RJCES ©Academy for Environment and Life Sciences, INDIA

Research Journal of Chemical and Environmental Sciences Res J. Chem. Environ. Sci. Vol 2 [3] June 2014: 05-16 Online ISSN 2321-1040 CODEN: RJCEA2 [USA] RJCES ©Academy for Environment and Life Sciences, INDIA

REVIEW ARTICLE

Actinomycetes: Potential Bioresource for Human Welfare: A Review

1Roshan Kumar*, 2Koushik Biswas, 3Vikas Soalnki, 4Pankaj Kumar, 5Avijit Tarafdar 1School of Biotechnology, Chemical and Biomedical Engineering (SBCBE), VIT University, Vellore, (India) 2Department of Agricultural Biotechnology, CSK Himachal Pradesh Agricultural University, Palampur, Himachal Pradesh (India) 3Department of Biotechnology, Beehive College of Advance Studies, Selaqui, Dehradun (India) 4Department of Agricultural Biotechnology, CSK Himachal Pradesh Agricultural University, Palampur, HP (India) 5Cytogenetics and Tissue Culture Unit, Department of Botany,University of Kalyani, Nadia, West Bengal (India) Corresponding Author- [email protected]

ABSTRACT Microbial natural products are the origin of most of the antibiotics in the market today. There is an alarming scarcity of new antibiotics currently under development in the pharmaceutical industry. Still, microbial natural products remain the most promising source of novel antibiotics, although new approaches are required to improve the efficiency of the discovery process. Actinomycetes which are the prolific producers of antibi-otics and important suppliers to the pharmaceutical and other industry they are well known for their ability to produce secondary metabolites many of which are active against pathogenic microorganisms. It is only more recently that actinomycetes have become recognized as a source of novel antibiotics and anticancer agents with unusual structures and properties. They are a promising source of wide range of important enzymes, some of which are produced on an industrial scale, but many other remained to be harnessed. The application of enzymes in diverse biotechnological industries indicates a positive trend which needs to be satisfied with the discovery of novel enzymes and metabolites. Since very few enzymes have been potentially utilize data the industrial level; there is a huge scope for the development of robust and low cost enzymes. Actinomycetes are a reservoir of important enzymes and metabolites due to their versatile genetic repertory. They perform microbial transformations of organic compounds, a field of great commercial value. Members of many genera of actinomycetes have potential for use in the bioconversion of underutilized agricultural and urban wastes into high-value chemical products. Keywords- Actinomycetes, bioactive compound, Antibiotics, Enzymes.

INTRODUCTION Actinomycetes are a wealthy source for the synthesis of medically and technically useful natural products. From the ancient times actinomycetes is mostly related to its use as an antibiotic. Its use as an anti fungal agent in the past is mostly responsible for its popularity with antibiotic research today. From 1914 to 1939, Selman A. Waksman had been systematically screening soil bacteria and fungi in an attempt to find an antibiotic for Tuberculosis. University of California (1939) discovered the effect of certain fungi, especially antinomycetes, on bacterial growth. Actinomycetes slowed bacterial growth because of the antibiotics they produce.Actinomycetes comprise a substantially larger group having wider range of applications in food and pharma. Actinomycetes are Gram-positive bacteria with a high G+C (>55%) content. Among others, representative genera include Micrococcus, Mycobacterium, Nocardia, Propionibacterium, and Streptomyces. Many actinomycetes, such as Streptomyces, grow as branching filaments and live in soil, as fungi do. Because of this resemblance, actinomycetes were originally classified as fungi. This was reflected on their name, where "mycetes" comes from the Greek for "mushroom, fungus". The actinomycetes represent a ubiquitous group of microbes that are widely distributed in natural ecosystems around the world and are particularly significant for their role in the recycling of organic matter. [1, 2] reported a bimodal distribution of actinomycetes in near shore tropical marine environments. The habitat of actinomycetes corresponds to its behavioral characteristics. Actinomycetes is a saprophyte, another word for a decomposing organism, which means it grows best in moist moderate to tropical atmospheres. This bacterium is also a heterotroph, meaning it draws its

