Ann Microbiol (2014) 64:421–429 DOI 10.1007/s13213-013-0699-5

REVIEW ARTICLE

Potential applications and emerging trends of of the genus : a review

Seyed Mohammad Taghi Gharibzahedi & Seyed Hadi Razavi & Mohammad Mousavi

Received: 19 May 2013 /Accepted: 25 June 2013 /Published online: 28 August 2013 # Springer-Verlag Berlin Heidelberg and the University of Milan 2013

Abstract Interest in attractive biological sources with et al. 2002), human clinical specimens (Yassin et al. 2006; multicriteria applications has been increasing during recent Jones et al. 2008; Kämpfer et al. 2012), plant tissues (Li et al. years. This study scrutinized the applications of Dietzia bac- 2008), soils (Mayilraj et al. 2006;Lietal.2009; Yamamura teria for future prospects. Apart from such present and well- et al. 2010), the air in a duck barn (Kämpfer et al. 2010)anda established applications—as in therapeutic biotreatments for traditional Korean food (Kim et al. 2011). adult paratuberculosis animals, production of carotenoid pig- Dietzia maris, D. natronolimnaea, D. psychralcaliphila, D. ments, and animal feed additives—their uses in biosurfactants cinnamea, D. kunjamensis, D. schimae and D. cerdiciphylli, and biodemulsifiers production, the pollutants bioremedia- D. papillomatosis, D. lutea, D. aerolata, D. timorensis, D. tion, biodegradation of petroleum hydrocarbons and crude alimentaria and D. aurantiaca are thirteen species from this oil and also production of extracellular polymeric substances genus at the time of writing. Many researchers showed that (EPSs) have been exploited. The use of these as a these bacteria are Gram-positive, aerobic, short rod- and biotechnological tool may lead to improve the optimization coccoid-like, non-motile, non-endospore forming, non-acid and quality assurance of food ingredients and products, the fast, oxidase-positive and catalase-positive. The colonies capability of degradation and remediation of environmental morphology of these species was small, smooth, circular pollutants, and the efficiency of bioconversion systems for and convex. Optimum growth temperature and pH for the energy recovery and bioprocessing of value-added products. different strains also were 25–30 °C and 7–8, respectively. Among all of the species, D. timorensis ID05-A0528T had Keywords Dietzia . Bioremediation . Biosurfactant . the lowest tolerance level to NaCl. This strain also was able Industrial fermentation . Carotenoid pigmentation to utilize a wide range of compounds individually as a sole carbon source. Short-chain mycolic acids were present in these bacteria. The cell wall contained meso-diaminopimelic Introduction acid, arabinose and galactose; the glycan moiety of the cell wall contained acetyl residues. The DNA G+C contents of Dietzia spp. are broadly distributed in nature. These bacteria strains ranged from 64.7 (D. alimentaria 72T) to 73 mol % have been isolated from widely different environments, even (D. maris DSM 43672T). The most important phospho- including the deepest sea mud of the Mariana Trench lipids in these strains were diphosphatidylglycerol, (Takami et al. 1997), deep-sea sediments (Colquhoun et al. phosphatidylglycerol, phosphatidylinositol mannoside, 1998), an alkaline East African soda lake (Duckworth et al. phosphatidylinositol and phosphatidylethanolamine. The 1998), a drain pool of a fish-egg processing plant (Yumoto species D. schimae (2.9 %), D. cinnamea (8.3 %), D. timorensis (48.0 %), D. aurantiaca (25.9 %), and D. psychralcaliphila (13.9 %) had the highest amounts of C , : : 14:0 S. M. T. Gharibzahedi S. H. Razavi (*) M. Mousavi C15:0,C16:0,C17:0,andC18:0 fatty acids, respectively (Rainey Bioprocess Engineering Laboratory (BPEL), Department of Food et al. 1995; Gharibzahedi et al. 2013a). Science, Engineering & Technology, Faculty of Agricultural Some strains identified as representing species of the Engineering and Technology, University of Tehran, P.O. Box 4111, Karaj 31587-77871, Iran genus Dietzia are potential human pathogens in immuno- e-mail: [email protected] competent (Pidoux et al. 2001) and immunocompromised 422 Ann Microbiol (2014) 64:421–429

