List of antibiotics produced from Actinobacteria
Actinobacteria, which share the characteristics of both bacteria and fungi, are widely distributed in both terrestrial and aquatic ecosystems, mainly in soil, where they play an essential role in recycling refractory biomaterials by decomposing complex mixtures of polymers in dead plants and animals and fungal materials. They are considered as the biotechnologically valuable bacteria that are exploited for its secondary metabolite production. Approximately, 10,000 bioactive metabolites are produced by Actinobacteria, which is 45% of all bioactive microbial metabolites discovered. Especially Streptomyces species produce industrially important microorganisms as they are a rich source of several useful bioactive natural products with potential applications. Though it has various applications, some Actinobacteria have its own negative effect against plants, animals, and humans. On this context, this chapter summarizes the general characteristics of Actinobacteria, its habitat, systematic classification, various biotechnological applications, and negative impact on plants and animals.
- Secondary metabolites
Actinobacteria are a group of Gram-positive bacteria with high guanine and cytosine content in their DNA, which can be terrestrial or aquatic. Though they are unicellular like bacteria, they do not have distinct cell wall, but they produce a mycelium that is nonseptate and more slender. Actinobacteria include some of the most common soil, freshwater, and marine type, playing an important role in decomposition of organic materials, such as cellulose and chitin, thereby playing a vital part in organic matter turnover and carbon cycle, replenishing the supply of nutrients in the soil, and is an important part of humus formation. Actinobacterial colonies show powdery consistency and stick firmly to agar surface, producing hyphae and conidia/sporangia-like fungi in culture media.
Actinobacteria produce a variety of secondary metabolites with high pharmacological and commercial interest. With the discovery of actinomycin, a number of antibiotics have been discovered from Actinobacteria, especially from the genus
2. Habitat of Actinobacteria
2.1. Terrestrial environment
Soil remains the most important habitat for Actinobacteria with streptomycetes existing as a major component of its population. According to numerous reports,
2.2. Aquatic environment
Actinobacteria are widely distributed in aquatic habitats, which may sometimes be washed in from surrounding terrestrial habitats. It is vitally important that the numbers and kinds of Actinobacteria are interpreted in the light of information on organisms, such as
Cross  in his study evidenced that Actinobacteria can readily be isolated from freshwater sites. Some of the major type of Actinobacteria dwelling in freshwater include
When comparing the Actinobacterial diversity in terrestrial environment, the greatest biodiversity lies in the oceans. The marine environment is an untapped source of novel Actinobacteria diversity and thus of new metabolites. Marine Actinobacteria dwelling in extremely different environment produce different types of bioactive compounds compared with terrestrial ones. Marine Actinobacteria had to adapt from extremely high pressure and anaerobic conditions at temperatures just below 0- 8 °C on the deep sea floor to high acidic conditions at temperatures of over 8- 100°C near hydrothermal vents at the mid-ocean ridges.
3. General characteristics of Actinobacteria
Actinobacteria comprises a group of branching unicellular microorganisms, most of which are aerobic-forming mycelium known as substrate and aerial. They reproduce by binary fission or by producing spores or conidia, and sporulation of Actinobacteria is through fragmentation and segmentation or conidia formation. The morphological appearance of Actinobacteria (Figure 1) is compact, often leathery, giving a conical appearance with a dry surface on culture media and are frequently covered with aerial mycelium.
3.1. Aerial mycelium
The aerial mycelium is usually thicker than the substrate mycelium (Figure 2a). The aerial mycelium shows sufficient differentiation that a miscellaneous assortment of isolates can be segregated into a number of groups having similar morphological characteristics under fixed condition. This is designated as one of the most important criteria for the classification of the genus
3.2. Substrate mycelium
The substrate mycelium of Actinobacteria varies in size, shape, and thickness (Figure 2b). Its color ranges from white or virtually colorless to yellow, brown, red, pink, orange, green, or black.
