Immobilization of heavy metals using sulphide-producing microorganisms has been reported as an effective means of treating some metal-contaminated sites Crawford and Crawford Laccase enzyme purified from thermophile, Geobacillus thermocatenulatus MS5 is of very higher catalytic activity and are economic, highly stable at different temperatures and pH levels and can be used widely and effectively in the removal of the dyes that cause environmental pollution. Verma and Shirkot investigated the purified laccase enzyme for the removal of some dyes used in industry i.
The results suggested the isolates as a useful tool for degradation of reactive dyes Sahni and Gupta Sporotricum thermophile LAR5 is an excellent fungal isolate having an ability to utilize crude agriculture based materials as carbon and nitrogen sources to produce significant cellulase titre.
Cellulase possesses desirable properties from industrial application point of view such as activity and stability over broad pH range and high temperatures and good saccharification ability on acid-pretreated rice straw.
It has been reported that considerable sugars are produced by enzymatic hydrolysis of acid-pretreated solids 3. Recombinant S. Thermophilic bacilli are used as hygiene indicators of processed product, within the dairy processing context. This is because of the ability of these strains to form endospores and biofilms.
The thermophilic bacilli, such as Anoxybacillus flavithermus and Geobacillus spp. Although these bacilli are generally not pathogenic, their presence in dairy products is an indicator of poor hygiene and high numbers are unacceptable to customers. In addition, their growth may result in milk product defects caused by the production of acids or enzymes, potentially leading to off-flavors Burgess et al.
Many strains of genera Lactobacillus and Bifidobacterium , as well as some enterococci and yeasts, have been shown to possess probiotic properties with potential for prophylaxis and treatment of a range of gastrointestinal disorders Varankovich et al. It is the first known thermophile that is able to degrade native feathers at high temperatures.
With the help of these enzymes, feathers could be converted to defined products such as the rare amino acids, serine, cysteine and proline Friedrich and Antranikian Asperjinone, a nor-neolignan, and Terrein, a suppressor of ABCG2-expressing breast cancer cells were isolated from thermophile Aspergillus terreus, which can restore drug sensitivity and could be the key to improve breast cancer therapeutics.
Terrein, displayed strong cytotoxicity against breast cancer MCF-7 cells. Treatment with terrein significantly suppressed growth of ABCG2-expressing breast cancer cells. This suppressive effect was achieved by inducing apoptosis via activating the caspase-7 pathway and inhibiting the Akt signaling pathway, which led to a decrease in ABCG2-expressing cells and a reduction in the side-population phenotype Liao et al.
Conventional chemotherapeutic agents are usually non specific towards cancerous cells and inhibit the progression of any dividing cells. The therapeutic potential of antitumor drugs is seriously limited by the manifestation of serious side effects and drug resistance. So there is a need of agents that are more effective, more selective and may not cause drug resistance.
According to Patent no. The increasing number of patents indicates that there is a growing interest in the commercial applications of thermophiles. The demand for thermostable enzymes has increased tremendously in the past few years. Since only a very few species from this group of microorganisms have been isolated till date, there seems to be a large number of hyperthermophilic catalysts with unique properties awaiting discovery. Brazil J Microbiol — Article Google Scholar.
Adv Biochem Eng Biotechnol — CAS Google Scholar. Can J Microbiol — Agric Biol Chem — J Mater Environ Sci 5 5 — J Bacteriol — Google Scholar. Curr Opin Biotechnol — Nat Chem Biol — Broad TE, Shepherd MG Purification and properties of glucosephosphate dehydrogenase from the thermophilic fungus Penicillium duponti.
Biochim Biophys Acta — Int J Food Microbiol — Int J Syst Evol Microbiol — Catterina G Beitrag zum Studium der thermophilen Bakterien. Zentr Bakt Parasitenk Infek Il — Chalaal O, Islam MR Integrated management of radioactive strontium contamination in aqueous stream systems. J Environ Manage — Chefetz B, Chen Y, Hadar Y Purification and characterization of laccase from Chaetomium thermophilium and its role in humification. Appl Environ Microbiol — US 6,, B1.