RJCES Vol 2 [3] June 2014 5 | P a g e © 2014 AELS, INDIA Kumar et al energy from surrounding sources such as dead and decomposing animal matter. These factors determine the habitat of actinomycetes. As a decomposer, actinomycetes is commonly found in compost piles and forest floor litter, and forms symbiotic relationships with red alders, a type of tree that forms anaerobic nodules in which actinomycetes fixes nitrogen for the tree. Actinomycetes bacteria are found in human and cattle bodies in the mouth, throat, and intestinal tract. Occurrence Actinomycetes occur in the soil in the spore stage as well as in the mycelial stage. As a result of comparative examination of the relative abundance of actinomycetes in the form of substrate growth and spores in soil, using the microscopic and plate methods. The mycelium developed most abundantly at 28 oC to 37 oC; at lower and higher temperatures growth was slower but eventually reached the same density. At higher temperatures, the mycelium underwent increasing fragmentation, giving rise to abundant formation of spores. Sporulation is also favored by a dry atmosphere. Enrichment of soil with bacteria leads to extensive actinomycete development; their excessive growth is due largely to the introduction of fresh supply of available nutrients in the form of bacterial cells. Addition of organic matter has in general a marked stimulating effect upon the development of Actinomycetes. When soils rich in organic matter such as peat bogs, are drained and aerated, actinomycetes are able to make extensive growth. Actinomycetes, including Streptomycetes and certain nocardiae, occur abundantly in and around the root systems of higher plants. (Table.1) contain list of actinomycetes strain present in particular plant and their function are given in. Some forms produce yellowish, orange, or black pigments in organic media. Some are spiral producing, others forming straight aerial mycelium. Certain horizons of different soil types were found to contain characteristic communities of Streptomycetes.Freshwater lakes, rivers, and sewage contain an abundance of actinomycetes, including thermophilic forms growing well at 60 oC. Diversity of marine actinobacteria An intriguing picture of the diversity of marine actinobacteria is beginning to emerge. Once largely considered to originate from dormant spores that washed in from land [3], it is now clear that specific populations of marine adapted actinobacteria not only exist but add significant new diversity within a broad range of actinobacterial taxa [ 4,5]. The first report on the marine actinobacteria was made by [6], when he observed and documented those in the salt mud. In 1969, Weyland [7] carried out an extensive survey on the distribution on marine actinobacteria in the sediments of North Sea and Atlantic Ocean and suggested that the marine actinobacteria are the best sources for isolation of unique bioactive compounds compared to terrestrial ones. After this, a number of researchers around the world have concentrated to isolate and identify the actinobacteria from the different marine habitats. Ecology of antibiotic producing actinomycetes Microbial diversity is a substantial leading edge and prospective goldmine for biotechnology industry because it offers countless of secondary metabolites to probe for enzymes, antibiotics, antioxidant, cytotoxic and so many other useful substances [8-10]. The actinomycetes occur in vast diversity of habitat either natural or artificial, growing on different kinds of substrate. The diversity of actinomycetes is of exceptional impact in several areas of pharmaceutical, medicine and agriculture, particularly, in antibiotic production [11]. Actinomycetes are ubiquitous and have been isolated from various locations, in the soil, fresh water, marine, hot spring, mining sites, and also in extreme environments.

Table 1. Examples of rizosphere some actinomycetes and their functions to plants. Rhizospheric Function Plant species References actinomycetes Micromonospora endolithica Phosphate solubilization to Bean (Phaseolus vulgaris [12] promote plant growth L.) Streptomyces griseus Protection against damping Wheat (Triticum spp.) [13] off disease caused by Pythium ultimum Frankia species Biological fixation of Actinorhizal plant [14] nitrogen (Casuarina equisetifolia) Norcardia levis Biological control of Sorghum (Sorghum bicolor) [15] Fusarium oxysporum wilt disease Streptomyces species Act as biocontrol against Tomato (Solanum [16] Rhizoctonia solani lycopersicum) Streptomyces species Bioremediation of Maize (Zea mays) [17] contaminated soil