3.3. Morphological appearance
Morphology has been an important characteristic to identify Actinobacteria isolates, which was used in the first descriptions of
4. Systematics of Actinobacteria
In the Bergey’s Manual of Determinative Bacteriology, Actinobacteria are included in several sections of volume four. All Actinobacteria are included under the order Actinomycetales. The order Actinomycetales is divided into four families—Streptomycetaceae, Actinomycetaceae, Actinoplanaceae, and Mycobacteriaceae . The “Bergey’s Manual of Systematic Bacteriology—2nd edition” for Actinobacteria classification has five volumes, which contain internationally recognized names and descriptions of bacterial species. Classification of Actinobacteria has been rearranged as follows:
|The Archaea and the Deeply Branching and Phototrophic Bacteria||
|The Bacteroidetes, Spirochaetes, Tenericutes (Mollicutes), Acidobacteria, Fibrobacteres, Fusobacteria||
In Volume 5, the phylum Actinobacteria is divided into six classes, namely Actinobacteria, Acidimicrobiia, Coriobacteriia, Nitriliruptoria, Rubrobacteria, and Thermoleophilia. The class Actinobacteria is further divided into 16 orders that are Actinomycetales, Actinopolysporales, Bifidobacteriales, Catenulisporales, Corynebacteriales, Frankiales, Glycomycetales, Jiangellales, Kineosporiales, Micrococcales, Micromonosporales, Propionibacteriales, Pseudonocardiales, Streptomycetales, Streptosporangiales, and Incertae sedis. In the order of abundance in soils, the common genera of Actinobacteria are
5. Types of Actinobacteria
5.1. Thermophilic Actinobacteria
Number of studies has been carried out by the researchers to confirm the existence of extremophilic and extreme tolerant soil Actinobacteria (acid tolerant and alkali tolerant, psychrotolerant and thermotolerant, and halotolerant and haloalkalitolerant or xerophilic). Mesophilic Actinobacteria can grow at an optimal temperature from 20°С to 42°С, among which thermotolerant species exist, which can survive at 50°С. Moderately thermophilic Actinobacteria have an optimum growth at 45°С–55°C , whereas strictly thermophilic Actinobacteria grow at 37°С–65°C with the optimum temperature at 55°С–60°C . Incubation temperatures of 28°С, 37°С, and 45°C are considered optimal for isolation of soil mesophilic, thermotolerant, and moderately thermophilic Actinobacteria.
5.2. Acidophilic Actinobacteria
Acidophilic Actinobacteria, which are common in terrestrial habitats such as acidic forest and mine drainage soil, grow in the pH range from about 3.5 to 6.5, with optimum rates at pH 4.5 to 5.5 [26, 27]. It has been shown that acidophilic Actinobacteria consistently form two distinct aggregate taxa (namely, the neutrotolerant acidophilic and strictly acidophilic cluster groups) based on numerical phenetic data; members of the two groups share common morphological and chemotaxonomic properties . Also some members of the strictly acidophilic group form a distinct taxon, such as the genus
5.3. Halophilic Actinobacteria
Halophilic Actinobacteria are categorized into different types based on their growth in media containing different concentrations of salt. Extreme halophiles grow best in media containing 2.5–5.2 M salt, whereas borderline extreme halophiles grow best in media containing 1.5–4.0 M salt, moderate halophiles grow best in media containing 0.5–2.5 M salt, and finally halotolerants that do not show an absolute requirement to salt for growth but grow well up to often very high salt concentrations and tolerate 100 g/l salt (equivalent to 1.7 M NaCl) at least. Seawater, saline soils, salt lakes, brines, and alkaline saline habitats are considered as the best habitats for isolating halophilic Actinobacteria. Generally, most of the halophilic Actinobacteria have been isolated from saline soils. Halophilic Actinobacteria isolated from marine environments are assigned to a few genera, including
5.4. Endophytic Actinobacteria
Endophytic Actinobacteria are defined as those that inhabit the internal part of plants, causing apparently no visible changes to their hosts. These Actinobacteria play specific roles, for instance, protecting the host plants against insects and diseases. Endophytic Actinobacteria constitute a large part of the rhizosphere, which are also found inside plants in which the extensively studied species are from the genus
5.5. Symbiotic Actinobacteria
About 15% of the world’s nitrogen is fixed naturally by the symbiotic relationships between various species of the
5.6. Endosymbiontic Actinobacteria
An endosymbiont is any organism that lives within the body or cells of another organism. Endosymbiosis process is sometimes obligate, that is, either the endosymbiont or the host cannot survive without the other. Members of the phylum Actinobacteria have been identified as abundant members of sponge-associated microbial communities.
