applications [8e11]. Extremophilic marine microorganisms living in hydrothermal vents, sea floors, salt pans, and hot springs have been found to be rich sources of extremozymes such as proteases, cellulases, amylases, and lipases [12]. Our laboratory has had excellent success with bioprospecting of marine enzymes from several such sources. Vashist et al. reported a novel strain of Microbulbifer sp. isolated from a mangrove area in Divar Island, Goa, India, which exhibited more than 13 ICP degrading activities [13]. Another important avenue of interest has been the marine biodegraders classed as thraustochytrids. Kanchana et al. [14] have reported alkaline lipase activity with potential biotechnological application from these marine protists. Similarly, Devasia & Muraleedharan [15] have reported multiple polysaccharide-degrading activities from thraustochytrids for the first time. Employing response surface methodology, Shirodkar & Muraleedharan have successfully optimized a medium for commendable production of amylolytic activities from a thraustochytrid strain [16,17]. Almost 75% of all industrially applied enzymes are of the hydrolytic category, out of which proteases, carbohydrases, and lipases lead the enzyme market, with more than 70% of all enzyme sales [1]. Among these, hydrolytic enzymes resourced from bacteria hold the majority share when compared to the overall bioactive compounds discovered from marine fungi and other domains [18]. The world of enzyme demand is satisfied by about 12 major producers and 400 minor suppliers [19]. Three top companies, namely, Denmark-based Novozymes and Danisco and Switzerland-based Roche satisfy about 75% of the total enzyme demand [1]. While the world market for industrial enzymes was to the tune of $4.2 billion in 2014, an expansion at a compound annual growth rate (CAGR) of approximately 7% is predicted over the next 5-year period, escalating the figure to nearly $6.2 billion [20]. In this chapter, attention is drawn to the significant role of hydrolytic enzymes from marine microbes involved in various fields of technical applications. Also reported are aspects of modern biotechnology for improved enzyme production, enabling various industrial processes at lower energy consumption and higher efficiency, besides enhancing properties of the product while ensuring environment friendliness.
7.2 Applications of marine hydrolases Hydrolases catalyze several reactions, including condensations, alcoholysis, and so on. The main advantages of this enzyme class are ready availability, lack of cofactor stereoselectivity, and tolerance to the addition of water-miscible solvents. In fact, the majority (about 75%) of currently used microbial enzymes in various industries fall into the hydrolytic category [21].
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7.2.1 Biorefineries Over several years, studies have been in progress in different parts of the world to find substitutes for the fractions of nonrenewable/fossil resources in use. Brazil, the United States, China, and India, covering the most-populated regions of the world at 44% of the total global population, use 45% of the total energy sources and account for 51% of the carbon dioxide release from the consumption of energy. Notwithstanding this, about 27% of primary energy is used worldwide for transportation, thus highlighting transportation fuels as notable targets for replacement with a renewable source [22]. Consistent and harmonized supply of raw materials is the most important factor in biorefinery productivity [23]. The use of aquaculture biomass (seaweeds) as a feed for biorefinery industries ensures a continuous, rapid, safer, and economic supply of feed for biorefinery industries compared to the use of lignocellulosic biomass from agricultural wastes and other biological materials. Algae have 20e80% oil content, most of which are triacylglycerols, but the robust algal cell walls present a challenge for the extraction of these lipids. Enzymatic lipid extraction has beencarried out withsnailase along with cellulase and lysozyme [24]. Novel and significant enzymatic properties of extremophiles and thermostable enzymes have boosted the biocatalysis in biofuel production [25]. In biorefineries for bioethanol, biogas, and biodiesel production, cellulases, hemicellulases, and ligninases are most widely used, primarily during pretreatment and saccharification of biomass. Some bacteria and fungi are capable of rapid and efficient degradation of cellulose [25,26]. Enhanced depolymerization of the cellulose fraction from lignocellulosic substrates would be favored by the use of a cocktail of enzymes such as exoglucanase, endoglucanase, b-glucosidase, cellobiase, xylanase, and other hemicellulases [27].
7.2.2 Pharmaceuticals and cosmeceuticals Enzymes play various vital roles in the realms of pharmaceuticals, cosmetics, and diagnostics. The pharmaceutical industry in particular requires a very high degree of substrate specificity on account of the stringent routes for desired product synthesis. Coupled with the effective processing cost, this propels enzymes to advantage over chemical catalysts [4]. Enzymes are extensively being used as large-scale therapeutic drugs mainly pertaining to health concerns associated with enzymatic deficiency, digestive disorders, and in ELISA-based diagnostics [28]. Marine natural products play an important role in drug development [29]. The enzymes involved in mycothiol biosynthesis and found in the Actinobacteria, Actinomycetes and Mycobacterium are of great interest as potential targets for the production of new drugs against Mycobacterium tuberculosis [30]. The applications of hydrolytic enzymes in medicine are growing rapidly, as in the removal of dead skin, and enzymes such as Vibrilase (vibriolysin)
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obtained from the marine microorganism Vibrio proteolyticus have been reported for effectiveness against denatured proteins found in burnt skin [31]. Lipases have been cited as the most repeatedly used enzymes in the synthesis of alcohols, acids, lactones, and esters that are optically active [32, 33]. On account of their digestive and inflammatory properties, proteases have been testified for a wide array of applications in medicine [34]. Chitosanase catalyzes the hydrolysis of chitosan, which is used as an antimicrobial, antioxidant, and in bringing down high blood cholesterol and blood pressure levels. It has also found use in the control of arthritis, besides enhancing antitumor properties and according protection against infections [35]. Actinobacteria are considered the most important group of organisms being studied extensively for the discovery and formulations of significant drugs and other bioactive metabolites with high specificity [36]. Glucanase isolated from a marine Bacillus has been reported suitable for oral and other health care [37]. Oligomers obtained from carrageenan degradation have demonstrated antitumor, antiviral, anticoagulative, and other activities such as boosting immune function and digestion [38e40]. Qiu et al. reported significant advantages of the marine bacterial protease from an N1-35 strain isolated after UV mutagenesis over those from terrestrial ones [41]. Enzymes such as urease and creatinase from Actinobacteria have been used in clinical diagnostics for quantitative evaluation of diabetes and other health disorders [42,43]. Enzymes have gained increasing popularity in the cosmetic industry by virtue of their relative safety and minimal side effects. Li et al. [1] have reported the use of enzymes as free radical scavengers in preparations for hair styling and dyeing, and in toothpastes, mouthwashes, and sunscreen products. In Japan, agar-oligosaccharide is being extensively used as a moisturizing cosmetic additive and and has also been reported for good hair conditioning effects [44]. Proteases have been used in skin creams to remove dead cells and thereby effect cleaning and smoothening of the skin [45]. Papain and endoglycosidase have found wide use in toothpastes and mouthwashes, serving to whiten and remove plaque as well as other odor-causing deposits on teeth and gum tissue [46]. Lipase and papain are used in skin care products [47]. Enzymes are also routinely being incorporated in contact lens cleaners to remove proteinaceous films [48].