(Click and Van Kampen 2010) patients. These bacteria also and most importantly, the lack of a successful protective have many applications in many industries, especially the vaccine or therapeutic drug treatment (Click and Van medical, chemical and food industries. Click and Van Kampen 2009; Click and Van Kampen 2010; Click 2011). It Kampen (2010) reported a number of strains of these bac- accounts a great economic problem on dairy, sheep and goat teria could be used as potential probiotic to inhibit farms in throughout worldwide (Click and Van Kampen Mycobacterium avium, subspecies paratuberculosis (MAP) 2010). under in vitro culture conditions. These strains, compared It is expected that the use of Dietzia bacteria as probiotics to the antimicrobial drugs, have fewer medical complica- with similar activities to antimicrobial drugs will result in tions. Some of the Dietzia species described to date have less medical complications. Recently it was reported that been revealed to degrade aliphatic hydrocarbons such as n- Dietzia subsp. C79793-74 as a probiotic could inhibit devel- alkanes (Alonso-Gutiérrez et al. 2011;Biharietal.2011; opment of factors indicative of bovine paratuberculosis after Rainey et al. 1995;Yumotoetal.2002). There have also potential in utero, birthing and colostrum exposure to MAP. In been reports on the aromatic compounds degradation by other words, the daily probiotic treatment of Dietzia success- Dietzia strains (Bødtker et al. 2009). Iwaki et al. (2008) fully increased the survival of cows with early-stage Johne’s found that a Dietzia strain isolated from enrichment cultures disease and in certain cases cured the animal (Click 2011). was able to utilize cyclohexylacetic acid. The surface-active compounds production related to hydrocarbon degradation has been reported (Banat 1995; Mukherjee et al. 2008). Production and application of natural pigment Takeishi et al. (2006) reported xylanolytic strains of the genus Dietzia isolated from the hindgut and faeces of Trypoxylus To date, attempts have been made to formulate the fermen- dichotomus larvae. Rashidi et al. (2009) with the effective tation media and optimization of environmental culture con- biotransformation of delta 9-tetrahydrocannabinol (Δ9-THC) ditions for increasing the production of carotenoid pigments by Dietzia sp. ENZHR1 concluded that alkane oxygenases by Dietzia sp. It was demonstrated that the CTX (4,4′- of Dietzia spp. can have a significant role in the produc- diketo-β-carotene) is the most common carotenoid produced tion of novel pharmaceuticals. Dietzia strains are the most by Dietzia bacteria. Identification of CTX as predominant promising sources for microbial production of carotenoid carotenoid in these microorganisms was previously carried pigments especially canthaxanthin (CTX) used in the dif- out using an ultraviolet (UV)-HPLC/atmospheric pressure ferent industries such as nutraceuticals, cosmetics, food chemical ionization-mass spectrometry (APCI-MS) method and feed industries (Goswami et al. 2012; Khodaiyan (Razavi et al. 2006). D. natronolimnaea HS-1 and D. maris et al. 2007; Khodaiyan et al. 2008; Nasri Nasrabadi and NIT-D identified as the best microbial strains in the CTX Razavi 2010a, c; Gharibzahedi et al. 2013b). production (Khodaiyan et al. 2007; Khodaiyan et al. 2008; Since there is no review article covering the complete Nasri Nasrabadi and Razavi 2010a, c;Gharibzahedietal. comprehensive information of possible applications of 2012b;Gharibzahedietal.2013b). The production and con- bacteria of the genus Dietzia, an attempt is made to write sumption of CTX has increased due to its beneficial health this review article with a main perspective on the use of attributes such as anti-oxidative, anti-inflammatory, anti- bacteria of this genus for creation of innovative industrial tumor, anticancer, or anti-dermatosis, and coloring agent applications and products, and to provide some view- (Gharibzahedi et al. 2012a). points on the current situation and suggestions for future Glucose was an excellent carbon source for the research directions. growth and CTX production by D. natronolimnaea HS-1. However, fructose, sucrose, mannose and also whey lactose were suitable carbon sources for the Potential applications growth and production of CTX by D. natronolimnaea HS-1 in a batch bioreactor. The combination of 10 g/l A summary of potential applications of the different strains peptone and 6 g/l yeast extract was the best nitrogen of the genus Dietzia is presented in Table 1. source among the different studied sources (Khodaiyan et al. 2007). However, Khodaiyan et al. (2008)inan- other investigation found that yeast extract revealed the Biological therapeutic application highest CTX production followed by casein acid hydro- lysate, bactotryptone, peptone and meat extract, respec- MAP can cause a chronic inflammatory bowel disease, tively. The optimum conditions to achieve the highest Johne’s disease, in ruminant animals. This disease may be CTX production (2.87 mg/l) in a batch bioreactor could attributed to the animal movement from one farm to another be obtained by using 55.54 g/l of whey lactose concen- farm, farm intensification and confinement, herd expansion tration and 7.36 g/l of yeast extract concentration. n irbo 21)64:421 (2014) Microbiol Ann