Int J Syst Bacteriol — Cook GM, Morgan HW Hyperbolic growth of Thermoanaerobacter thermohydrosulfuricus Clostridium thermohydrosulfuricus increases ethanol production in pH controlled batch culture. Appl Microbiol Biotechnol — Cambridge: Cambridge University Press. Curvers S, Koln DE, Svetlichnyi V Single step bioconversion of lignocellulosic biomass to biofuels using extreme thermophilic bacteria. US Patent A1. Mol Biotechnol 10 3 — Bioresour Technol — Enzyme Microb Tech — Egorova K, Antranikian G Industrial revelance of thermophilic.
Archaea Curr Opin Microbiol — Eichler J Biotechnological uses of archaeal extremozymes. Biotech Adv — US ,,A. Falcioni D I germi termofili nelle acque del Bullicame. Arch Farmacol Speri —5. In: Seckbach J ed Enigmatic microorganisms and life in extreme environments. Kluwer Academic Publishers, Dordrecht, pp — Chapter Google Scholar. Friedrich AB, Antranikian G Keratin degradation by Fervidobacterium pennavorans , a novel thermophilic anaerobic species of the order Thermotogales.
Appl Environ Microbiol 62 8 — J Mol Evol 44 6 — Ganghofer D, Kellermann J, Staudenbauer WL, Bronnenmeier K Purification and properties of an amylopullulanase, a glucoamylase, and an alphaglucosidase in the amylolytic enzyme system of Thermoanaerobacterium thermosaccharolyticum.
Biosci Biotech Bioch — Georgevitch P b Bacillus thermophilus vranjensis. Arch Hyg Sci — Gilbert Ueber Actinomyces thermophilue und andere Aktinomyceten. Z Hyg Infektionskrankh — In summary, T. More recent studies involving genetic manipulation of T. The focus has now shifted away from development of the genetic system to applying these mature techniques to answer a variety of scientific questions.
Examples of the extent to which genetic modification is possible in T. Additional metabolic engineering tools available in T. Efforts to further improve hydrogen production are ongoing Kanai et al.
Strains optimized for overexpression and secretion of enzymes have also been developed Takemasa et al. There are currently no reports of expressing full heterologous metabolic pathways in T. Robust and fast-growing, Pyrococcus furiosus is convenient to work with, and also happens to be the highest temperature organism for which a versatile genetic system is available.
Able to grow on a variety of peptide and carbohydrate substrates, P. It has already been genetically engineered to express a variety of heterologous metabolic pathways, despite the fact that the first reports of a functional genetic system did not appear until It is a euryarchaeal heterotroph capable of growth on peptides and some oligo- and polysaccharides.
Sulfur S 0 is required for peptide utilization, resulting in the production of organic acids and H 2 S as byproducts. Carbohydrates can be utilized either with S 0 or without, where reducing equivalents are disposed of as H 2 Adams et al. The genetic system in P. In fact, the first successful genetic manipulation in P. A subsequent effort applied counter-selection via 6-MP to perform a single nucleotide deletion Kreuzer et al.
These are the only reports of genetic transformations in wild-type P. Compared to the wild-type P. The uracil auxotroph for this transformation was generated by simvastatin selection, while agmatine Hopkins et al. Promoters available for protein overexpression include the strong constitutive promoter of the S-layer protein P slp Hopkins et al. Recently, entire operons up to 17 kilobases long have been transformed into P. As with most new genetic systems, the first demonstrations of genetic modification in P.
The genes encoding two soluble hydrogenases, believed to be essential to P. The ancestral ability of P. But the most impressive examples of metabolic engineering in this organism have been in the expression of heterologous pathways. The core metabolic strategies available at the extreme temperatures where P.
For example, alcohol production is very rare among extreme thermophiles, possibly because the archaea, which predominate at high temperatures, are not known to produce significant amounts of any alcohol naturally Basen et al.
Therefore, heterologous pathways to be inserted into P. Another single-enzyme insertion gave P. More complex multi-enzyme metabolic engineering has also been demonstrated in P. Deletion of a competing pathway roughly doubled 3HP titers over the parent strain Thorgersen et al.
