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Haloarcula Marismortui, Eighty-Four Years After Its Discovery in The International Journal of Engineering Research & Technology (IJERT) ISSN: 2278-0181 Vol. 3 Issue 6, June - 2014 Haloarcula Marismortui, Eighty-Four Y ears after its Discovery in the Dead Sea, Review David Isidoro Camacho Córdova Juan Carlos Mateos Díaz Departamento de Ing. Química, CUCEI, Universidad de Centro de Investigación y Asistencia en Tecnología y Guadalajara. Diseño del Estado de Jalisco A.C. CIATEJ Guadalajara, Jalisco, Mexico Guadalajara, Jalisco, Mexico Rosa María Camacho Ruíz Jesús Antonio Córdova López Centro de Investigación y Asistencia en Tecnología y Departamento de Química, CUCEI, Universidad de Diseño del Estado de Jalisco A.C. CIATEJ Guadalajara. Guadalajara, Jalisco, Mexico Guadalajara, Jalisco, Mexico Jorge Alberto Rodríguez González Centro de Investigaci ón y Asistencia en Tecnología y Diseño del Estado de Jalisco A.C. CIATEJ Guadalajara, Jalisco, Mexico Abstract— Haloarcula marismortui is one of the few halophilic archaea are well adapted to proliferate in media halophilic microorganisms from the Archaea domain to have containing NaCl concentrations close to saturation. been thoroughly studied. This microorganism has been Halophiles are disseminated over the world, in hyper-saline considered as a model for many researcher groups to environments, such as natural hypersaline brines in arid, understand how it is able to proliferate at extremely high salt coastal and deep-sea locations as well as in artificial salterns concentration. Such studies include the purification of the first used as source of salts from the sea [3]. The use of halophiles haloarchaeal enzyme, the estimation of intracellular salt for industrial purposes is expanding. Many potential uses concentrations, the first crystal structure for haloarchaeal have been proposed and the number of patents issued has ribosome, the sequencing of the second haloarchaeal genome,IJERTIJERT increased. Due to the necessity for more environmentally the sizeable (4.27 Mb) and incredibly complex genome friendly technologies, halophilic microorganisms are architecture, consisting of nine replicons, and many more perceived as a source for enzymes [4], β–carotene and ectoine examples that continue to providing interest insight into this (used as a stabilizer for enzymes and for cosmetic products) microorganism. This review discusses and analyzes essential publications concerning Haloarcula marismortui to be taken and they are also utilized to produce traditional fermented account for future works with other halophilic microorganisms foods [5]. and an additional effort is made to collect physiological Haloarcula marismortui is a halophilic archaeon isolated characteristics and biotechnological applications of this from the Dead Sea by Elazari-Volcani in the 1930s [6], organism. whose genome was sequenced in 2004 [7]. Different biomolecules isolated from Haloarcula marismortui (lipids, Keywords—Haloarcula marismortui, Halobacterium carotenoids, bacterioruberins, rhodopsins, polymers, halocins marismortui, Halobacterium of the Dead Sea, ATCC and proteins) reveals it as an extremely important research T 43049 , Halophilic archaea. topic. INTRODUCTION I. TIME-LAPSE In the 1930s, Elazari-Volcani isolated a novel strain of the Archaea is a domain which comprises numerous genus Halobacterium described as “Halobacterium microorganisms having the ability to survive extreme marismortui” from the Dead Sea. However, this strain was conditions for life, called “extremophiles” [1]. Archaea, more never deposited in a culture collection and is believed to be than other organisms, are exposed to a variety of stressors in lost. During the 1960s, a new Halobacterium strain was their natural habitats: physical (high and low temperatures, and radiation), mechanical (high and low pressures and isolated from the Dead Sea by Ginzburg et al [8]. This strain shearing) and chemical (sudden shift from anaerobiosis to was originally referred as “Halobacterium of the Dead Sea”. aerobiosis or vice versa; high and low pH, high osmolarity In 1975, the International Code of Nomenclature of Bacteria and salinity) [2]. The halophiles can be loosely classified as proposed the name Haloarcula marismortui (Volcani) sp. slightly, moderately or extremely halophilic, depending on nov., nom. rev [6]. Between 1986 and 1988, the isolated their requirement on NaCl to live. In particular, extremely strain by M. Ginzburg was deposited in the American Type IJERTV3IS061260 www.ijert.org 1257 International Journal of Engineering Research & Technology (IJERT) ISSN: 2278-0181 Vol. 3 Issue 6, June - 2014 Culture Collection by L. I. Hochstein (ATCC 43049)[9]. In Furthermore, optimal culture conditions (temperature, pH, 1990, Oren et al compiled valuable physiological