5.7. Gut Actinobacteria
Though Actinobacteria are found in various diverse habitats, some are also known to form intimate associations with invertebrates and vertebrates. Symbiotic interactions are essential mainly for the survival and reproduction because they play a crucial role in nutrition, detoxification of certain compounds, growth performance, and protection against pathogenic bacteria. Many studies have shown that some symbiotic Actinobacterial species, that is probiotics, control bacterial diseases in livestock, poultry, and aquaculture. They also take part in host health by converting the feedstuffs into microbial biomass and fermentation end products that can be utilized by the animal host. Tan et al  isolated
6. Applications of Actinobacteria
Actinobacteria are well recognized for their production of primary and secondary metabolites that have important applications in various fields. They are also a promising source of wide range of important enzymes, which are produced on an industrial scale. A large fraction of antibiotics in the market is obtained from Actinobacteria. They produce enzyme inhibitors useful for cancer treatment and immunomodifiers that enhance immune response. They have the ability to degrade a wide range of hydrocarbons, pesticides, and aliphatic and aromatic compounds. They perform microbial transformations of organic compounds, a field of great commercial value. Members of many genera of Actinobacteria can be potentially used in the bioconversion of underutilized agricultural and urban wastes into high-value chemical products. Actinobacteria are also important in plant biotechnology as strains with antagonistic activity against plant pathogens are useful in biocontrol. Their metabolic potential offers a strong area for research. Here, we have a brief description about important applications of Actinobacteria (Figure 5).
Actinobacteria hold a significant role in producing variety of drugs that are extremely important to our health and nutrition. Recently, diseases due to multidrug-resistant pathogenic bacteria are sturdily increasing, and thus search for new antibiotics is effective against the multidrug-resistant pathogens. Natural products having novel structures have been observed to possess useful biological activities . Nature always remains the richest and the most versatile propitious source for new antibiotics, though there is considerable progress within the fields of chemical synthesis and engineered biosynthesis of antibacterial compounds. Toxic nature of certain antibiotics led to their limited usage although thousands of antibiotics have been discovered till date. To get through this problem, search of new antibiotics that are more effective and that do not have any toxic side effects is in progress. As already mentioned, one of the major healthcare problems is the antibiotic resistance. One approach to solve this problem is to search for new antibiotics with new mechanism of action. Figure 6 shows that a majority of antibiotics are derived from microorganisms, especially from the species Actinobacteria. Almost 80% of the world’s antibiotics are known to be derived from Actinobacteria, mostly from the genera
Antagonistic Actinobacteria produce a variety of antibiotics that vary in chemical nature, in antimicrobial action, in toxicity to animals, and in their chemotherapeutic potentialities. Some of the antibiotics that have been isolated so far from Actinobacteria are crude preparations, whereas others have been crystallized, and considerable information has been gained concerning their chemical nature, which includes lysozyme, actinomycin, micromonosporin, streptothricin, streptomycin, and mycetin. Some Actinobacteria produce more than one antibiotic substance (e.g.,
|2-Allyloxyphenol||Antimicrobial; food preservative; oral disinfectant||
|Arenicolides A–C||Mild cytotoxicity||
|Bafilomycin||ATPase inhibitor of microorganisms, plant and animal cells||
|Chloramphenicol||Antibacterial, inhibitor of protein biosynthesis||
|Lincomycin||Antibacterial, inhibitor of protein biosynthesis||
|Marinomycins A–D||Antimicrobial; anticancer||
|Mitomycin C||Antitumor, binds to double-stranded DNA||
|Pacificanones A & B||Antibacterial||
|Resistoflavin methyl ether||Antibacterial; antioxidative||
|Salinispyrone A & B||Mild cytotoxicity||
|Salinosporamide A||Anticancer; antimalarial||
|Salinosporamide B & C||Cytotoxicity||
|Valinomycin||Ionophor, toxic for prokaryotes and eukaryotes||
|Elaiomycins B and C||Antitumor||
|N-[2-hydroxyphenyl)-2-phenazinamine (NHP),||Anticancer; antifungus||
|Chromomycin B, A2, A3||Antitumor||