A synthetic butanol pathway consisting of six genes from three thermophilic bacteria enabled P. Heterologous expression of the massive 17 kb, gene formate dehydrogenase operon from Thermococcus onnurineus allowed P.
Insertion of another large operon from T. With an incredible diversity of functional engineered pathways, P. However, additional improvements will be necessary to turn current strains, which often produce only trace amounts of the target chemicals, into viable industrial hosts.
Some progress has already been made in this area, particularly in 3HP production, where additional enzymes and improved growth conditions increased titers nearly ten-fold Hawkins et al.
Three Sulfolobus species, S. All three species are obligate aerobes, grow well on rich media, and single colonies can be isolated on solid substrates. Combined with their genetic tractability, these traits make them excellent model organisms.
Sulfolobus species have been used extensively to elucidate the mechanisms and cellular machinery of transcription in archaea, as a model host for archaeal viruses, and as a source of easily crystallized thermophilic proteins. So far, no member of Sulfolobus has been metabolically engineered to produce a commercially-desirable chemical product.
Following the initial isolation of S. This, combined with difficulties obtaining pure cultures, led to the use of mixed and misidentified strains during the early stages of Sulfolobus research Grogan, The first isolation report for a Sulfolobus species described its ability to grow autotrophically by oxidation of sulfur Brock et al. However, current laboratory strains of S.
A comparison of the two strains during heterotrophic growth indicates that, while both grow well on complex protein sources and starch, S. The original S. The first genetic modifications in Sulfolobus relied on nutrient selection. Lactose selection Worthington et al. Uracil selection, which has been used in all three species Albers et al. Genetic transformations have also been accomplished in S. Electroporation is the standard means of transformation across all three species.
DNA methylation is used in some methods involving S. Genome sequences are available for the type strain of S. None of the islandicus strains are available from culture collections and no type strain has been designated Zuo et al. Sulfolobus species, particularly S.
As a result, a variety of viral-based vectors are available for genetic manipulation Aucelli et al. Inducible promoter systems are available Berkner et al. While production of novel products has yet to be demonstrated in Sulfolobus , an S. Therefore, if product pathways are developed, S. There are many other examples of homologous or heterologous overexpression of proteins in Sulfolobus , sometimes because the protein cannot be expressed in functional form otherwise, but often simply as demonstration of a new method of genetic modification.
Genetic system development in Sulfolobus has been underway for over a decade, but remains a research focus. Like other thermophiles, Ts. Additionally, strain HB8 can grow anaerobically by denitrification Cava et al. Genome sequences are available for Ts. The HB8 strain appears to be polyploid, like the closely related Deinococcus radiodurans , with cells able to maintain two different antibiotic resistance genes at the same location on the chromosome Ohtani et al. The presence of multiple chromosome copies was proposed to present a significant challenge to genetic manipulation, but more recent reports of genetic modification in strain HB27 Leis et al.
Early genetics work in Ts. The genetic system of Ts. So far, the only metabolic engineering reported in Ts. It is an aerobe, capable of autotrophic growth by oxidizing sulfidic ores or hydrogen, heterotrophic growth on peptides, or a combination of the two Auernik and Kelly, The genome sequence has been published Auernik et al. Despite current limitations to the genetic system, M. Its uniquely archaeal carbon fixation pathway Kockelkorn and Fuchs, , coupled with the ability to grow on hydrogen gas, makes M.
While this ability to use lignocellulosic biomass is appealing, improving its ethanol yield is necessary to exploit T. To accomplish this, glycerol dehydrogenase from Thermotoga maritima was expressed in T. Recently re-classified from its original designation as Anaerocellum thermophilum, C. It has a published genome sequence Kataeva et al. One added challenge has been the presence of restriction enzymes, necessitating either the use of methylated transformation constructs, or strains that lack the relevant enzymes Chung et al.