7.2.3 Food industry The present rate of world average for per capita food consumption is 3000 kcal/day [19]. The estimated food consumption is expected to go up to 22.8 1012 kcal/day by 2020; when compared to the energy, it is roughly 16 million barrels of oil equivalent (boe) per day i.e., an approximate 70% increase by 2050 [19]. Meeting these requirements in time without compromising on food quality could be managed by enzymes application. These biocatalysts have been reported as well as used efficiently for improving food
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production, quality and components such as flavor, aroma, color, texture, appearance, fat modification, and in sweetener technology [1]. Application of enzymes in the food industry is principally for improving the shelf life of products, besides their stability and texture. Agar oligosaccharides produced by the action of agarases are being used for the production of beverages, bread, and some low-calorie food [49,50]. Isolates such as Microbulbifer CMC-5, 10A, and LK2 have been reported for promising and extensive agarolytic activity that can be further exploited [51,52]. Agarose-derived neoagarooligosaccharides produced by b-agarase from the seaweed-decomposing bacterium Microbulbifer sp. strain CMC-5 showed a promising biotechnological application [52]. Similarly, esterases and lipases have been reported for betterment of bread quality by change in lipids in the dough used, in butter, cheese, and margarine flavor enhancement, in the production of crackers, pasta, etc., degumming of vegetable oils, synthesis of infant formula and nutraceuticals, increasing the concentration of polyunsaturated fatty acids in vegetable oils, as well as for improving the digestibility of natural lipids [53]. Recently Li et al. isolated a novel esterase of the hormone-sensitive lipase family, from a metagenomic library of more than 10,500 cosmid clones from the South China Sea [54]. A cold-active lipase from the marine bacterium Janibacter sp. strain HTCC2649 was isolated and characterized by Yuan and coworkers with sn-1/3 specificity toward monoand di- acylglycerols [55]. Recent reviews have endorsed the application of xylanases, lipases, and amylases in the baking industry, for improving freshness and shelf life of the products, elasticity of gluten, enhancing handling, and increasing stability of the dough, as well as improved bread volumes and absorption/redistribution of water and flavor; higher content of arabinoxylooligosaccharides in bread with a positive effect on health, lighter cream crackers, improved texture, palatability, and uniformity of wafers are other benefits that have been derived [1,56,57]. Cheese, yogurt, and other milk products have been improved upon with respect to organoleptic characteristics such as aroma, flavor, color, and yield [58]. The use of enzymes (protease, catalase, lipase, esterase, lactase, etc.) in the dairy market is well recognized for reduction of allergenic properties of milk products [59]. The effectiveness of collagenase and protease has been proven in meat tenderization and further processing and protease MCP-01, an abundant extracellular serine protease produced by the deep-sea psychrophilic Pseudoalteromonas sp. SM9913, is reported to be one such psychrotolerant enzyme [60]. Damare et al. have described a detergent-compatible alkaline, coldtolerant protease from a deep-sea fungus from the Central Indian Basin [61]. Enzymes have been employed in the beverage industry to control the brewing process, and to produce beer of consistently high quality. Improvement of the juice yield, aroma, and color are other positive outcomes.
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Such enzymes also serve to enhance the nutritional and functional characteristics of proteins of animal and vegetable origin that are used in the process [62]. The first stage in a brewing industry for extraction of material would be to digest the plant cell wall using microbial enzymes [63]. Inulinase form Pichia guilliermondii OUC1 has been reported for the production of ultrahigh-fructose syrups, while Yarrowia lipolytica OUC2 could be used for the production of inulooligosaccharides [64,65]. It has been reported very effective to use amylases, cellulases, arabinases, naringinase, pectinases, and xylanases during fruit juice/beverage processing to enhance yield, in juice clarification, for improved extraction, stability, texture, sensory characteristics, and cost/time management [1,66,67]. Tannases from Aspergillus awamori BTMFW032 have been used for producing coffeeflavored soft drinks and instant tea, for the clarification of fruit juice and beer, and for food detannification [68]. Many reports describe b-glucosidases as widely used during processing of fruit juices, tea, wine, and beer to effect enhancement of their organoleptic properties and essential oil content [69e72]. One specific study on b-glucosidases from Thermotoga maritime has been documented by Goyal et al. [73].
7.2.4 Feed industry The continuously increasing trend in worldwide milk and meat consumption invariably escalates the demand for the feedstock for animals, and to meet the requirements of quantity and quality, the search for feed enzymes for diet formulations necessarily had to pick up the pace. Enzymes incorporated in animal feeds aid to increase the digestibility of nutrients and ensure better feed utilization [74]. The practice of supplementation of animal diets with enzymes increased exponentially from the 1980s to the subsequent decades. The world market for feed enzymes, assessed at $899.19 million in 2014, is estimated to touch the $1.3 billion mark by 2020, at a CAGR of 7.3% over the period [1]. The protein concentration in poultry feeds has been enhanced by the application of feed enzymes such as a-galactosidases, phytases, a-amylases, proteases, glucanases, xylanases, and polygalacturonases [1]. Faced with the inability of monogastric animals to digest plant-based feeds rich in hemicelluloses and cellulose, xylanase and b-glucanase have been added to the feeds to enable total degradation and digestion [75,76]. To the same end, b-glucosidases have been documented as favored additives in cellulose-based feeds for animals such as chicken and pigs [77].
7.2.5 Biopolymer industry Involving microbial enzymes in the synthesis of biodegradable polymers appears the best solution to circumvent the increased requirement of polymers that have proved detrimental to human health and environment. The applications of biopolymers such as polyesters, polycarbonates, and polyphosphates have been
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reported for various biomedical purposes, e.g., orthopedic devices, tissue engineering, adhesion barriers, and controlled drug delivery [78]. The biopolymer market, which had grown substantially over the recent years, was expected to touch $3.6 billion by the end of 2018 [1]. The ever-increasing clamor for packaging materials and the concomitant environmental safety concerns could both be addressed by the flourishing biodegradable polymer industry. Lipases and laccases catalyze polymerization reactions to produce polyesters or polycarbonates and cross-links in biopolymers [79,80]. Y. lipolytica lipase when customized by immobilization and protein engineering has been shown to improve the performance in biopolymer synthesis [81].
7.2.6 Detergent industry Successful integration of hydrolytic enzymes into detergent formulations is cited as the key to the production of cost-effective and environmentally gentle detergents [82]. In today’s laundry detergents, enzymes such as proteases and amylases are deeply entrenched among the active ingredients. A recipe of alkaline proteases and cellulases isolated from microbial sources and that can effectively remove protein stains as well as cleave damaged cotton fibers is now a vital component of various detergents produced on a commercial scale by manufacturers such as Novozymes SA, Kao Corporation, and Genencor International [83]. Greene and his colleagues had reported cleansing properties of an alkaline protease isolated from a bacterial strain symbiotic with the gland of Deshayes of a marine shipworm [84]. Aureobasidium pullulans and Bacillus mojavensis A21 isolated from sea saltern of the China Yellow Sea and seawater, respectively, produced high yields of detergent-stable alkaline protease exhibiting excellent compatibility with a variety of commercial liquid and solid detergents [85,86]. The high specificity of enzymatic reactions indisputably mitigates the typical damages to fabrics and surfaces following the use of chemically harsh detergent components [19]. Besides, the types and ratios of enzymes in detergent mixes can be optimized to favor specific detergent applications. For instance, since dishwashing detergents would need to ensure the removal of starchy food and fat/oil deposits, they often contain varying degrees of amylase and lipase [1,19]. The industrial application of alkaline cellulases as a potential additive to laundry detergents is being actively pursued with a view to selectively contact the cellulose within the interior of fibers and facilitate soil removal from the interfibrillar spaces [87].