Role of actinomycetes in the marine environment The marine environment is characterized by the hostile parameters such as high pressure, salinity; low temperature, absence of light, etc. and the marine actinobacteria have adapted themselves to survive in this environment. They require Na+ for growth because it is essential to maintain the osmotic environment for protection of cellular integrity. Oligotrophy is also one more adaptation because of the smaller amount of available nutrients. However, actinobacterial action promotes organic degradation, decomposition and mineralization processes in the sediments and in the overlying water column and releases of the dissolved organic and inorganic substances. The mineralization of organic matter, which is derived from the primary producers, results in its being recycled, so that these substances are again available for the primary producers. Distribution of actinobacteria depends on changes in water temperature, salinity and other physico-chemical parameters. Actinobacteria also serve as important source of food for a variety of marine organisms. Thus, actinobacteria not only maintain the pristine nature of the environment, but also serve as biological mediators through their involvement in biogeochemical processes. Breakdown of organic matter Marine actinobacteria play a decisive role in the cycling of matter in water, as they are able to breakdown all natural organic compounds into the compounds from which they have originated. Decomposition of protein is done by proteolytic actinobacteria e.g. Streptomyces galbus. Cellulose is decomposed by cellulotic actinobacteria e.g. Streptomyces actuosus. Chitin, which is synthesized by several marine organisms as extra-cellular material algae, cell walls of some chlorophytes [18] exoskeleton, including molts from copepods and other marine invertebrates [19] is a structural polysaccharide. However, it is not degraded easily [20] and there is a report on chitin preservation in fossils [21]. However, this biopolymer is degraded by chitinolytic or chitinoclastic actinobacteria by their exoenzyme chitinase. Starch is also decomposed by numerous actinobacteria [22-23]. Biodegradation of pollutants Marine actinobacteria have a marked ability to degrade complex molecules such as petroleum. Rhodococcus sp. and Mycobacterium sp. are some of the marine actinobacteria found to degrade a range of hydrocarbons [24]. Large oil spills are one of the most dramatic and terrible environmental disasters. Some of the oil is degraded by marine actinobacteria, which degrades the hydrocarbon in oil by using as a carbon source. Scientists have been experimenting with the use of oil degrading marine actinobacteria in oil spill clean ups. Marine actinobacteria, which naturally degrade oil, grow much more slowly than other bacterial strains.

Actinomycetes as a potential revenue resource Antibiotics Actinobacteria are unsurpassed in their ability to produce many antibiotics that have pharmaceutically useful properties. In 1940, Selman Waksman discovered that the soil bacteria contain actinomycin, which granted him the Nobel Prize. Since then, hundreds of naturally occurring antibiotics have been discovered in the terrestrial microorganisms. Marine actinobacteria also constitute an important and potential source of novel antibiotics [25]. Since environmental conditions of the sea are extremely different from the terrestrial conditions, they produce different types of antibiotics. Several antibiotics have been isolated from marine actinobacteria by many researchers which are summarized in (Table 2). The isolated antibiotics are entirely new and unique when compared to those from the terrestrial ones [26]. Anti-cancer compounds Marine actinobacteria are the diversified groups which are capable of producing different types to anticancer compounds. [27-36] isolated different kinds of cytotoxic compounds form the marine actinobacteria (Table 2). The isolated compounds showed significant activity against different cancer cell line.

Table 2. List of the bioactive compounds derived from the marine actinobacteria. S. Bioactive compound Species Activity Reference no. 1. Abyssomicin Verrucosispora sp. Antibacterial and antitumor [37] 2. Actinoflavoside Streptomyces sp. Antifungal [38] 3. Actinofuranones A-B Streptomyces sp. Antibacterial and antitumor [39] 4. Altemicidin S. sioyaensis Antibacterial and antitumor [40] 5. Analogs-metacycloprodigiosin Saccharopolyspora sp. Anticancer [41] 6. Analogs-undecylprodigiosin Saccharopolyspora sp. Anticancer [41] 7. Aplasmomycins A-C Streptomyces griseus Antibacterial [42, 43] 8. Benzanthraaquinone Chainia purpurogena Antibacterial and antitumor [44]