Despite its recent development, the genetic system has already been used to metabolically engineer C. In addition, a heterologous gene encoding an archaeal tungsten-containing enzyme was successfully expressed in C. Members of the genus Thermotoga , including T. Members of the genus contain a remarkable number of sugar utilization genes, allowing for growth on a wide variety of carbohydrates Chhabra et al. The early publication of the genome sequence of T. A plasmid isolated from Thermotoga strain RQ7 was used to transform T.
Ten years later, T. RQ7 were finally stably transformed with an E. Transformation has been accomplished with liposomes Yu et al. The only demonstration of metabolic engineering thus far involved recombinant expression of cellulases from Caldicellulosiruptor saccharolyticus , which were fused with native signal peptides for export, leading to cellulolytic activity in the cell supernatant; however, expression was not stable long term Xu et al.
In order to understand the potential of extreme thermophiles for bioprocessing, as well as how far they still have to go, it is worth considering the current state of the art, which is still reliant on mesophilic hosts.
While many chemicals and fuels of interest have been successfully created in microorganisms, production has been hindered by a few key factors. One of the most notable can be attributed to the high proportion of water necessary for all biological processes. Thus, separating low concentrations of the target molecule from massive quantities of water requires significant energy inputs for heating, cooling, distillation, and transportation.
Another hurdle to renewable chemical production is the high cost of feedstocks, although, as shown in Table 3 , U. Various other feedstocks with potential for use by biological organisms, such as hydrogen and natural gas, are also available at competitive rates.
Table 3. Commodity prices of fuel feedstocks, including common biomass, and fossil-derived sources. In the landscape of chemicals being targeted for production via biological organisms, ethanol stands alone as the only chemical to have rivaled an industrial commodity on a volume and economic basis. Over 14 billion US gallons 53 billion liters of ethanol were produced in the United States in , mostly from corn starch, with Brazil adding over six billion gallons 23 billion liters from sugar RFA, No other biologically produced chemical or fuel has approached the one billion gallon mark.
Ethanol benefits from a number of advantages, especially the natural ability of yeast to metabolize sugars to ethanol at high titers and with high efficiency, thus avoiding the need for extensive genetic engineering. However, the success of ethanol proves that bio-based chemical production at scale is possible, and recent progress by industry startups generating a variety of other chemicals Table 4 confirms this. Table 4. Commercial scale biochemical production excluding 1st generation ethanol.
Production of 1,3-propanediol from corn starch commenced in and, nearly a decade later, the company reports progress on expanding production. Recent years have brought more commercial facilities onto the scene Table 4 , representing billions of dollars in capital investment. The move from demonstration to commercial-scale shows that the investment community is optimistic about the future prospects of biological chemical production. While none of the examples in Table 4 uses a thermophilic host, most of the research on these commercial scale projects was started a decade ago, before significant tools were available for genetic manipulation of thermophiles.
The experiences gained in these initial projects may provide further evidence of the advantages of thermophilic processes. The genetic systems for several species of extreme thermophiles are now advanced enough to begin developing metabolically engineered strains to produce industrially relevant chemicals, but so far only P. Thermophiles research has historically been focused on answering the basic questions of how life functions at high temperature: how do transcription and translation occur, how do protein molecules fold, how can metabolism proceed through thermolabile intermediates, and could the earliest life have evolved at high temperatures?
However, the emerging sector of industrial biotechnology has added the additional focus of producing fuels and chemicals by exploiting thermophilicity.
Some thoughts on this subject follow. A number of advantages become available when working with a thermophilic host. One that has already been applied in P. Figure 3. Temperature-shift strategy involving a hyperthermophilic host expressing more moderately thermostable recombinant enzymes.
Reduced temperatures result in a transition from growth to production phase. The hosts enzymes, naturally optimized for higher temperatures, become less active, and the cell growth rate stalls. Meanwhile, the recombinant enzymes from less thermophilic sources then re-fold and begin producing chemicals.
Enzyme production can furthermore be coupled to temperature shift through the use of cold-induced promoters. A similar temperature-shift has been performed with E. In contrast, a cold-shock expression in thermophiles could be accomplished using ambient water or air as a heat acceptor. By far the most significant benefit of thermophilic production is expected to be minimized contamination risk.