Table 1 Useful applications of the different strains of the genus Dietzia

Dietzia strain Application type Reference

Biological Biocolourant Biosurfactant Biodemulsifier Biodegradation/ therapeutic synthesis production production Bioremediation

Dietzia subsp. C79793-74 Bovine paratuberculosis –– – – Click and Van Kampen (2010) –

inhibition Click (2011) 429 D. natronolimnaea HS-1 – CTX production –––Khodaiyan et al. (2007) D. natronolimnaea HS-1 – CTX production –––Khodaiyan et al. (2008) D. natronolimnaea HS-1 – CTX production –––Nasri Nasrabadi and Razavi (2010a, b, c) D. natronolimnaea HS-1 – CTX production –––Gharibzahedi et al. (2012b) D. maris NIT-D – CTX production –––Goswami et al. (2012) D. maris WR-3 ––Biosurfactant production – n-alkane biodegradation Nakano et al. (2011) in culture containing n-hexadecane and NaNO3 Dietzia sp. S-JS-1 ––– Biodemulsifier synthesis Liu et al. (2009) in cultures containing waste frying oil and paraffin D. cinnamea strain P4 –––– n-alkane biodegradation Von der Weid et al. (2007) Dietzia sp. A14101 –––– n-alkane biodegradation Bødtker et al. (2009) Dietzia sp. E1 –––– n-alkane biodegradation Bihari et al. (2011) D. maris DSM 43672T –––– n-alkane biodegradation Rainey et al. (1995) D. psychralcaliphila –––– n-alkane biodegradation Yumoto et al. (2002) DSM 44820T D. natronolimnaea –––– n-alkane biodegradation Yassin et al. (2006) DSM 44860T Dietzia sp. CBMAI 705 ––Biosurfactant production – n-alkane biodegradation and Vasconcellos et al. (2011) MEOR processes D. maris AM3 –––– Bioremediation of soils Pleshakova et al. (2008) polluted with crude oil D. maris strain 53 –––– Hydrocarbon biodegradation Alvarez (2003) Dietzia DQ12-45-1b –––– Biodegradation of Wang et al. (2011) n-alkanes, crude oil and aromatic compounds D. psychralcaliphila –––– n- and branched alkane- Yumoto et al. (2002) Biodegradation Dietzia H0B –––– n-alkane biodegradation Alonso-Gutiérrez et al. (2011) D. natronolimnaea JQ-AN –––– Aniline biodegradation Jin et al. (2012) 423 424 Ann Microbiol (2014) 64:421–429