7.2.7 Textile industry The use of enzymes in the textile industry is one of the most rapidly growing fields in industrial enzymology. Enzymes used in the textile industry offer
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various advantages as they accelerate the reaction by operating under milder conditions, providing an alternative to the polluting chemicals otherwise used. Also, they are biodegradable and easy to control, acting only on specific substrates. Current applications in the textile industry lie in the areas of biopolishing and bioscouring of fabric, antifelting of wool, softening and desizing cotton, finishing of denim and woolen items, and modification of synthetic fibers, for which the focus is primarily on hydrolases such as amylase, cellulase, protease, pectinase, and lipase/esterase [88]. There are two well-established enzyme applications in the textile industry. Firstly, in the preparatory finishing area, amylases are commonly used during the desizing process; and secondly, cellulases are used for softening in the finishing area, biostoning, and reducing of pilling propensity for cotton goods. The enzymatic desizing of cotton with a-amylases has been a state-of-the-art procedure for many decades [89]. Moreover, cellulases, pectinases, hemicellulases, lipases, and catalases are used in different cotton pretreatment and finishing processes [90]. Enzymes are emerging in a big way in the field of textile wet processing. If their cost could be managed, enzymes can be put to use in a much bigger way for textile-processing applications.
7.2.8 Leather industry For facilitating procedures and enhancing leather quality, enzymes are required during the various stages that are integrated in the processing of leather, such as curing, soaking, liming, dehairing, bating, picking, degreasing, and tanning [91]. The usage of enzymes as substitutes to chemicals has proved efficacious in improving leather quality and in reducing environmental pollution. Lipases as well as alkaline and neutral proteases are primarily used. By removing nonfibrillar proteins during soaking and bating, alkaline proteases ensure that the finished leather is soft, supple, and pliable. Water wastage is minimized by the use of neutral and alkaline proteases during the dehairing step [92]. Removal of fats during degreasing is ensured by the addition of lipases [93]. Stainless pelt, low biological oxygen demand and chemical oxygen demand in effluents, reduced odor, and enhanced hair recovery are the benefits derived by substituting chemicals with enzymes during the liming process [1].
7.2.9 Paper and pulp industry Aiming to diminish adverse effects on the ecosystem, the use of microbial enzymes in the paper and pulp industry has recorded steady growth. Besides saving on time and energy consumption, incorporation of enzymes cuts down on the use of chemicals during processing. Enzymes are also being used to enrich deinking and bleaching [94] and xylanases and ligninases, in particular, serve to enhance the value of the pulp by removing lignin and hemicelluloses
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[95]. Raghukumar et al. had reported the potential application of crude culture filtrate of a marine fungal isolate possessing thermostable, cellulase-free alkaline xylanase activity in biobleaching of paper pulp [96]. Amylases are used by paper and pulp industries for starch coating, improving paper cleanliness, and drainage [97]. Lipases find use in deinking and enhancing pitch control, while cellulases are involved in deinking, improving softness, and drainage [97]. The role of cellulases in the bioprocess development for recycling of used printed paper is also on record [98].
7.2.10 Organic synthesis The application of biocatalysts has invited growing attention over the past few years, and heavy demands have been placed on identifying new biocatalysts for the organic synthesis of novel compounds. The catalysis of numerous organic reactions directs the significance and high potential of this field of research. Enzymes perform an important role as biocatalysts in the synthesis of vital intermediates for the pharmaceutical and chemical industry, and new enzymatic technologies and processes have been recognized [99]. Through the last three decades, using hydrolases for the catalysis of environmentally friendly organic processes under mild reaction conditions has been well documented. The hydrolases have presented themselves as ideal tools for acceleration of synthetic transformations due to their broad substrate specificity, high stability, commercial availability, and catalytic efficiency in a wide spectrum of biocatalyzed processes. In recent years novel examples associated with nonconventional reactions catalyzed by hydrolytic enzymes have been witnessed. Amongst the biocatalysts in organic synthesis, lipases and acylases have earned great attention as promiscuous biocatalysts displaying good levels of reactivity in unique hydrolytic reactions, Ceheteroatom bond formation, CeC bond formation, and oxidative processes [100]. In particular, lipases perform enantioselective hydrolytic reactions and catalyze the formation of a varied range of ester and amide bonds [101]. Furthermore, lyases are involved in the organic synthesis of acrylamide from acrylonitrile, cyanohydrins from ketones, and malic acid from fumaric acid. The use of glucose isomerase has facilitated the food and beverage industry with an annual commercial production of a multimillion ton of high-fructose corn syrup as an alternative sweetener to sucrose [19].
7.2.11 Waste treatment Numerous enzymes are involved in the degradation of toxic pollutants and their usage in waste management has been extensive. In addition to the issues of domestic waste, industrial effluents foul the ecosystem with many lethal or toxic chemical entities. Wastewater from slaughterhouses and dairies contains high levels of proteins and fats that present low biodegradability.
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Several pretreatment systems are employed to remove oil and grease to avoid many of the problems that might otherwise arise in the biological process and decrease the effectiveness of the treatment station [102]. Alone or in combinations, microbial enzymes have been employed in industrial effluent treatment by degradation or bioconversion of toxic compounds to less harmful products [103]. The enzymes involved in waste treatment are amylases, amyloglucosidases, cellulases, glucoamylases, lipases, pectinases, and proteases [19]. These enzymes are also used to convert starch to sugar, to recover additional oil from oil seeds, and to convert whey to numerous useful products [104]. A wide variety of scientific investigations are ongoing for the development of enzymatic hydrolysis processes to precede traditional biological treatment.
7.2.12 Nanoparticle synthesis Metal nanoparticles synthesized using biomolecules are an attractive prospect owing to their variety in shapes and sizes as well as stability in colloidal solutions. The broad range of nanoparticle utility centers around their small size yet larger surface area. While a host of approaches are available for their synthesis [105], the usage of harmful radiations and chemical processes being discouraged in the current scenario, green and sustainable approaches gain favor, wherein enzymes are fast finding a place. Silver nanoparticles (AgNPs) have been extensively used in many commercial products. They are important modules for research in biomedicine, electronics, optics, magnetics, catalysis, mechanics, energy science, and so on [106]. Gold nanoparticles (AuNPs) have huge prospects as drug carriers, in gene therapy for gene delivery, and also in optical biosensors. Pure forms of alpha-amylase have been used in the biosynthesis of AgNPs, as the enzyme reduces the silver ions, which results in the construction of stable AgNPs [107]. They are also used in the synthesis of AuNPs by reduction of tetrachloroaurate, ensuring stabilizing of the nanoparticles by capping in colloidal solution [107]. In vitro synthesis of AgNPs and AuNPs using cellulase enzyme in a single-step reaction has also been reported [108]. Enzyme-assisted green synthesis of nanoparticles is extremely dependent on the enzyme itself, the nature of the metal salt, as well as pH of the solution. The nature of capping proteins and the strength of interaction of proteins with metal nanoparticles would decide the end stability. This leads to diversity in morphologies and size control and hence monodispersity index. Enzyme-based metal nanoparticle synthesis being still in its infancy, the probability of uncovering vast applications cannot be underestimated.