Biorefineries experience two main types of contamination. Chronic, low level infections can reduce yields, but at large scale they are so difficult to prevent that they have been accepted as a fact of life at many corn ethanol plants.
The harsh pre-treatments sometimes used to solubilize biomass would be expected to eliminate wild-type bacteria that might compete with the production host, but contamination remains a major industrial problem. The load of bacterial contaminants was found to be comparable between two dry-grind ethanol plants that incorporated a high-temperature saccharification pre-treatment, and a wet-mill plant which did not Skinner and Leathers, Bacterial load was also not significantly affected by differing degrees of antibiotic use among the three processing plants, although the diversity of bacterial contaminants declined with increased antibiotics.
Therefore, having a sterile feedstock or antibiotics in the bioreactor is not enough to prevent opportunistic bacteria from taking advantage of the abundant nutrients and mild conditions found in a bioreactor operated under mesophilic conditions.
The use of an extremely thermophilic host is expected to reduce the risk of bacterial contamination in biorefineries, but the possibility of viral infections remains. Viruses affecting both bacterial and archaeal hyperthermophiles have been identified, although interestingly the archaeal species infected are limited to the crenarchaeoata Pina et al. No virus has yet been reported for a member of the hyperthermophilic euryarchaeota, although the presence of a CRISPR system in both T. This indicates that T.
One of the oft-cited concerns for the use of thermophiles in industrial fermentations is the energy required to heat the process. There are two major thermal energy requirements for an industrial fermentation: sterilization and fermentor temperature maintenance. For a thermophilic fermentation, the sterilization process heat inputs would be no higher, and could potentially be reduced; hardy bacterial spores may be able to survive exposure to high temperatures, but they cannot grow at them.
Fermentor temperature maintenance actually provides an opportunity for energy savings if a thermophile is used. All organisms produce heat as a byproduct of metabolic processes. At large scales, the metabolic heat produced outweighs heat lost to the environment through the fermentor walls or evaporation Yang et al. As a result, cooling is required to maintain the fermentor at a constant temperature. The cooling duty for a large bioreactor with a high-density culture is extremely high, and can be one of the limiting factors for fermentation scale-up Yang, A further difficulty for mesophilic fermentations is the small thermal driving force between the fermentation temperature and the ambient environment, which limits heat removal, often making refrigeration necessary, a further energy cost Abdel-Banat et al.
The metabolic heat generated, and thus the heat removal required, are primarily dependent on the metabolic activity of the culture and not on the fermentor temperature or organism used Blanch and Clark, Thus, a thermophilic fermentation would require a similar amount of heat removal as a mesophilic fermentation.
Furthermore, this heat would be much easier to remove because of the large temperature differential between a thermophilic fermenation and the environment, providing the possibility for substantial cost savings Abdel-Banat et al.
An additional opportunity for energy savings in thermophilic industrial fermentations is product separation, which can be the most energy intensive part of a process, since it is often carried out at elevated temperatures.
In particular, thermophilic production of volatile products, such as fuel alcohols, allows for the possibility of facilitated product removal. The use of thermophilic organisms would be a favorable match for most separations processes recovering volatile products from fermentation broth, including distillation, gas stripping, and pervaporation Vane, While metabolic engineering opens up the possibility of engineering desired chemical pathways into any genetically tractable host, it is worth remembering that S.
Just because the appropriate enzymes can be inserted into an organism does not mean the resulting mutant will be industrially useful. Therefore, desirable hosts should be selected not only for what they can be engineered to do, but also for what they already do well.
Fortunately, the group of extreme thermophiles discussed above exhibit a variety of desirable properties natively. Many are capable of metabolizing a diverse set of sugar polymers and monomers, and C. Coupled with production of ethanol as either a minor or major natural metabolite, this makes them promising candidates for bio-ethanol production from non-food feedstocks.
Fuels are typically highly reduced organic molecules, so various efforts to maximize biofuel titers have focused on tuning the redox pathways within mesophilic hosts to favor the production of reduced end products Liu et al.
