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Araújo J, Rocha J, Filho M. O, Matias S, Júnior S. O, Padilha C, Santos E. Rhamnolipids Biosurfactants from Pseudomonas Aeruginosa - A Review. Biosci Biotech Res Asia 2018;15(4).
Manuscript received on : 21-Sep-2018
Manuscript accepted on : 27-Nov-2018
Published online on:  24-12-2018

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Rhamnolipids Biosurfactants from Pseudomonas Aeruginosa - A Review

Jaciara Araújo, Juliene Rocha, Marcos Oliveira Filho, Stephanie Matias, Sérgio Oliveira Júnior, Carlos Padilha and Everaldo Santos

Chemical Engineering Department – Federal University of Rio Grande do Norte, Candelária, Natal, Brazil.

Corresponding Author E-mail: everaldo@eq.ufrn.br


DOI : http://dx.doi.org/10.13005/bbra/2685

ABSTRACT: Studies addressing for ecological compatible products have been increased along time, especially, on biosurfactant field. Biosurfactants are extracellular amphiphilic compound that are mainly produced by microorganisms and are classified into five main groups, including the glycolipids one. Rhamnolipids are included in the latter and are anionic biosurfactants produced predominantly by Pseudomonas aeruginosa being classified as mono- and di-rhamnolipids. In addition, their production may occur from different carbon sources, which may be obtained from renewable and low-cost residue. Therefore, it is possible to reduce the rhamnolipids production cost, since this has been the main bottleneck for replacing the chemical surfactants. In addition, to meeting a bona fide industrial application some limitations such as low productivity as well as recovery and/or purification that represent from 60 to 80% of total production cost should be improved. Therefore, this review covers different ways for producing rhamnolipids covering their application in many fields such as pharmaceutical, agricultural, petrochemical and so on; demonstrating the versatility of these biological compounds.

KEYWORDS: Agro-Industrial Waste; Application; Rhamnolipid; Synthesis

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Araújo J, Rocha J, Filho M. O, Matias S, Júnior S. O, Padilha C, Santos E. Rhamnolipids Biosurfactants from Pseudomonas Aeruginosa - A Review. Biosci Biotech Res Asia 2018;15(4).

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Araújo J, Rocha J, Filho M. O, Matias S, Júnior S. O, Padilha C, Santos E. Rhamnolipids Biosurfactants from Pseudomonas Aeruginosa - A Review. Biosci Biotech Res Asia 2018;15(4). Available from: http://www.biotech-asia.org/?p=32191


Surfactants are chemical compounds synthesized by petroleum derivatives and capable of reducing the surface tension between two immiscible phases due to their amphiphilicity. Structurally, a surfactant molecule is composed of a hydrophilic and a hydrophobic moiety. 1,2 The polar moiety can be formed by carbohydrates, amino acids, carboxylic acids, phosphates or alcohols, while the apolar portion consists of carbon chains.3 This characteristic is essential in applications requiring emulsification, lubrication, foaming, solubilization of immiscible compounds or phase dispersion.4

In contrast, biosurfactants are metabolites produced by bacteria, filamentous fungi or yeasts. These amphiphilic and extracellular compounds were discovered in the 1960s through the fermentation of hydrocarbons and have many advantages compared to chemical surfactants. In the last 10 years, the biosurfactants received a lot of attention due to their low toxicity, high selectivity and biodegradability, low critical micellar concentration (CMC) and stability in drastic conditions of pH, salinity and temperature.5–7

Although biological surfactants have a wide range of structures, can be produced by different strains of microorganisms and water immiscible and miscible substrates, its low productivity and recovery hinders industrial scale production. Their upstream process costs can represent up to 30% of the total production cost. Meanwhile, recovery and purification steps amount between 60 and 80% of the total operating value, which explains the high values of marketable products based on biosurfactants (BS) and bioemulsifiers (BE). Thus, the use of renewable and low-cost substrates appears as an alternative to reducing these costs.8–10

Biosurfactants are divided into five main groups: glycolipids, phospholipids, lipopeptides, fatty acids and polymeric biosurfactants.11 Rhamnolipids (RLs) are one of the glycolipids, whose molecules are formed by a hydrophilic portion, containing one or two rhamnoses, and a lipophilic region, consisting of saturated or unsaturated fatty acids. In addition, depending on the amount of rhamnoses they can be classified into mono- and di-rhamnolipids.12