7.3 Prospecting the use of hydrolytic enzymes from marine microbes Currently, just about 5% of all chemical products are synthesized/produced using biocatalysts, and there are various doors to be yet opened to combat the
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numerous crises related to growing population, pollutants, or cost and time effectiveness as well as resource consumption. In recent years, the marine biological progress and enzyme bioprospecting activity have given a new source and options to humans, acquiring new potency and effectiveness especially for exploitation of bioactive compounds from marine microbial communities. As enzymes have incomparable advantages, many industries are keenly interested in adapting enzymatic methods to the requirements of their processes. Prospects are excellent for continuing to increase the usage of currently available enzymes in ongoing applications, as also in the use of novel enzymes for other purposes. Japan, Canada, Spain, Finland, Russia, and other countries have been focusing more on marine bioenzyme research. Taken as a whole, on account of the marine biological diversity and the specificity of biological metabolism, while the study on a global scale is but just beginning, this area has huge potential for development and applications with industrial benefits.
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116 Advances in Biological Science Research [53] Ferreira-Dias S, Sandoval G, Plou F, Valero F. The potential use of lipases in the production of fatty acid derivatives for the food and nutraceutical industries. Electron J Biotechnol 2013;16:1e38. [54] Li PY, Ji P, Li CY, Zhang Y, Wang GL, Zhang XY. Structural basis for dimerization and catalysis of a novel esterase from the GTSAG motif subfamily of the bacterial hormonesensitive lipase family. J Biol Chem 2014;289:19031e41. [55] Yuan D, Lan D, Xin R, Yang B, Wang Y. Biochemical properties of a new cold-active mono- and di- acylglycerol lipase from marine member Janibacter sp. strain HTCC2649. Int J Mol Sci 2014;15:10554e66. [56] Sharma M, Kumar A. Xylanases: an overview. Br Biotechnol J 2013;3:1e28. [57] Qureshi MA, Khare AK, Pervez A. Enzymes used in dairy industries. Int J Appl Res 2015;1:523e7. [58] Kieliszek M, Misiewicz A. Microbial transglutaminase and its application in the food industry. A review. Folia Microbiol 2014;59:241e50. [59] Zhao GY, Zhou MY, Zhao HL, Chen XL, Xie BB, Zhang XY, et al. Tenderization effect of cold-adapted collagenolytic protease MCP01 on beef meat at low temperature and its mechanism. Food Chem 2012;134:1738e44. [60] Law BA. The nature of enzymes and their action in foods. In: Whitehurst RJ, Law BA, editors. Enzyme in food technology. Sheffileld: Sheffield Academic Press; 2002. p. 1e18. [61] Damare S, Raghukumar C, Muraleedharan UD, Raghukumar S. Deep-sea fungi as a source of alkaline and cold-tolerant proteases. Enzym Microb Technol 2006;39:172e81. [62] Karlund A, Moor U, Sandell M, Karjalainen RO. The impact of harvesting, storage and processing factors on health-promoting phytochemicals in berries and fruits. Processes 2014;2:596e624. [63] Kumar S. Role of enzymes in fruit juice processing and its quality enhancement. Adv Appl Sci Res 2015;6:114e24. [64] Guo M. Functional foods: principles and technology. Cambridge: Woodhead Publishing; 2009. https://doi.org/10.1533/9781845696078. [65] Lima DM, Fernandes P, Nascimento DS, Ribeiro RCLF, de Assis SAA. Fructose syrup: a biotechnology asset. Food Technol Biotechnol 2011;49:424e34. [66] Garg G, Singh A, Kaur A, Mahajan R. Microbial pectinases: an ecofriendly tool of nature for industries. 3 Biotech 2016;6:47e59. [67] Molkabadi EZ, Hamidi-Esfahani Z, Sahari MA, HoseinAzizi M. A new native source of tannase producer, Penicillium sp. EZ-ZH190: characterization of the enzyme. Iran J Biotechnol 2013;11:244e50. [68] Choct M. Enzymes for the feed industry: past, present and future. World Poult Sci J 2006;62:5e15. [69] Kang W, Xu Y, Qin L, Wang Y. Effects of different b-D-Glycosidases on bound aroma compounds in muscat grape determined by HS-SPME and GC-MS. J Inst Brew 2010;116:70e7. [70] Mojsov K. Use of enzymes in wine making: a review. Int J Market Technol 2013;3:112e27. [71] Wang Y, Zhang C, Li J, Xu Y. Different influences of b-glucosidases on volatile compounds and anthocyanins of Cabernet Gernischt and possible reason. Food Chem 2013;140:245e54. [72] Su E, Xia T, Gao L, Dai O, Zhang Z. Immobilization of b glucosidase and its aroma increasing effect on tea beverage. Food Bioprod Process 2010;88:83e9.
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[73] Goyal K, Selvakumar P, Hayashi K. Characterization of a thermostable b-glucosidase (BglB) from Thermotogamaritima showing transglycosylation activity. J Mol Catal B Enzym 2001;15:45e53. [74] Bhat MK. Cellulases and related enzymes in biotechnology. Biotechnol Adv 2000;18:355e83. [75] Kirk O, Borchert TV, Fuglsang CC. Industrial enzyme applications. Curr Opin Biotechnol 2002;13:345e51. [76] Ulery BD, Nair LS, Laurencin CT. Biomedical applications of biodegradable polymers. J Polym Sci B Polym Phys 2011;49:832e64. [77] Zhang Z, Marquardt RR, Wang G, Guenter W, Crow GH, Han Z, et al. A simple model for predicting the response of chicks to dietary enzyme supplementation. J Anim Sci 1996;74:394e402. [78] Kobayashi S. Lipase-catalyzed polyester synthesis - a green polymer chemistry. Proc Jpn Acad Ser B Phys Biol Sci 2010;86:338e65. [79] Gurung N, Ray S, Bose S, Rai V. A broader view: microbial enzymes and their relevance in industries, medicine, and beyond. BioMed Res Int 2013. https://doi.org/10.1155/2013/ 329121. Article ID 329121. [80] Yan J, Han B, Gui X, Wang G, Xu L, Yan Y, et al. Engineering Yarrowiali polytica to simultaneously produce lipase and single cell protein from agro-industrial wastes for feed. Sci Rep 2018;8:758. [81] Dewan SS. Global markets for enzymes in industrial applications. MA, USA: BCC Research: Wellesly; 2017. [82] Kumar D, Savitri, Thakur N, Verma R, Bhalla TC. Microbial proteases and application as laundry detergent additive. Res J Microbiol 2008;3:661e72. [83] Singh A, Kuhad RC, Ward OP. Industrial application of microbial cellulases. In: Kuhad RC, Singh A, editors. Lignocellulose Biotechnology: future prospects. New Delhi, India: I.K. International Publishing House; 2007. p. 345e58. [84] Greene RV, Grifin HL, Cotta1 MA. Utility of alkaline protease from marine shipworm bacterium in industrial cleansing applications. Biotechnol Lett 1996;18:759e64. [85] Chi ZM, Ma C, Wang P, Li HF. Optimization of medium and cultivation conditions for alkaline protease production by the marine yeast Aureobasidium pullulans. Bioresour Technol 2007;98:534e8. [86] Haddar A, Agrebi R, Bougatef A, Hmidet N, Sellami-Kamoun A, Nasri M. Two detergent stable alkaline serine-proteases from Bacillus mojavensis A21: purification, characterization and potential application as a laundry detergent additive. Bioresour Technol 2009;100:3366e73. [87] Araujo R, Casal M, Cavaco-Paulo A. Application of enzymes for textiles fibers processing. Biocatal Biotechnol 2008;26:332e49. [88] Marcher D, Hagen HA, Castelli S. ITB Veredlung 1993;39:20e32. [89] Meyer-Stork LS. Maschen Ind 2002;52:32e40. [90] Mojsov K. Applications of enzymes in the textile industry: a review. In: 2nd international congress: engineering, ecology and materials in the processing industry. Jahorina, Bosnia and Herzegovina: Tehnol. Oski Fakultet Zvornik; 2011. p. 230e9. [91] Rao MB, Tanksale AM, Ghatge MS, Deshpande VV. Molecular and biotechnological aspects of microbial proteases. Microbiol Mol Biol Rev 1998;62:597e635. [92] Choudhary RB, Jana AK, Jha MK. Enzyme technology applications in leather processing. Ind J Chem Technol 2004;11:659e71.