|1,4-dihydroxy-2-(3-hydroxybutyl)-9, 10-anthraquinone 9, 10-anthrac||Antibacterial||
A wide variety of biologically active enzymes are produced by both marine and terrestrial Actinobacteria (Figure 7; Table 2). They secrete amylases to the outside of the cells, which helps them to carry out extracellular digestion. This enzyme is of great significance in biotechnological applications such as food industry, fermentation, and textile to paper industries because of their ability to degrade starch . Another important aspect of Actinobacteria is the production of cellulases, which are a collection of hydrolytic enzymes that hydrolyze the glucosidic bonds of cellulose and related cello-digosaccharide derivatives. Lipase is produced from various Actinobacteria, bacteria, and fungi and is used in detergent industries, foodstuff, oleochemical, diagnostic settings, and also in industries of pharmaceutical fields . Many Actinobacteria have been isolated from various natural sources, as well as in plant tissues and rhizospheric soil. Biological functions of Actinobacteria mainly depend on sources from which the bacteria are isolated. Actinobacteria, particularly streptomycetes, are known to secrete multiple proteases in the culture medium . Similarly, Actinobacteria have been revealed to be an excellent resource for L-asparaginase, which is produced by a range of Actinobacteria, mainly those isolated from soils, such as
|Clarification- low calorie beer||Brewing|
|Treatment of blood clot||Medicine|
||Removal of stains||Detergent|
|Denim finishing, softening of cotton||Textile|
|Deinking, modification of fibers||Paper and pulp|
||Removal of stains||Detergent|
|Stability of dough and conditioning||Baking|
||Conditioning of dough||Baking|
|Bleach boosting||Paper and pulp|
||Removal of stains||Detergent|
|Softness of bread softness and volume||Baking|
|Deinking, drainage improvement||Paper and pulp|
|Production of glucose and fructose syrups||Starch industry|
|Removal of starch from woven fabrics||Textile|
||Strengthening of dough||Baking|
||Feather degradation||Animal feed|
||Phytate digestibility||Animal feed|
Another interesting application of the Actinobacteria is the use of their secondary metabolites as herbicides against unwanted herbs and weeds.
Probiotics are the live microbial adjunct that has a beneficial effect on the host by various means, such as modifying the host associated or ambient microbial community, by ensuring the improved use of the feed or enhancing its nutritional value, by enhancing the host response towards disease, or by improving the quality of its ambient environment. Despite several other important applications, marine Actinobacteria have been given its attention for their use as probiotics. The potential of Actinobacteria against shrimp pathogenic
6.4.1. Aggregative peptide pheromones
Aggregation is one of the most important criteria for the selection of a good probiotic candidate, which is the process of reversible accumulation of cells with one or more strains. For this aggregating process to take place, pheromone production is one of the main criteria that involves defense against predators, mate selection, and in overcoming host resistance by mass attack. In particular, sex pheromone peptides in culture supernatants have been shown to promote aggregation not only with the same species but also with related species [60–62]. Thus, the auto-aggregating ability of a probiotic is a prerequisite for colonization of the gastrointestinal tract, whereas coaggregation provides a close interaction with pathogenic bacteria. Though there are a number of studies in accordance with peptide pheromone–mediated signaling, it is lacking in the case of Actinobacteria, and thus a novel report on isolation and purification of diffusible aggregation promoting factor, that is, pheromones from potent Actinobacterial probiont
Biosurfactants are the microbially derived compounds that share hydrophilic and hydrophobic moieties that are surface active. When compared with chemically derived surfactants, biosurfactants are independent of mineral oil as a feedstock; they are readily biodegradable and can be produced at low temperatures. Biosurfactants can be applied in various areas, such as the nutrient, cosmetic, textile, varnish, pharmaceutical, mining, and oil recovery industries [64–66]. The lipopeptide antibiotic daptomycin is an Actinobacterial biosurfactant that has already entered the market and is used in the treatment of diseases caused by Gram-positive pathogens and has been marketed as Cubicin by Cubist Pharmaceuticals. Diverse types of biosurfactants or bioemulsifiers have been described to be produced within the class