9. Bioxalomycins Streptomyces sp. Antibacterial [45] 10. Butenolides Streptomyces sp. Antitumor [46] 11. Chalcomycin-B Streptomyces sp. Antibacterial [47] 12. Chandrananimycin Actinomadura sp. Antibacterial & antifungal [48] 13. Chinikomycins Streptomyces sp. Anticancer [49] 14. Chromomycin-A3 Streptomyces sp. Antibacterial [50] 15. Cyanosporasides A-B Salinispora sp. Antitumor [51] 16. Cyanthiwigins A-E Streptomyces spheroids Antibavterial & antifungal [51] 17. Cyclic tetrapeptide Nocardiopsis sp. Antibacterial [53] 18. Debromomarinon Streptomyces sp. Antibacterial [54] 19. Delta-indomycinone Streptomyces sp. Antibacterial [55] 20. Diazepinomicin Micromonospora sp. Antifungal [56] 21. Diphosphatidylglycerol Micromonospora sp. Antitumor [57] 22. Glucosylmannosyl-glycerolipid Microbacterium sp. Antitumor [58] 23. Glycoglycerolipids Microbacterium sp. Antitumor [59] 24. Halichoblelide Streptomyces sp. Antitumor [60] 25. Himalomycin A-B Streptomyces sp. Antitumor [61] 26. Holyrines A-B Streptomyces sp. Antibacterial & antitumer [62] 27. Istaycines A-B S. tenjimariensis Antibacterial [63] 28. Lajollamycin S. nodosus Antitumor [33] 29. Loesaponarin II Streptomyces sp. Antitumor [64] 30. Lorneamide A-B Streptomyces sp. Antifungal [65] 31. Kahakamides A-B Nocardiopsis dassonvillei Antibacterial [66] 32. Komodoquinone Streptomyces sp. Antibacterial and antifungal [67] 33. Macrolide Micromonospora Antitumor [68] 34. Marinomycins A-D Marinispora sp. Antitumor [69] 35. Marinone Streptomyces sp. Antibacterial [54] 36. Mechercharmycin Thermoactinomyces sp. Antitumor [70] 37. Menaquinone Streptomyces sp. Antibacterial [71] 38. Neomarinone Streptomyces sp. Antitumor [72] 39. Octalactins A-B Strptomyces sp. Antibacterial and antitumor [73] 40. Oxolonomycin Streptomyces sp. Antibacterial [74] 41. Parimycin Streptomyces sp. Antitumor [75] 42. Phencomycin Streptomyces sp. Antibacterial [76] 43. Pyrrolosesquiterpenes Streptomyces sp. Antibacterial [77] 44. Resistoflavine S. chibaensis Antibacterial [78] 45. Resistomycin S. corchorusii Antibacterial [79] 46. Tetracenomycin S. corchorusii Antibacterial [79] 47. Tetrodotoxin Streptomyces sp. Antibacterial [80] 48. Thiocoraline Micromonospora sp. Antimicrobialand antitumor [81]

49. Thiocoraline Micromonospora sp. Antitumor [82] 50. Trioxacarcins A Streptomyces sp. antibacterial and antitumor [83] ,antimaleria 51. Salinamides A- B Streptomyces sp. Antibacterial [84] 52. Salinosporamide A Salinispora tropica Cytotoxic proteasome inhibitor [85] 53. Staurosporine Micromonospora sp. Antitumor [86] 54. Streptokordin Streptomyces sp. Antitumor [87] 55. Sporolides Salinispora tropica Antitumor [88] 56. Wailupemycins A-C Streptomyces sp. Antbacterial [89] 57. L-methionine S. maritimus Antbacterial [90] 58. N-acetyl-gamma-hydroxyvaline lactone Streptomyces sp. Antbacterial [91] 59. N-carboxamido-staurosporine Streptomyces sp. Antitumor [92] 60. 1,6-dihydroxy-8- Streptomyces sp. Antibacterial [63] hydroxymethylanthraquinone 61. 1-Hydroxy-1-norresistomycin Streptomyces chibaensis Antibacterial and antitumor [93] 62. 3-epi-Sdeoxyenterocin Streptomyces sp. Antibacterial [89] 63. 10-methyl-6-dodecanolide Streptomyces sp. Antitumor [94] 64. Essramycin Streptomyces sp. anti-inﬂammatory [95] 65. Lynamicins Marinispora sp. anti-inﬂammatory [96] 66. Marinopyrroles Streptomyces sp. Cytotoxic [97] 67. Caboxamycin Streptomyces sp. Cytotoxic [98] 68. Tirandamycin Streptomyces sp Antifungal; anticance [99] 69. TP-1161 Nocardiopsis sp. Antifungal; anticancer [100] 70. Tirandamycins Streptomyces sp. Antifungal; anticancer [99] 71. 1,4-Dihydroxy-2-(3-hydroxybutyl)-9,10- Streptomyces sp. Antifungal; anticancer [101] anthraquinone 72. N-(2-hydroxyphenyl)-2-phenazinamine Nocardia dassonvillei. anticancer [110] (NHP) 73. Aureolic acid Streptomyces sp. Antibacterial [102]