118 Advances in Biological Science Research [93] Srivastava N, Singh P. Degradation of toxic pollutants from pulp & paper mill effluent. Discovery 2015;40:221e7. [94] Maijala P, Kleen M, Westin C, Poppius-Levlin K, Herranen K, Lehto JH, et al. Biomechanical pulping of softwood with enzymes and white-rot fungus Physisporinus rivulosus. Enzym Microb Technol 2008;43:169e77. [95] Kuhad RC, Gupta R, Singh A. Microbial cellulases and their industrial applications. Enzym Res 2011:1e10. https://doi.org/10.4061/2011/280696. [96] Raghukumar C, Muraleedharan U, Gaud VR, Mishra RJ. Xylanases of marine fungi of potential use for biobleaching of paper pulp. J Ind Microbiol Biotechnol 2004;31:433e41. [97] Kirk TK, Jeffries TW. Role of microbial enzymes for pulp and paper processing. In: ACS symposium series: Am. Chem. Soc. Washington, DC; 1996. p. 2e14. [98] Patrick K. Enzyme technology improves efficiency, cost, safety of stickies removal program. Pap Age 2004;120:22e5. [99] Eugene AL, Uchechukwu OC, Mariagoretti UO. A review on biological catalysts in organic synthesis. Int J Adv Eng Res Appl 2016;2:296e321. [100] Busto E, Gotor-Ferna´ndez V, Gotor V. Hydrolases: catalytically promiscuous enzymes for non-conventional reactions in organic synthesis. Chem Soc Rev 2010;39:4504e23. [101] Davis BG, Boyer V. Biocatalysis and enzymes in organic synthesis. Nat Prod Rep 2001;18:618e40. [102] Cammarota MC, Freire DMG. A review on hydrolytic enzymes in the treatment of wastewater with high oil and grease content. Bioresour Technol 2006;97:2195e210. [103] Pandey D, Singh R, Chand D. An improved bioprocess for synthesis of acetohydroxamic acids using DTT (dithiothreitol) treated resting cells of Bacillus sp. APB-6. Bioresour Technol 2011;102:6579e86. [104] Kalia VC, Rashmi, Lal S, Gupta MN. Using enzymes for oil recovery from edible seeds. J Sci Ind Res 2001;60:298e310. [105] Iravani S, Korbekandi H, Mirmohammadi SV, Zolfaghari B. Synthesis of silver nanoparticles: chemical, physical and biological methods. Res Pharm Sci. 2014;9:385e406. [106] Borase HP, Salunke BK, Salunkhe RB, Patil CD, Hallsworth JE, Kim BS, et al. Plant extract: a promising biomatrix for ecofriendly, controlled synthesis of silver nanoparticles. Appl Biochem Biotechnol 2014;173:1e29. [107] Khan I, Duttaa JR, Ganesanb R. Enzymes’ action on materials: recent trends. J Cell Biotechnol 2016;1:131e44. [108] Mishra A, Sardar M. Cellulase assisted synthesis of nano-silver and gold: application as immobilization matrix for biocatalysis. Int J Biol Macromol 2015;77:105e13.
Further reading [1] Kamm B, Kamm M. Principles of biorefineries. Appl Microbiol Biotechnol 2004;64: 137e45.
Biotechnological implications of hydrolytic enzymes from marine microbes Poonam Vashist1, R. Kanchana2, Vazhakatt Lilly Anne Devasia1, #, Priyanka V. Shirodkar1, Usha D. Muraleedharan1, * 1
Department of Biotechnology, Goa University, Goa, India; 2Department of Biotechnology, Parvatibai Chowgule College of Arts and Science -Autonomous, Margao, Goa, India; # Present address: Department of Biotechnology, Hindustan College of Arts and Science, Padur, Kelambakkam, Chennai, India * Corresponding author: [email protected].
7.1 Introduction Biocatalyst and biotechnology are intervolved terms that have camouflaged almost every socioenvironmental activity of our living style in the 21st century. To date, almost 4000 enzymes are known, out of which approximately 5% of microbial original types are used commercially [1]. The marine microbial population, while still largely unexplored because of its diversities and extreme environment, is being screened with the help of high-throughput technologies such as shotgun sequencing or pyrosequencing for deeper exploration [2], resulting in the reporting of approximately 20,000 species per liter of marine water samples [3]. Microbial enzymes have proved superior over chemical catalysts and enzymes derived from plants or animals, by virtue of their versatility, stability, enantioselectivity, reduced process time, intake of low-energy input, and costeffectiveness, besides their nontoxic and eco-friendly characteristics [4]. Competition amongst microorganisms for space and nutrients in the marine environment is a powerful selective force that has led to the generation of multifarious enzyme systems to adapt to the complicated environments. Many of them are thus endowed with desirable features, from a general biotechnological perspective. However, these features of microbes as well as enzymes can be enhanced with the help of techno-studies such as recombinant DNA/ protein engineering [5].
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Marine microorganisms, whose genetic and biochemical diversity studies are not quite out of their infancy, are of significant interest as a potential source for novel applications [6,7]. Researchers the world over have successfully isolated and characterized a variety of enzymes with novel activities from marine bacteria, actinomycetes, fungi, and other marine microorganisms, of which a few products have already found large-scale industrial applications [8e11]. Extremophilic marine microorganisms living in hydrothermal vents, sea floors, salt pans, and hot springs have been found to be rich sources of extremozymes such as proteases, cellulases, amylases, and lipases [12]. Our laboratory has had excellent success with bioprospecting of marine enzymes from several such sources. Vashist et al. reported a novel strain of Microbulbifer sp. isolated from a mangrove area in Divar Island, Goa, India, which exhibited more than 13 ICP degrading activities [13]. Another important avenue of interest has been the marine biodegraders classed as thraustochytrids. Kanchana et al. [14] have reported alkaline lipase activity with potential biotechnological application from these marine protists. Similarly, Devasia & Muraleedharan [15] have reported multiple polysaccharide-degrading activities from thraustochytrids for the first time. Employing response surface methodology, Shirodkar & Muraleedharan have successfully optimized a medium for commendable production of amylolytic activities from a thraustochytrid strain [16,17]. Almost 75% of all industrially applied enzymes are of the hydrolytic category, out of which proteases, carbohydrases, and lipases lead the enzyme market, with more than 70% of all enzyme sales [1]. Among these, hydrolytic enzymes resourced from bacteria hold the majority share when compared to the overall bioactive compounds discovered from marine fungi and other domains [18]. The world of enzyme demand is satisfied by about 12 major producers and 400 minor suppliers [19]. Three top companies, namely, Denmark-based Novozymes and Danisco and Switzerland-based Roche satisfy about 75% of the total enzyme demand [1]. While the world market for industrial enzymes was to the tune of $4.2 billion in 2014, an expansion at a compound annual growth rate (CAGR) of approximately 7% is predicted over the next 5-year period, escalating the figure to nearly $6.2 billion [20]. In this chapter, attention is drawn to the significant role of hydrolytic enzymes from marine microbes involved in various fields of technical applications. Also reported are aspects of modern biotechnology for improved enzyme production, enabling various industrial processes at lower energy consumption and higher efficiency, besides enhancing properties of the product while ensuring environment friendliness.