Vitamin B12 as it exists in nature may be produced by bacteria or Actinobacteria . Isolation of vitamin B12 from Actinobacteria fermentations [68, 69] stirred up considerable interest in possible production of vitamin by microbial fermentations. Addition of cobalt salts to the media apparently acts as a precursor for all Actinobacteria to produce vitamin. As cobalt is a rather effective bactericidal agent, this precursor must be added carefully. The fermentations producing the antibiotics streptomycin, aureomycin, grisein, and neomycin will produce some vitamin B12 as well if the medium is supplemented with cobalt without apparently affecting the yields of antibiotic substances. Several other studies suggested that some Actinobacteria that are non–antibiotic-producing cultures produce more of this vitamin than those producing antibiotics. Actinobacteria have been shown to produce other water soluble vitamins, with special studies on production of thiamine and the pteroylglutamic acid derivative that is active in promoting the growth of certain strains of
As synthetic dyes have some limitations such as usage of hazardous chemicals for their production, creating worker safety concerns and generation of hazardous wastes, microbe-oriented pigments are of great concern. Specially, Actinobacteria are characterized by the production of various pigments on natural or synthetic media (Figure 8) and are considered as an important cultural characteristic in describing the organisms. Any phenotypic changes induced by environmental influences will help Actinobacteria as they boast distinctive colony morphologies and produce variety of pigments and aerial branching filaments called hyphae, which give them a characteristic fuzzy appearance. These pigments usually comes in various shades of blue, violet, red, rose, yellow, green, brown, and black, which may be dissolved into the medium or it may be retained in the mycelium. The pigments produced by
6.8. Nanoparticle synthesis
Nanoparticles are of great scientific interest as they bridge the gap between bulk materials and atomic or molecular structures. Generally, the chemical methods are low cost for high volume; however, their drawbacks include contamination from precursor chemicals, use of toxic solvents, and generation of hazardous byproducts. Hence, there is an increasing need to develop high-yield, low-cost, nontoxic, and environmentally benign procedures for synthesis of metallic nanoparticles. Therefore, the biological approach for synthesis of nanoparticles becomes important. In fact, Actinobacteria are efficient producers of nanoparticles, which show a range of biological properties, namely antibacterial, antifungal, anticancer, antibiofouling, antimalarial, antiparasitic, and antioxidant.
||Zinc, copper, manganese|
Actinobacteria possess many properties that make them good candidates for application in bioremediation of soils contaminated with organic pollutants. In some contaminated sites, Actinobacteria represent the dominant group among the degraders . They play an important role in the recycling of organic carbon and are able to degrade complex polymers. Sanscartier et al  reported that the greater use of petroleum hydrocarbons that are widely used in our daily life as chemical compounds and fuel has become one of the most common contaminants of large soil surfaces and eventually is considered as a major environmental problem. Some reports suggests that
6.10. Control of plant diseases
The worldwide efforts in the search of natural products for the crop protection market have progressed significantly, and Actinobacteria, especially genus
EF-76 and FP-54
|Grass seedling disease||
||Nigericin and guanidylfungin A|
|Root rot of Pea||
|Asparagus root diseases||
|Rice blast disease||
|Broad range of plant diseases||
|Sheath blight of rice||
|Brown rust of wheat||
|Phytophthora blight of pepper||
|Phytophthora blight of pepper||
|Damping-off of cabbage||
|Rice sheath blight||
||Polyoxin B and D|
|Rice root disease||
||2,3-dihydroxybenzoic acid, phenylacetic acid, cervinomycin A1 and A2|
|Blotch of wheat||
|Powdery mildew of cucumber||
||Neopeptin A and B|
6.11. Nematode control
It has been known for decades that effective control of plant-parasitic nematodes is dependent on chemical nematicides. Due to its ill effects with respect to the environmental hazards, hazardous nematicides have emphasized the need for new methods to control nematodes. Today, numerous microorganisms are recognized as antagonists of plant-parasitic nematodes. Especially, Actinobacteria have potential for use in biological control as they are known to produce antibiotics. The production of avermectins by a species of