74. Elaiomycins B and C Streptomyces sp. Phycotoxicity [103] 75. Salinipyrones Salinispora pacifica Phycotoxicity [104] 76. Pacificanones Salinispora pacifica Phycotoxicity [104] 77. Mansouramycin C Streptomyces sp. Phycotoxicity [105] 78. Usabamycins Streptomyces sp. Phycotoxicity [106] 79. Pyridinium Amycolatopsis alba. Antimicrobial [107] 80. ML-449 (macrolactam Streptomyces sp. Antimicrobial [108] 81. Albidopyrone Streptomyces sp Cytotoxic (inhibitor of [109] protein–tyrosine phosphatise) 82. Proximicins Verrucosispora sp. Cytostatic activity [110] 83. Cyclomarins Streptomyces sp. Anti-inflammatory activity [111] 84. Dermacozines A-G Dermacoccus Antitumor; Antiprotozoal [112] 85. Lipocarbazoles Tsukamurella Antiprotozoal [110] pseudospumae 86. 2-Allyloxyphenol Streptomyces sp. Antimicrobial [112] 87. Streptopyrrolidine Streptomyces sp. Anti-angiogenesis activity [114] 88. Cyclo-(l-Pro-l-Met) Nocardiopsis sp. Anti-angiogenesis activity [115]

Actinomycetes as a excellent sources of commercial enzymes The commercial value for enzymes has increased substantially with the uses including confectionary, detergents, industry etc. Marine actinobacteria have a diverse range of enzyme activity and are capable of catalyzing various biochemical reactions with novel enzymes. The applications of few commerciallysignificant enzymes are enlisted in (Table 3). Amylase, protease and cellulose enzymes are important and production of these enzymes and formation of fermentative products by Streptomyces spp. are important for their commercial application in textile, paper, laundry, confectionary and sugar industries [116] L- glutaminase, L-asparaginase and -galactosidase also play an important role in biocycling of carbon and nitrogen in natural water and sediments. L-glutaminase and L-asparaginase have shown antitumor activities and were optimized from the marine actinobacteria [117,118]. Golbally the enzyme market has also increased, India is also taking active effort. The global industrial enzyme market has evolved continounsly due to numerous mergers and acquisitions. Different sector like food, beverage enzyme are taking active part in these decades which are shown in Figure 1. There are also increase in patent number in the decades in linear scale which given in figure 2.

Figure 1: Global enzyme industry market in the years 2011 and 2016 [119]

Figure 2: Growth in number of patents issued for important industrial enzymes over past few decades. [119]

Table 3. Commercially relevant enzymes produced by actinomycetes and their application Enzyme Actinomycetes Strains Use Industrial of References application Protease S. galbus Detergents Detergents [120] Cheese making Food Clarication- low calorie beer Brewing Dehiding Leather Treatment of blood clot Medicine Cellulase S. actuosus Removal of stains, Denim nishing, Denim nishing, soening [121] Soening of Detergent Deinking, modication of of bers Paper and pulp Cotton Lipase S. griseochromogenes Removal of stains Detergent [122] Stability of dough and conditioning Baking Cheese avoring Dairy Deinking, cleaning Textile Xylanase S. rameus Conditioning of dough Baking [123] Digestibility Animal feed Bleach boosting Paper and pulp Pectinase S. fradzae Clarication, mashing Beverage [124] S nztrosporeur Scouring Textile Amylase S. aureofasciculus, Soness of bread soness Detergen [23] S. galilaeus Removal of stains volume Baking Deinking, drainage improvement Paper and pulp Production of glucose and fructose syrups Starch industry Removal of starch from woven fabrics Textile Glucos oxidase Streptomyces sp. Strengthening of dough Baking [125] Lipoxygenase Streptomyces sp. Bread whitening Baking [126]