7.2 Applications of marine hydrolases Hydrolases catalyze several reactions, including condensations, alcoholysis, and so on. The main advantages of this enzyme class are ready availability, lack of cofactor stereoselectivity, and tolerance to the addition of water-miscible solvents. In fact, the majority (about 75%) of currently used microbial enzymes in various industries fall into the hydrolytic category [21].
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7.2.1 Biorefineries Over several years, studies have been in progress in different parts of the world to find substitutes for the fractions of nonrenewable/fossil resources in use. Brazil, the United States, China, and India, covering the most-populated regions of the world at 44% of the total global population, use 45% of the total energy sources and account for 51% of the carbon dioxide release from the consumption of energy. Notwithstanding this, about 27% of primary energy is used worldwide for transportation, thus highlighting transportation fuels as notable targets for replacement with a renewable source [22]. Consistent and harmonized supply of raw materials is the most important factor in biorefinery productivity [23]. The use of aquaculture biomass (seaweeds) as a feed for biorefinery industries ensures a continuous, rapid, safer, and economic supply of feed for biorefinery industries compared to the use of lignocellulosic biomass from agricultural wastes and other biological materials. Algae have 20e80% oil content, most of which are triacylglycerols, but the robust algal cell walls present a challenge for the extraction of these lipids. Enzymatic lipid extraction has beencarried out withsnailase along with cellulase and lysozyme [24]. Novel and significant enzymatic properties of extremophiles and thermostable enzymes have boosted the biocatalysis in biofuel production [25]. In biorefineries for bioethanol, biogas, and biodiesel production, cellulases, hemicellulases, and ligninases are most widely used, primarily during pretreatment and saccharification of biomass. Some bacteria and fungi are capable of rapid and efficient degradation of cellulose [25,26]. Enhanced depolymerization of the cellulose fraction from lignocellulosic substrates would be favored by the use of a cocktail of enzymes such as exoglucanase, endoglucanase, b-glucosidase, cellobiase, xylanase, and other hemicellulases [27].
7.2.2 Pharmaceuticals and cosmeceuticals Enzymes play various vital roles in the realms of pharmaceuticals, cosmetics, and diagnostics. The pharmaceutical industry in particular requires a very high degree of substrate specificity on account of the stringent routes for desired product synthesis. Coupled with the effective processing cost, this propels enzymes to advantage over chemical catalysts [4]. Enzymes are extensively being used as large-scale therapeutic drugs mainly pertaining to health concerns associated with enzymatic deficiency, digestive disorders, and in ELISA-based diagnostics [28]. Marine natural products play an important role in drug development [29]. The enzymes involved in mycothiol biosynthesis and found in the Actinobacteria, Actinomycetes and Mycobacterium are of great interest as potential targets for the production of new drugs against Mycobacterium tuberculosis [30]. The applications of hydrolytic enzymes in medicine are growing rapidly, as in the removal of dead skin, and enzymes such as Vibrilase (vibriolysin)
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obtained from the marine microorganism Vibrio proteolyticus have been reported for effectiveness against denatured proteins found in burnt skin [31]. Lipases have been cited as the most repeatedly used enzymes in the synthesis of alcohols, acids, lactones, and esters that are optically active [32, 33]. On account of their digestive and inflammatory properties, proteases have been testified for a wide array of applications in medicine [34]. Chitosanase catalyzes the hydrolysis of chitosan, which is used as an antimicrobial, antioxidant, and in bringing down high blood cholesterol and blood pressure levels. It has also found use in the control of arthritis, besides enhancing antitumor properties and according protection against infections [35]. Actinobacteria are considered the most important group of organisms being studied extensively for the discovery and formulations of significant drugs and other bioactive metabolites with high specificity [36]. Glucanase isolated from a marine Bacillus has been reported suitable for oral and other health care [37]. Oligomers obtained from carrageenan degradation have demonstrated antitumor, antiviral, anticoagulative, and other activities such as boosting immune function and digestion [38e40]. Qiu et al. reported significant advantages of the marine bacterial protease from an N1-35 strain isolated after UV mutagenesis over those from terrestrial ones [41]. Enzymes such as urease and creatinase from Actinobacteria have been used in clinical diagnostics for quantitative evaluation of diabetes and other health disorders [42,43]. Enzymes have gained increasing popularity in the cosmetic industry by virtue of their relative safety and minimal side effects. Li et al. [1] have reported the use of enzymes as free radical scavengers in preparations for hair styling and dyeing, and in toothpastes, mouthwashes, and sunscreen products. In Japan, agar-oligosaccharide is being extensively used as a moisturizing cosmetic additive and and has also been reported for good hair conditioning effects [44]. Proteases have been used in skin creams to remove dead cells and thereby effect cleaning and smoothening of the skin [45]. Papain and endoglycosidase have found wide use in toothpastes and mouthwashes, serving to whiten and remove plaque as well as other odor-causing deposits on teeth and gum tissue [46]. Lipase and papain are used in skin care products [47]. Enzymes are also routinely being incorporated in contact lens cleaners to remove proteinaceous films [48].