6.12. Enhancement of plant growth
Despite the well-documented history of
6.13. Phytohormone production
The production of the plant hormone indole-3-acetic acid (IAA) and the pathways of its synthesis by various
Extensive use of chemical insecticides for controlling malaria, filaria, dengue, chickungunya, Japanese encephalitis, and other mosquitoes have resulted in hazards to the environment and caused development of resistance in vector mosquitoes. Accordingly, various biological control agents have gained importance with innumerable advantages over the chemical insecticides. At very low doses, these biolarvicides are highly effective against mosquito larvae and are completely safe to other nontarget organisms, environment, man, and wild life. Several varieties of microorganisms, including fungi, bacteria, and nematodes have been reported as strategies to biologically control the vectors. Specifically, Actinobacteria produce many important bioactive compounds of high commercial value and continue to be routinely screened for new bioactive substances. In a study made by Vijayan and Balaraman , extracellular secondary metabolites were produced from 35 different Actinobacterial isolates that showed high larvicidal activity against
6.15. Odor and flavor compounds production
Actinomycetes have long been associated with musty odors in water but their actual contribution to odor in freshwater was unknown. But in late 1960s, secondary metabolites, geosmin and 2-methylisoborneol (MIB), were identified from actinomycete cultures  after which actinomycetes have gained considerable importance throughout the water industry as major sources of drinking water taste and odor. Gaines and Collins  studied the metabolites of
2-isobutyl-3-methoxypyrazine or 2-isopropyl-3-methoxypyrazine
||Dimethyl trisulfide||Potato like|
7. Harmful effects
7.1. Actinobacterial plant diseases
A number of significant plant diseases are caused by Actinobacteria. Actinobacteria currently assigned to the genus
||B light of holly (flex opaca)|
||Wilt and leaf spot of red beet (
||Wilt of bean (
||Wilt and stunting of alfalfa (
||Canker of tomato (
||Wilt and blight of corn|
||Spot of tulip leaves and bulbs|
||Stem canker and leaf spot of poinsettia (
||Gumming of cereals|
||Wilt and tuber rot of potato (
||Galls and bud proliferation in blueberry
||Leaf gall in many plants, fasciation of sweet pea|
||Common scab of potato|
||Sweet potato scab|
||Common and russet scab of potatoes, sugar beet, etc|
7.2. Actinobacterial human and animal diseases
Actinobacteria have proved to be the causal agents of many human and animal infections, which include a number of common and intensively studied diseases, such as diphtheria, tuberculosis, and leprosy. There is also a wide range of infections that are less well known; some, like actinomycosis and nocardiosis, are proving to be more clinically significant than previously thought. In addition, it is becoming increasingly evident that
||Feet, legs, upper extremities, and other sites|
||Cervicofacial, thoracic, abdominal, and uterine regions|
|Bacterial kidney disease||
||Kidney, liver, spleen, and other internal organs|
|Dermatophilosis & streptothricosis||
||Throat, occasionally wounds|
||Lungs, lymph nodes, and skin|
||Lung, central nervous system, kidney, muscle, and other tissues|
||Any part of body surface, especially the extremities|
|Purulent infections including abscesses||
||Abscess formation in various organs (brain, spinal cord, and joints)|
|Pyelonephritis in cattle
Actinobacteria is one of the dominant groups of microorganisms that produce industrially important secondary metabolites. A wide range of antibiotics in the market is obtained from Actinobacteria. Products such as enzymes, herbicides, vitamins, pigments, larvicides, phytohormones, and surfactants are produced by these several genera of Actinobacteria, which are of great commercial value. They are capable of degrading a wide range of hydrocarbons, pesticides, and feather waste, and their metabolic potential offers a strong area for research. However, many of the rare genera of Actinobacteria have been neither discovered from unexplored locations nor employed for their biotechnological and industrial potential. Thus, studies on unique ecological environments could yield molecules that could become future harbingers of green technology.
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