Gharibzahedi et al. (2012b), using the selection of su- and 120 rpm, initial pH and percentage inoculum being 5.5 and perior mutant type in combination with optimization of 2 %, respectively. nutrient medium components, demonstrated that the op- CTX are applied commercially as food colorants, timization of D-glucose, mannose and Fe3+ concentra- nutraceuticals, and for potential industrial and pharma- tions can lead to increase of the CTX biosynthesis ceutical applications. However, it like most carotenoids is (7.67 mg/l) in a batch culture by the mutant strains a highly unsaturated molecule, and thus is very susceptible to induced by ethyl methane sulfonate (EMS). The authors environmental conditions. Hojjati and coauthors during 2011– showed that the mutants induced by EMS had higher 2012 successfully used a spray-drying technique to microen- survival and growth rates compared to UV mutants. The capsulate and enhance the light, thermal and oxidative stabil- appropriate color and colony properties obtained from ity of CTX produced by D. natronolimnaea HS-1. The differ- method of EMS mutagenesis also showed moderate to ent wall materials such as gum arabic, maltodextrin and sol- high growth rates. uble soybean polysaccharide (SSPS) were applied. They Nasri Nasrabadi and Razavi (2010a) also found that the use proved that the best microencapsulation efficiency was of optimum amounts of three ions of Fe3+ (30 ppm), Cu2+ obtained when the ratio of CTX/SSPS was at minimum level (28.75 ppm) and Zn2+ (27 ppm) can cause enhancement of the of this study (0.25). Moreover, light and high temperatures CTX production (8,923 μg/l) by D. natronolimnaea HS-1 in a were drastic factors to the stability of microencapsulated sam- fed-batch fermentation process. They also reported that the ples because of the acceleration of carotenoids degradation CTX biosynthesis by D. natronolimnaea HS-1 is not affected (Hojjati et al. 2011; Hojjati et al. 2012). Gharibzahedi et al. by heavy metal of cobalt. However, Khodaiyan et al. (2008) (2012a) developed an emulsion model system containing found that KH2PO4 content on CTX production by this CTX produced by bacterium D. natronolimnaea HS-1. They bacterium had a negative effect. These researchers have found that a combination of fenugreek gum (0.49 % w/w), recently reported their effort to increase the CTX produc- coconut oil (CO, 6.28 % w/w) and CO/CTX ratio (50:1) can tion by this bacterium in a fed-batch bioreactor by optimi- lead to produce the stable CTX emulsions. Gharibzahedi et al. zation of tricarboxylic acid (TCA) cycle intermediates. The (2013c) using RSM determined the optimum formulation for results showed that three TCA cycle intermediates, namely production of a stable O/W emulsion containing CTX alpha-ketoglutarate, oxaloacetate and succinate, have a signif- biosynthesized by D. natronolimnaea HS-1 using 25 kHz icant effect on the CTX production and cell biomass (Nasri ultrasonic emulsification. They found that the most suitable Nasrabadi and Razavi 2010c). They also investigated the combination of variables for the higher stability (98.6 %) and influence of lycopene cyclase inhibitors on carotenogenesis CTX entrapment efficiency (93.1 %) was 1.20 % w/w, in order to achieve high-level accumulation of lycopene 3.30 % w/w and 5.43 % w/w for whey protein isolate (WPI), (8.26 mg/l) in the bacterium D. natronolimnaea HS-1 cul- psyllium husk gum (PHG) and CO concentrations, respectively. tured in a fed-batch process. The optimum set of the indepen- This natural pigment can also use as a valuable dent variables for the highest lycopene level using response supplementary for the aquaculture and animal feeding surface methodology (RSM) was obtained 24.74 ppm imid- applications. CTX has been approved for use in fish azole, 28 ppm nicotinic acid, 24.05 ppm pyridine, 27.6 ppm feed, for example, in the EU (maximum 25 mg/kg) and piperidine and 23.22 ppm triethylamine (Nasri Nasrabadi and in the US (maximum 80 mg/kg) (Breithaupt 2007). Razavi 2010b). Since fishes such as salmon are able to transport and The production and accumulation of CTX by D. deposit this pigment at specific sites in their muscle, natronolimnaea HS-1 can notably increase by the inten- the commercial preparations of this pigment in the sity of illumination and white-light irradiation due to manufacture of pelleted feeds could be possible for this fact that light affect as a stimulant on the microor- aquaculture industries (Baker 2002). Esfahani- ganism growth and the activity of existing enzymes in Mashhour et al. (2009) evaluated the influence of dif- carotenoid biosynthesis (Khodaiyan et al. 2007). The ferent levels of extracted pigment from D. optimal temperature and pH for this CTX-containing natronolimnaea HS-1 biomass as a CTX source in microorganism were 31 °C and 7, respectively. Moreover, comparison with synthetic CTX on egg yolk pigmenta- the biomass and CTX production by bacterial strain of D. tion. The results showed that hens feeding with a diet natronolimnaea HS-1 was enhanced in a medium without containing CTX produced by D. natronolimnaea HS-1 led NaCl (Khodaiyan et al. 2007). Goswami et al. (2012)also to a significant effect on the color of egg yolks and color by studying the effects of four process parameters including scores of egg yolks from this treatment were higher com- temperature, shaker speed, pH and percentage inoculum on pared with the control group. They demonstrated that the the biomass and CTX yield found that the CTX biosynthesis CTX biosynthesized by this bacterium is a proper natu- by D. maris NIT-D increased up to 121.6 mg/l in a batch ral pigment which could create suitable egg yolk color bioreactor when this bacterium was incubated for 120 h at 25 °C (Esfahani-Mashhour et al. 2009). Ann Microbiol (2014) 64:421–429 425