7.2.3 Food industry The present rate of world average for per capita food consumption is 3000 kcal/day [19]. The estimated food consumption is expected to go up to 22.8 1012 kcal/day by 2020; when compared to the energy, it is roughly 16 million barrels of oil equivalent (boe) per day i.e., an approximate 70% increase by 2050 [19]. Meeting these requirements in time without compromising on food quality could be managed by enzymes application. These biocatalysts have been reported as well as used efficiently for improving food
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production, quality and components such as flavor, aroma, color, texture, appearance, fat modification, and in sweetener technology [1]. Application of enzymes in the food industry is principally for improving the shelf life of products, besides their stability and texture. Agar oligosaccharides produced by the action of agarases are being used for the production of beverages, bread, and some low-calorie food [49,50]. Isolates such as Microbulbifer CMC-5, 10A, and LK2 have been reported for promising and extensive agarolytic activity that can be further exploited [51,52]. Agarose-derived neoagarooligosaccharides produced by b-agarase from the seaweed-decomposing bacterium Microbulbifer sp. strain CMC-5 showed a promising biotechnological application [52]. Similarly, esterases and lipases have been reported for betterment of bread quality by change in lipids in the dough used, in butter, cheese, and margarine flavor enhancement, in the production of crackers, pasta, etc., degumming of vegetable oils, synthesis of infant formula and nutraceuticals, increasing the concentration of polyunsaturated fatty acids in vegetable oils, as well as for improving the digestibility of natural lipids [53]. Recently Li et al. isolated a novel esterase of the hormone-sensitive lipase family, from a metagenomic library of more than 10,500 cosmid clones from the South China Sea [54]. A cold-active lipase from the marine bacterium Janibacter sp. strain HTCC2649 was isolated and characterized by Yuan and coworkers with sn-1/3 specificity toward monoand di- acylglycerols [55]. Recent reviews have endorsed the application of xylanases, lipases, and amylases in the baking industry, for improving freshness and shelf life of the products, elasticity of gluten, enhancing handling, and increasing stability of the dough, as well as improved bread volumes and absorption/redistribution of water and flavor; higher content of arabinoxylooligosaccharides in bread with a positive effect on health, lighter cream crackers, improved texture, palatability, and uniformity of wafers are other benefits that have been derived [1,56,57]. Cheese, yogurt, and other milk products have been improved upon with respect to organoleptic characteristics such as aroma, flavor, color, and yield [58]. The use of enzymes (protease, catalase, lipase, esterase, lactase, etc.) in the dairy market is well recognized for reduction of allergenic properties of milk products [59]. The effectiveness of collagenase and protease has been proven in meat tenderization and further processing and protease MCP-01, an abundant extracellular serine protease produced by the deep-sea psychrophilic Pseudoalteromonas sp. SM9913, is reported to be one such psychrotolerant enzyme [60]. Damare et al. have described a detergent-compatible alkaline, coldtolerant protease from a deep-sea fungus from the Central Indian Basin [61]. Enzymes have been employed in the beverage industry to control the brewing process, and to produce beer of consistently high quality. Improvement of the juice yield, aroma, and color are other positive outcomes.
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Such enzymes also serve to enhance the nutritional and functional characteristics of proteins of animal and vegetable origin that are used in the process [62]. The first stage in a brewing industry for extraction of material would be to digest the plant cell wall using microbial enzymes [63]. Inulinase form Pichia guilliermondii OUC1 has been reported for the production of ultrahigh-fructose syrups, while Yarrowia lipolytica OUC2 could be used for the production of inulooligosaccharides [64,65]. It has been reported very effective to use amylases, cellulases, arabinases, naringinase, pectinases, and xylanases during fruit juice/beverage processing to enhance yield, in juice clarification, for improved extraction, stability, texture, sensory characteristics, and cost/time management [1,66,67]. Tannases from Aspergillus awamori BTMFW032 have been used for producing coffeeflavored soft drinks and instant tea, for the clarification of fruit juice and beer, and for food detannification [68]. Many reports describe b-glucosidases as widely used during processing of fruit juices, tea, wine, and beer to effect enhancement of their organoleptic properties and essential oil content [69e72]. One specific study on b-glucosidases from Thermotoga maritime has been documented by Goyal et al. [73].
7.2.4 Feed industry The continuously increasing trend in worldwide milk and meat consumption invariably escalates the demand for the feedstock for animals, and to meet the requirements of quantity and quality, the search for feed enzymes for diet formulations necessarily had to pick up the pace. Enzymes incorporated in animal feeds aid to increase the digestibility of nutrients and ensure better feed utilization [74]. The practice of supplementation of animal diets with enzymes increased exponentially from the 1980s to the subsequent decades. The world market for feed enzymes, assessed at $899.19 million in 2014, is estimated to touch the $1.3 billion mark by 2020, at a CAGR of 7.3% over the period [1]. The protein concentration in poultry feeds has been enhanced by the application of feed enzymes such as a-galactosidases, phytases, a-amylases, proteases, glucanases, xylanases, and polygalacturonases [1]. Faced with the inability of monogastric animals to digest plant-based feeds rich in hemicelluloses and cellulose, xylanase and b-glucanase have been added to the feeds to enable total degradation and digestion [75,76]. To the same end, b-glucosidases have been documented as favored additives in cellulose-based feeds for animals such as chicken and pigs [77].
7.2.5 Biopolymer industry Involving microbial enzymes in the synthesis of biodegradable polymers appears the best solution to circumvent the increased requirement of polymers that have proved detrimental to human health and environment. The applications of biopolymers such as polyesters, polycarbonates, and polyphosphates have been
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reported for various biomedical purposes, e.g., orthopedic devices, tissue engineering, adhesion barriers, and controlled drug delivery [78]. The biopolymer market, which had grown substantially over the recent years, was expected to touch $3.6 billion by the end of 2018 [1]. The ever-increasing clamor for packaging materials and the concomitant environmental safety concerns could both be addressed by the flourishing biodegradable polymer industry. Lipases and laccases catalyze polymerization reactions to produce polyesters or polycarbonates and cross-links in biopolymers [79,80]. Y. lipolytica lipase when customized by immobilization and protein engineering has been shown to improve the performance in biopolymer synthesis [81].
7.2.6 Detergent industry Successful integration of hydrolytic enzymes into detergent formulations is cited as the key to the production of cost-effective and environmentally gentle detergents [82]. In today’s laundry detergents, enzymes such as proteases and amylases are deeply entrenched among the active ingredients. A recipe of alkaline proteases and cellulases isolated from microbial sources and that can effectively remove protein stains as well as cleave damaged cotton fibers is now a vital component of various detergents produced on a commercial scale by manufacturers such as Novozymes SA, Kao Corporation, and Genencor International [83]. Greene and his colleagues had reported cleansing properties of an alkaline protease isolated from a bacterial strain symbiotic with the gland of Deshayes of a marine shipworm [84]. Aureobasidium pullulans and Bacillus mojavensis A21 isolated from sea saltern of the China Yellow Sea and seawater, respectively, produced high yields of detergent-stable alkaline protease exhibiting excellent compatibility with a variety of commercial liquid and solid detergents [85,86]. The high specificity of enzymatic reactions indisputably mitigates the typical damages to fabrics and surfaces following the use of chemically harsh detergent components [19]. Besides, the types and ratios of enzymes in detergent mixes can be optimized to favor specific detergent applications. For instance, since dishwashing detergents would need to ensure the removal of starchy food and fat/oil deposits, they often contain varying degrees of amylase and lipase [1,19]. The industrial application of alkaline cellulases as a potential additive to laundry detergents is being actively pursued with a view to selectively contact the cellulose within the interior of fibers and facilitate soil removal from the interfibrillar spaces [87].
7.2.7 Textile industry The use of enzymes in the textile industry is one of the most rapidly growing fields in industrial enzymology. Enzymes used in the textile industry offer
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various advantages as they accelerate the reaction by operating under milder conditions, providing an alternative to the polluting chemicals otherwise used. Also, they are biodegradable and easy to control, acting only on specific substrates. Current applications in the textile industry lie in the areas of biopolishing and bioscouring of fabric, antifelting of wool, softening and desizing cotton, finishing of denim and woolen items, and modification of synthetic fibers, for which the focus is primarily on hydrolases such as amylase, cellulase, protease, pectinase, and lipase/esterase [88]. There are two well-established enzyme applications in the textile industry. Firstly, in the preparatory finishing area, amylases are commonly used during the desizing process; and secondly, cellulases are used for softening in the finishing area, biostoning, and reducing of pilling propensity for cotton goods. The enzymatic desizing of cotton with a-amylases has been a state-of-the-art procedure for many decades [89]. Moreover, cellulases, pectinases, hemicellulases, lipases, and catalases are used in different cotton pretreatment and finishing processes [90]. Enzymes are emerging in a big way in the field of textile wet processing. If their cost could be managed, enzymes can be put to use in a much bigger way for textile-processing applications.