information salt concentration and composition media) for H. marismortui about Haloarcula marismortui. are showed in Table II. The optimal incubation temperature, pH and salt concentration ranges reported for H. marismortui II. GENOME cultures are 40-50°C, as the family Halobacteriaceae that need for optimal growth slightly elevated temperatures (38– In 2004, Baliga et al reported the complete sequence of 45°C) [3, 15] 7.0-7.4, and 3.4-5.1 M, respectively. This last the 4.27 Mb genome of Haloarcula marismortui, which is feature denotes the marked halophilic character of H. one of the largest known genomes from the archaea domain, marismortui. Differences in biomass concentration values of only surpassed by Methanosarcina barkeri (4.86 Mb) and H. marismortui reported by several research groups, could be Methanosarcina acetivorans (5.75 Mb) [10]. Haloarcula explained by the different tested culture conditions (aeration, marismortui is the second sequenced genome from halophilic light, agitation, temperature, pH, type and concentration of archaea. The genome is organized into nine circular replicons nutrients in the culture media) [16]. of varying G·C compositions with an average of 57% [7]. This G·C composition is similar to other halophilic archaea TABLE II. OPTIMAL CULTURE CONDITIONS FOR such as Haloferax volcanii (average of 55%) [11] and HALOARCULA MARISMORTUI. Halobacterium sp. NRC-1 (average of 58%) [12]. Temperature Aeratio Furthermore, 1,310 proteins identified from mid-log and late- NaCl (M) pH Ref log phase of Haloarcula marismortui cultures were analyzed (°C) n (vvm) in metabolic and cellular process networks after being 40-50 3.4-3.9 7.0 * [14] exposed to environmental stresses by integrative genomics 42.5 3.6 7.4 1 [18] and proteomics analyzes [13]. Culture media composition (g/L) III. PHYSIOLOGICAL STUDIES NaCl, 234; MgS04.7H20, 53.2; CaCl2 2H20, 0.1; [8] MnCl2, 0.02mg; KCl, 0.51; K3(PO4), 0.31; yeast o Physiological studies of H. marismortui have been extract, 10. OD400-465nm of 0.2 (at 37 C). performed and the results are summarized in Table I. H. NaCl, 206; MgS04.7H20, 36; KCI, 0.373; [14] marismortui catabolizes sugars by glycolysis and Entner- CaCI2.2H20, 0.5; MnCI2, 0.013 mg; yeast extract, Doudoroff pathways and some amino acids. H. marismortui 5.0. Doubling time of 4.5-5 h (at 40°C). is a facultative aerobe, requiring growth factors; NaCl, 250; MgSO47H2O, 20; trisodium citrate, 3; [18] chemoorganotroph, oxidase and catalase positive. It can grow KCl, 2; tryptone, 5; yeast extract, 3; 0.1 ml of o anaerobically with nitrate as an electron acceptor. It utilizes a oligoelements. OD 600 nm of 3.3 (at 37 C). range of organic compounds as sole carbon and energy NaCl, 125; MgCl2.6H2O, 160; K2SO4, 5; [20] sources (glucose, fructose, sucrose, glycerol, acetate, CaCl2.2H2O, 0.1; yeast extract, 1; casamino-acids, 1; succinate and malate). It produces organic acids fromIJERTIJERT starch, 2. Cultures were performed at 160 rpm and o glucose, fructose, ribose, xylose, maltose, sucrose, mannitol, 37 C, during five days. sorbitol and glycerol. It hydrolyzes starch very slowly and * Cultures were made using Erlenmeyer flasks does not produce indole. It is susceptible to bacitracin and novobiocin [6, 14]. IV. BIOTECHNOLOGICAL APPLICATIONS OF HALOARCULA MARISMORTUI TABLE I. PHYSIOLOGICAL CHARACTERISTICS OF HALOARCULA MARISMORTUI. A few studies have been performed on the use of halophilic microorganisms for industrial waste treatment. Morphology Flat disk-shape [14] Haloarcula sp. IRU1 produced polyhydroxybutirate (PHB) Pleomorphic, flat and disk-shape [6] Mode of motility Rarely observed, motile cells rotate around [14] using petrochemical wastes and a saline culture medium [21]. their axis H. marismortui MTCC 1596 produced PHB, using vinasses Nonmotility (but rarely cells are observed [6] (ethanol industry liquid waste) and a saline culture medium. to rotate around their axis) In that study, H. marismortui cells accumulated 23 and 30% Motility (helicity flagellar filaments) [17] of the PHB (cell dry weight), using 10% of raw or pre-treated Size 1-2 and 2-3µm [6] (with activated carbon) vinasses, respectively [22]. One Intracellular ions Na+ 1.2-3M, K+ 3.77-5.5M, Cl- 2.3-4.2M [8] unconventional application using H. marismortui IR is to Sensitivity to Bacitracin and novobiocin [7, cure salted anchovies. The fermented anchovies exhibited antibiotics 17] low loads of Staphylococci, Enterobacteriaceae and lactic Starch digestion Positive [14] acid bacteria. They also had a reduced content of histamine Slow [6] and an improved organoleptic acceptance [23]. Remarkably, Utilization of Glucose, fructose, ribose, alfa- [7, industrial residual waters from the manufacture of cheese, carbon sources methylglucoside, maltose, sucrose, 17] corn flour, fish and
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