Biosurfactant and biodemulsifier production low production yield, high cost, and functionality (Banat 1995; Huang et al. 2009). A great amount of the high cost of Biosurfactants or microbial surfactants are surface-active biodemulsifier production was contributed to the cost of cul- compounds synthesized by a wide group of microorganisms. ture medium, especially carbon sources (about 30–50 %). They are amphiphilic molecules containing hydrophobic and Therefore, increase of the volumetric production and use of hydrophilic moieties and tend to interact with surfaces of cheap carbon sources can be solved the economic problems in different polarities and reduce the surface and interfacial production of biodemulsifier (Liu et al. 2009). For most of the tension of solutions (Mukherjee et al. 2008). Some of the demulsifying strains, hydrophobic substrates are preferred as advantages, which make biosurfactants promising alterna- carbon source in the synthesis of biodemulsifiers, such as tives to chemically synthesized surfactants, are their high crude oils, tetradecane, cetane, hexadecane and kerosene stability at extremes of pH, salinity and temperature, low (Singh et al. 2007). toxicity, high foaming, low irritancy, high biodegradability, Liu et al. (2009) found that Dietzia sp. S-JS-1, a Dietzia suitable environmental compatibility, and high compatibility strain isolated from petroleum contaminated soils (Huang with human skin (Banat 1995). Nakano et al. (2011) for the et al. 2009), was able to produce the biodemulsifier. They first time reported that D. maris WR-3, as a marine bacterium compared two cheap carbon sources including waste frying of the family Dietzia, can reduce the surface tension of oil (WFO-II) and paraffin in the biodemulsifier synthesis. culture broth to 31 mN/m when cultured using n-hexadecane This strain had 93 % of homology with D. natronolimnaea