Phytase Phytate S. ambofaciens digestibility Animal feed [127] S. lienomycini. Peroxidase Thermomonospora fusca Removal of excess dye Textile [128] S. viridosporus β-galactosidase Streptomyces sp. Enzymatic hydrolysis of lactose either from Dairy [129] milk/whey or pure lactose L-asparaginase S. aureofasciculus, Reduce the formation of acrylamide, a Food Industry [130, 131] S. canus, carcinogen found in starchy food products S. chattanoogenesis, S. hawaiiensis, S. olivoviridid, S. orientalis, S. plicatus L-glutaminase S. rimosus, S. galbus Flavor enhancing agent in food Food Industry [132, 133]

Keratinase Doretomycetes animal feed poultry Industry [134] microsporus

Petinase Thermomonospora ﬂisca retting and degumming of fiber crops textile industry [134, 135] S. viridochromogenes

Enzyme inhibitors Enzyme inhibitors have received increasing attention as useful tools, not only for the study of enzyme structures and reaction mechanisms but also for potential utilization in pharmacology [136]. Marine actinobacteria are the potential source for production of enzyme inhibitors [137, 139] reported different types of enzyme inhibitors viz. -glucosidase, N-acetyl--D-glucosaminidase, pyroglutamyl peptidase and - amylase inhibitors from marine actinobacteria. Single cell protein feed Marine actinobacteria can be used as the substitute for fishmeal. Actinobacteria can produce some secondary metabolities which may enhance the growth of juvenile fish, shrimp and prawn. Some of the secondary metabolites are organometalic compounds such as ferrioxamines, magnesidin with bleomycin and beron containing compounds such as boromycin and aplasmomycin [140] and usual amino acids such as alanosine, amino dichlobutyric acid, azaleucine, 4-oxalysine etc. [141]. Juveniles of prawns and shrimps fed on actinobacteria incorporated feeds showed higher growth percentage, more food conversion efficiency and higher protein content [142]. Hence, among the unconventional protein sources, single cell protein (SCP) of microbial origin appears to be a promising substitute for fishmeal,

RJCES Vol 2 [3] June 2014 10 | P a g e © 2014 AELS, INDIA Kumar et al which can replace up to 25-50% fishmeal in aquaculture operations. Marine actinobacteria are the potential microorganisms which have both ecological and biotechnological importance. Till now, 83 species of actinobacteria belonging to 28 genera have been recorded from the marine environment and most of these genera are new to science. If in-depth studies are carried out, the diversity of the marine actinobacteria will be increased. Nearly 64 novel bioactive compounds have been isolated from the marine actinobacteria which show higher antimicrobial and anticancer activities. Apart from this, different commercial enzymes and enzyme inhibitors have been also reported from the marine actinobacteria. Further, some of these compounds after clinical trials have not shown any toxicity and side effect. But it is a pity that none of the compounds have been commercialized.

CONCLUSION Actinomycetes play a significant role in the production of antimicrobial agents and other industrial important product like enzymes, drugs and other source of natural pigment and compound. It is essential to have a good knowledge about their taxonomy and ecology for maximum exploration, since they are of great use for economic and industrial development. In soil ecology, they are also active in bioremediation, biofertilizer, biocontrol and as plant growth promoters, making them indispensable in agricultural practice. Actinomycetes are the potential microorganisms which have both ecological and biotechnological importance. Till now, several species of actinomycetes have been discovered from the marine environment and most of these genera are new to science. Several type of novel bioactive compounds have been isolated from the actinobacteria which show higher antimicrobial and anticancer activity. Since the need for antimicrobials is going up day by day due to emergence of new pathogens or due to drug resistance, so efforts are to be taken to discover newer and potent antimicrobials to combat these emerging diseases. Actinomycetes being the efficient producers could be exploited for the production of these drugs and we can diversify the range of antimicrobials only if we explore more and more un discovered or unexploited species.Although, a great work has been carried out on actinomycetes, more comprehensive studies are still needed in the area of taxonomy and ecology. This will help to predict the productivity of the members of this order and possible exploitation.

ACKNOWLEDGEMENTS The authors are thankful to Chancellor and authorities of VIT University, Vellore, Tamil Nadu for support during compilation of this review article.

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Cite this article Roshan K., Koushik B., Vikas S., Pankaj K., Avijit T. Actinomycetes: Potential Bioresource for Human Welfare: A Review. Res. J. Chem. Env. Sci. Vol 2 [3] June 2014. 05-16