7.2.8 Leather industry For facilitating procedures and enhancing leather quality, enzymes are required during the various stages that are integrated in the processing of leather, such as curing, soaking, liming, dehairing, bating, picking, degreasing, and tanning [91]. The usage of enzymes as substitutes to chemicals has proved efficacious in improving leather quality and in reducing environmental pollution. Lipases as well as alkaline and neutral proteases are primarily used. By removing nonfibrillar proteins during soaking and bating, alkaline proteases ensure that the finished leather is soft, supple, and pliable. Water wastage is minimized by the use of neutral and alkaline proteases during the dehairing step [92]. Removal of fats during degreasing is ensured by the addition of lipases [93]. Stainless pelt, low biological oxygen demand and chemical oxygen demand in effluents, reduced odor, and enhanced hair recovery are the benefits derived by substituting chemicals with enzymes during the liming process [1].
7.2.9 Paper and pulp industry Aiming to diminish adverse effects on the ecosystem, the use of microbial enzymes in the paper and pulp industry has recorded steady growth. Besides saving on time and energy consumption, incorporation of enzymes cuts down on the use of chemicals during processing. Enzymes are also being used to enrich deinking and bleaching [94] and xylanases and ligninases, in particular, serve to enhance the value of the pulp by removing lignin and hemicelluloses
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[95]. Raghukumar et al. had reported the potential application of crude culture filtrate of a marine fungal isolate possessing thermostable, cellulase-free alkaline xylanase activity in biobleaching of paper pulp [96]. Amylases are used by paper and pulp industries for starch coating, improving paper cleanliness, and drainage [97]. Lipases find use in deinking and enhancing pitch control, while cellulases are involved in deinking, improving softness, and drainage [97]. The role of cellulases in the bioprocess development for recycling of used printed paper is also on record [98].
7.2.10 Organic synthesis The application of biocatalysts has invited growing attention over the past few years, and heavy demands have been placed on identifying new biocatalysts for the organic synthesis of novel compounds. The catalysis of numerous organic reactions directs the significance and high potential of this field of research. Enzymes perform an important role as biocatalysts in the synthesis of vital intermediates for the pharmaceutical and chemical industry, and new enzymatic technologies and processes have been recognized [99]. Through the last three decades, using hydrolases for the catalysis of environmentally friendly organic processes under mild reaction conditions has been well documented. The hydrolases have presented themselves as ideal tools for acceleration of synthetic transformations due to their broad substrate specificity, high stability, commercial availability, and catalytic efficiency in a wide spectrum of biocatalyzed processes. In recent years novel examples associated with nonconventional reactions catalyzed by hydrolytic enzymes have been witnessed. Amongst the biocatalysts in organic synthesis, lipases and acylases have earned great attention as promiscuous biocatalysts displaying good levels of reactivity in unique hydrolytic reactions, Ceheteroatom bond formation, CeC bond formation, and oxidative processes [100]. In particular, lipases perform enantioselective hydrolytic reactions and catalyze the formation of a varied range of ester and amide bonds [101]. Furthermore, lyases are involved in the organic synthesis of acrylamide from acrylonitrile, cyanohydrins from ketones, and malic acid from fumaric acid. The use of glucose isomerase has facilitated the food and beverage industry with an annual commercial production of a multimillion ton of high-fructose corn syrup as an alternative sweetener to sucrose [19].
7.2.11 Waste treatment Numerous enzymes are involved in the degradation of toxic pollutants and their usage in waste management has been extensive. In addition to the issues of domestic waste, industrial effluents foul the ecosystem with many lethal or toxic chemical entities. Wastewater from slaughterhouses and dairies contains high levels of proteins and fats that present low biodegradability.
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Several pretreatment systems are employed to remove oil and grease to avoid many of the problems that might otherwise arise in the biological process and decrease the effectiveness of the treatment station [102]. Alone or in combinations, microbial enzymes have been employed in industrial effluent treatment by degradation or bioconversion of toxic compounds to less harmful products [103]. The enzymes involved in waste treatment are amylases, amyloglucosidases, cellulases, glucoamylases, lipases, pectinases, and proteases [19]. These enzymes are also used to convert starch to sugar, to recover additional oil from oil seeds, and to convert whey to numerous useful products [104]. A wide variety of scientific investigations are ongoing for the development of enzymatic hydrolysis processes to precede traditional biological treatment.
7.2.12 Nanoparticle synthesis Metal nanoparticles synthesized using biomolecules are an attractive prospect owing to their variety in shapes and sizes as well as stability in colloidal solutions. The broad range of nanoparticle utility centers around their small size yet larger surface area. While a host of approaches are available for their synthesis [105], the usage of harmful radiations and chemical processes being discouraged in the current scenario, green and sustainable approaches gain favor, wherein enzymes are fast finding a place. Silver nanoparticles (AgNPs) have been extensively used in many commercial products. They are important modules for research in biomedicine, electronics, optics, magnetics, catalysis, mechanics, energy science, and so on [106]. Gold nanoparticles (AuNPs) have huge prospects as drug carriers, in gene therapy for gene delivery, and also in optical biosensors. Pure forms of alpha-amylase have been used in the biosynthesis of AgNPs, as the enzyme reduces the silver ions, which results in the construction of stable AgNPs [107]. They are also used in the synthesis of AuNPs by reduction of tetrachloroaurate, ensuring stabilizing of the nanoparticles by capping in colloidal solution [107]. In vitro synthesis of AgNPs and AuNPs using cellulase enzyme in a single-step reaction has also been reported [108]. Enzyme-assisted green synthesis of nanoparticles is extremely dependent on the enzyme itself, the nature of the metal salt, as well as pH of the solution. The nature of capping proteins and the strength of interaction of proteins with metal nanoparticles would decide the end stability. This leads to diversity in morphologies and size control and hence monodispersity index. Enzyme-based metal nanoparticle synthesis being still in its infancy, the probability of uncovering vast applications cannot be underestimated.
7.3 Prospecting the use of hydrolytic enzymes from marine microbes Currently, just about 5% of all chemical products are synthesized/produced using biocatalysts, and there are various doors to be yet opened to combat the
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numerous crises related to growing population, pollutants, or cost and time effectiveness as well as resource consumption. In recent years, the marine biological progress and enzyme bioprospecting activity have given a new source and options to humans, acquiring new potency and effectiveness especially for exploitation of bioactive compounds from marine microbial communities. As enzymes have incomparable advantages, many industries are keenly interested in adapting enzymatic methods to the requirements of their processes. Prospects are excellent for continuing to increase the usage of currently available enzymes in ongoing applications, as also in the use of novel enzymes for other purposes. Japan, Canada, Spain, Finland, Russia, and other countries have been focusing more on marine bioenzyme research. Taken as a whole, on account of the marine biological diversity and the specificity of biological metabolism, while the study on a global scale is but just beginning, this area has huge potential for development and applications with industrial benefits.
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Further reading [1] Kamm B, Kamm M. Principles of biorefineries. Appl Microbiol Biotechnol 2004;64: 137e45.