(as carbon source) and NaNO3 (as nitrogen source). The wax strain LL 51 (DQ821754) and D. psychralcaliphila strain ester-like compounds were evaluated as main surface-active JCM 10987 (AB159036), and 92 % homology with D. compounds produced by this strain using high-performance natronolimnaea strain TPL19 (EU373398) according to the thin-layer chromatography (HPTLC) and gas chromatogra- phylogenetic analysis of 16S rRNA (Huang et al. 2010). The phy–mass spectrometry (GC-MS). These biosurfactant com- biomass concentration after 7 days of cultivation with WFO- pounds and their role were probably originated the degrada- II was higher than the same amount of paraffin as carbon tion process of n-alkane and low water-solubility of the source (2.4 times). The efficiency of produced biodemulsifier substrates, respectively (Nakano et al. 2011). Thus, the in- with WFO-II culture to break the emulsions was more than its creased attachment of cells to hydrocarbons via enhanced counterpart in paraffin culture under the same cultivation cell hydrophobicity can lead to accelerate bioavailability and conditions. In this case, oil separation ratio in W/O emulsion biodegradation of water immiscible substrates. and water separation ratio in O/W emulsion within 5 h were Biodemulsifiers produced by microorganisms can break obtained 88.3 and 76.4 %, respectively (Liu et al. 2009). water-in-oil (W/O) emulsions produced in the recovery and processing of crude petroleum, because quantity rather than purity matters in this application. In order to separate Bioremediation and biodegradation applications water from oil phase of these emulsions, high quantities of biodemulsifiers are needed (Liu et al. 2009). Also, these Crude oil is a series of complex hydrocarbons with low demulsifiers have the potential to be widely used in many bioavailability, and is persistent in soil. Crude oil pollution commercial applications in the environmental protection, met- is one of the most important environmental problems faced allurgy, transportation, textile, biomedical, food processing by oil-producing countries of the world, especially middle and pharmaceutical industries (Li et al. 2012). eastern countries. Since most oil spillages are due to anthro- Previous observations showed that bacteria belonging to pogenic causes such as pipeline vandalism, quantification of the genera Nocardia (N. amarae), Corynebacterium (C. real contamination details of the global ecosystem by oil in petrophilum; C. lepus), Rhodococcus (R. aurantiacus; R. these countries is difficult (Wang et al. 2012). In the USA, erythropolis; R. rhodochrous), Bacillus (B. mojavensis the cost of bioremediation to clean up petroleum hydrocar- XH1; B. subtilis), Pseudomonas (P. aeruginosa MSJ;P. bon contaminated soils is expected to exceed US$1 trillion paucimobilis), Torulopsis (T. bombicola), Alcaligenes (A. per year (Stroud et al. 2007). Thus, development of appropriate latus), Micrococcus sp., and Mycobacterium sp., have been methods to clean up contaminated environments continues examined in demulsification investigations with satisfactory to be a vital issue in terms of environmental restoration and results (Huang et al. 2010; Li et al. 2012). protection. Compared to conventional chemical demulsifiers, biologi- Duetotheprocesscomplexityinnaturalmedia,the cal demulsifiers are generally characterized by following at- efficiency of remediation methods has depending on many tributes: good surface activity, low toxicity, ecological safety factors such as pollutant composition, concentration, and and easy biodegradability, and high demulsifying efficiency in exposure time; soil type; and other ambient factors: tem- extreme conditions. However, there are some limiting factors perature, moisture, pressure, atmospheric conditions, etc. that continue to prevent their practical application, including (Pleshakova et al. 2008). However, microbial remediation 426 Ann Microbiol (2014) 64:421–429 is a better method than the physical and chemical methods Efficiencies of introduction of an oil-oxidizing D. maris because of its cost effectiveness along with low environ- strain and stimulation of natural microbial communities in mental impact. Bioaugmentation involving introduction of remediation of polluted soils by Pleshakova et al. (2008) cultured microbial degraders and biostimulation based on were investigated. They showed that the addition of D. maris growth and activity stimulation of usual microorganisms in AM3 to soil freshly polluted with oil accelerated its remedi- polluted soils are two common techniques for the microbial ation twofold due to high dehydrogenase and catalase activ- remediation (Wang et al. 2012). The marine ecosystems ities within the first month in comparison with the stimula- harbor bacteria such as Pseudomonas, Pseudoalteromonas, tion. This strain had not significant differences in the rate of Kocuria, Marinobacter, Mycobacterium, Psychrobacter, oil degradation for soils with aged pollution. Therefore, the Isoptericola, Rhodococcus, Alcanivorax, Dietzia,etc.are authors pointed out stimulation of the aboriginal microflora known for their hydrocarbon-degradation potential (Al- in the case of aged pollution are preferable. Awadhi et al. 2007; Harwati et al. 2007). Extracellular Alvarez (2003) investigated relationship between β- polymeric substances (EPSs) obtained from the cellular oxidation pathway and the hydrocarbon-degrading pro- degradation of hydrocarbons have a vast potential for file in actinomycetes bacteria like D. maris strain 53. The using in biotechnological processes such as microbial studied hydrocarbons were alkylcycloalkanes, alkylbenzenes, enhanced oil recovery (MEOR) and bioremediation. cycloalkanes, normal- and branched-alkanes, polyaromatics Dietzia strains are identified as interest bacteria increasing and monoaromatics. These bacteria had more affinity to hydrocarbon-degrading activities of the surrounding bacteria degrade pentadecane, hexadecane, 2,6,10,14-tetramethyl increasing the hydrogenase and catalase activities of other pentadecane (pristane), phenyldecane, gas–oil and kero- key oil-degrading species, and therefore speeding up the sene which were contained an alkyl-side chain suscepti- process of biodegradation (Bihari et al. 2011; Pleshakova ble to be degraded through β-oxidation pathway. Author et al. 2008) and for their long-term viability in the environ- indicated that these hydrocarbons were applied to biosynthesis ment even under dry, resource-limited conditions (Radwan cellular lipids such as triacylglycerols during nutrient starva- et al. 2010). The n-C11 to n-C36 alkanes, n-C6 to n-C26 tion conditions. Therefore, D. maris strain 53 was able to alkanes and n-C12 to n-C38 alkanes as sole carbon and transform hydrocarbons into cellular lipids under unbalanced energy source during aerobic growth were degraded by D. growth conditions. This fact makes it potential candidates for cinnamea strain P4, Dietzia sp. A14101 and Dietzia sp. E1, the bioremediation of contaminated environments. respectively (Bihari et al. 2011; Bødtker et al. 2009; von der Wang et al. (2011) for the first time reported the various Weid et al. 2007). Prior to this, it was known that n-C6 to behaviors for a bacterial genus to degrade crude oil and a n-C23 alkanes, n-C13 to n-C24 alkanes and paraffin can wide range of hydrocarbons. They isolated Dietzia DQ12- consume by D. maris DSM 43672T (Rainey et al. 1995), 45-1b which had the highest 16S rRNA sequence similarity D. psychralcaliphila DSM 44820T (Yumoto et al. 2002) with D. cercidiphylli (99.35 %). This genus was able to use and D. natronolimnaea DSM 44860T (Yassin et al. 2006), a broad range of n-alkanes (C6–C40), crude oil and aro- respectively. matic compounds (such as benzoate, carbazole, fluoran- In recent years, many research groups focused on the thene, naphthalene phenanthrene, quinoline and toluene) identification of new strains of Dietzia bacteria with hydro- as the sole carbon sources for growth. Nie et al. (2011) carbons utilization and their bioremediation potential (Wang identified two alkane hydroxylase-rubredoxin fusion gene et al. 2011). homologs including alkW1 and alkW2 from this strain were Vasconcellos et al. (2011) found that Pseudomonas identified. They found that the cloned gene of alkW1 in- sp. CBMAI 754 and Dietzia sp. CBMAI 705 isolated creased growth on and degradation of n-alkanes up to C32 from a Brazilian petroleum reservoir among the studied in length. However, degradation of aromatic compounds strains had the highest tendency for hydrocarbon bio- such as benzoate (Maeda et al. 1998), carbazole, quinoline, degradation. However, both strains could not deplete fluoranthene (Kumar et al. 2011), phenanthrene (Brito et al. phenanthrene. This fact showed that this single com- 2006; Al-Awadhi et al. 2007), disodium terephthalate pound is possibly being co-metabolized by both strains (Sugimori et al. 2000), naphthalene and toluene (von der through the n-alkane biodegradation pathway. Moreover, Weid et al. 2007; Bødtker et al. 2009) were previously the supernatant of the Dietzia sp. CBMAI 705 growth reported. culture compared to other evaluated strains had the Yumoto et al. (2002) showed that D. psychralcaliphila highest indices of surface tension reduction and the was an n- and branched alkane-degrading bacterium. Dietzia suitable emulsification of hydrophobic compounds. H0B is a quite similar genus to D. psychralcaliphila by Therefore, these multiple abilities make Dietzia sp. studying 16S rRNA gene sequence information (Alonso- CBMAI 705 an interesting candidate for future applica- Gutiérrez et al. 2011). Alonso-Gutiérrez et al. (2011) report- tion in bioremediation and MEOR processes. ed that Dietzia H0B was capable to grow on n-alkanes Ann Microbiol (2014) 64:421–429 427