More oxidized products, such as lactic acid for production of biodegradable plastics, can be selected for by shifting metabolism in the other direction Lee et al. Substantial progress toward redox-tuning in thermophiles has already been made. The redox pathways of several extreme thermophiles are well understood Schut et al. The ability to select for the production of alcohols, rather than organic acids, has been demonstrated in recombinant P. While anaerobes like P. This causes fermentative anaerobes to exist in a constant state of energy limitation, where even seemingly minor energetic costs, such as export of a final product, can be problematic Maris et al.
Therefore, expression in anaerobic hosts should be focused on pathways that are either energy-neutral or energy-yielding.
It is now apparent that fossil fuels cannot continue to be used at their current rate without causing irreparable environmental harm.
The shift away from petroleum will necessitate dramatic changes to the current motor-fuel regime, but will also significantly alter the production of plastics, solvents, and other specialty chemicals that are currently generated in chemical refineries. Extreme thermophiles, because of their unique advantages and the recent expansion of genetic systems allowing for metabolic engineering, are perfectly positioned to fill the need for massive chemical production from renewable feedstocks.
They are able to survive the high temperatures that can result from heat generated in large-scale bioreactors, and when operated at these temperatures are less likely to be contaminated by ambient microorganisms and phages.
Many also exhibit unique metabolic properties as a product of their extreme environment. Much work remains to be done before the promise of using thermophilic hosts to produce large quantities of renewable fuels and chemicals can be realized, but the genetic tools are now in place to allow that work to be carried out. 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.
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Adams, M. Key role for sulfur in peptide metabolism and in regulation of three hydrogenases in the hyperthermophilic archaeon Pyrococcus furiosus. Ahring, B. Methanogenesis in thermophilic biogas reactors. Anton Leeuw 67, 91— Production of ethanol from wet oxidised wheat straw by Thermoanaerobacter mathranii.
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FEMS Microbiol. Atomi, H. Description of Thermococcus kodakaraensis sp. Archaea 1, — Biotechnological applications of thermophiles Thermophiles have shown tremendous promise in terms of their applications in modern biotechnology. Bioconversion of lignocellulose to hydrogen Although many reported microorganisms possess the capability of cellulose hydrolysis or hydrogen production H 2 , no conclusive research has been able to clarify that both of these capabilities are possessed in a single microorganism.
Conversion of glycerol to lactate Bioprospecting efforts for exploring novel biocatalytic molecules with unique properties have inspired the design and construction of a wider variety of artificial metabolic pathways Bond-Watts et al.
Conversion of d -xylose into ethanol Thermophilic anaerobic bacteria could be promising candidates for conversion of hemicellulose or its monomers xylose, arabinose, mannose and galactose into ethanol with a satisfactory yield and productivity. Biodegradation of petroleum hydrocarbons Thermophiles have also been utilized for the microbial degradation of crude oil and refined petroleum pollutants.
Recovery of heavy metals As a result of increasing industrial activities, heavy metal contamination is a problem. Remediation of textile dyes Laccase enzyme purified from thermophile, Geobacillus thermocatenulatus MS5 is of very higher catalytic activity and are economic, highly stable at different temperatures and pH levels and can be used widely and effectively in the removal of the dyes that cause environmental pollution.
Saccharification of agricultural residues Sporotricum thermophile LAR5 is an excellent fungal isolate having an ability to utilize crude agriculture based materials as carbon and nitrogen sources to produce significant cellulase titre. Thermophilic bacilli in dairy processing Thermophilic bacilli are used as hygiene indicators of processed product, within the dairy processing context. Cancer treatment Asperjinone, a nor-neolignan, and Terrein, a suppressor of ABCG2-expressing breast cancer cells were isolated from thermophile Aspergillus terreus, which can restore drug sensitivity and could be the key to improve breast cancer therapeutics.
Conclusions The increasing number of patents indicates that there is a growing interest in the commercial applications of thermophiles. Acknowledgments Authors would like to acknowledge M. Compliance with ethical standards Conflict of interest There exists no conflict of interest regarding publication of this manuscript. Exploring the biotechnological applications in the archeal domain.
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