ranging from C12 to C38 of crude oil and branched alkanes degradation pathways and organic compounds biotransfor- such as pristane and phytane. They found that the present of mation are still limited. Therefore, further investigations will 8-hexadecene as an intermediate of hexadecane degradation be conducted in order to understand the various interactions by this bacterium is a new alkane-degrading pathway. and microbial mechanisms that take place during hydrocar- Jin et al. (2012) isolated D. natronolimnaea JQ-AN from bon biodegradation processes. Moreover, further studies in industrial wastewater which was degraded 87 % of the ani- order to expose the relationship between carbon sources and line in a 300 mg/l aniline solution using an ortho-cleavage the chemical composition of produced biosurfactants and pathway with catechol intermediate. This bacterium had a biodemulsifiers by the Dietzia strains as well as their emul- higher homology with the aniline-degraders of the genus sification and demulsification abilities are needed. Rhodococcus among most aniline-degraders. The optimal pH and salinity for aniline biodegradation were pH 8.0 and Acknowledgments This work was supported by a grant from the 0–6 % (w/v) NaCl. Therefore, low concentrations of sodium University of Tehran. The authors would like to thank the constructive comments of two anonymous reviewers. acetate (40 mM) had a stimulating effect on the degradation. It is demonstrated that R. koreensis sp. nov. (Yoon et al. Conflict of interest The authors have declared no conflict of interest. 2000), D. papillomatosis sp. nov. (Jones et al. 2008) and Dietzia spp. (Bihari et al. 2011) can also apply sodium acetate to use as an easily assimilable carbon source for the improved microbial growth. 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