RLs are produced by different strains of Pseudomonas e.g. P. chlororaphis, P. plantarii, P. putida, P. fluorescens and P. aeruginosa. The latter being the most used in the studies. In order to produce the RLs, the submerged fermentation is a mode of production extensively explored. However, solid-state fermentation has advantages such as lower energy expenditure during cultivation, less use of solvent for extraction and no need for agitation unit. In this mode, the nutrient source is a solid residue, so the choice of the substrate is a very relevant point, since it will have to contain all the nutrients the microorganism needs to express the biosurfactant.13,14

The biosynthesis of RLs in Pseudomonas aeruginosa is controlled by environmental factors and the quorum sensing system (QS).15 This system is responsible for the regulation of approximately 10% of the P. aeruginosa genes. It coordinates several functions, including the formation of virulence agents, motility and production of exopolysaccharides. The QS also controls the synthesis of fundamental compounds for biofilms, such as rhamnolipids, lectins and siderophores.16

Among the properties of RLs, they are able to emulsify oils, reduce water surface tension from 72 mN/m to approximately 25-30 mN/m, reduce the interfacial tension between compounds of different polarities, and decrease the CMC to values between 10 and 200 mg/L.17–19 Due to these characteristics, the bio-product can be applied in agriculture and in the pharmaceutical, food, cosmetic and petrochemical industries. Studies have also shown that these compounds exhibit antimicrobial activities against Gram-positive and Gram-negative bacteria and fungi, and many of these micro-organisms are pathogenic.19–21

Thus, this study aims to highlight important points in the production of rhamnolipids by Pseudomonas aeruginosa, emphasizing the use of renewable and low-cost substrates, production methods (submerged and solid state), as well as their main applications.

Fermentative Process 

The choice of the fermentative process is of great importance, given that the conditions employed directly influence the productivity of the target biomolecule.22 The conditions of the applied fermentation process must be optimized and controlled so that the process can succeed and its increase of scale is favorable for the production of the biosurfactants, being economically feasible when compared to chemical surfactants.23

Submerged Fermentation (SmF)

SmF is a fermentative process that uses a liquid fermentative medium, composed of soluble nutrients, where the microorganisms develop and release the biomolecules of interest.24

For the most part, the production of biosurfactants is developed through this process. The volume of biosurfactants produced by SmF varies according to the type and the magnitude of the process, the reaction medium and the culture conditions involved.25

Some difficulties have been found in pilot or large-scale reactors in the production processes of these molecules.25,26 A great amount of foam is formed, when the biosurfactant are produced, as they are conducted with agitation and forced aeration, leading to the loss of biomass, nutrients and products contained in the foam that is expelled from the reactor, which decreases the production parameters or in extreme cases it makes the process unfeasible.27,28

Solid-State Fermentation (SSF)

SSF is a process that simulates the natural habitat of microorganisms. It occurs in the absence or near absence of free water, so the substrate must have sufficient moisture to maintain and grow the microorganism.29 The substrates used in SSF are quite diverse, namely, wheat bran, lemon and orange peel,30 sugar cane bagasse and coffee husk,31 corn bran,25 among others.

The differences between this process and the SmF are in the restricted availability of water, which may stimulate the excretion of some specific metabolites that are not produced in a liquid medium.29

This process presents several advantages, higher yields and volumetric productivity, lower operating costs, inexpensive culture medium (the use of agroindustrial waste as a substrate), greater oxygen distribution and, in general, lower energy demand.32,33

Thus, SSF presents a viable alternative for the production of high added value metabolic products, among them are biologically active secondary metabolites (toxins, antibiotics), enzymes, organic acids, amino acids, vitamins, ethanol and biopesticides.29,31–34

There are few papers in the literature addressing the production of biosurfactants by SSF, the only records found are related to the production of biosurfactants by bacteria or filamentous fungi. 25,35,36 SSF is a technique of simple application, especially in bench scale. In surfactant production, higher concentrations can be achieved, and the formation of foam is avoided as in SmF. However, there are obstacles to overcome, the increase of scale is hampered by heat and mass transfer in bioreactors because materials’ heterogeneity. Thereby, this process should be further explored to improve the application and production of biosurfactants. The development and projection of bioreactors capable of operating under the conditions the closest to optimal is ideal to the strengthening of SSF.14,37,38

Agroindustrial Waste 

Researchers are increasingly interested in biosurfactants, since they present physicochemical and surfactant characteristics that allow them to be applied to several areas such as petroleum (recovery, emulsification and refining), cosmetic. They also can act as antimicrobial and biomedical agents, in bioremediation, as food additives, in cleaning products and others.39–41

The successful implementation of biosurfactants in the industrial field is due to the efforts made by Jeneil Biosurfactant Co. (Saukville, Wisconsin), which carried out a batch fermentation process of up to 20,000 gallons.41,42

Interestingly, microorganisms are versatile to mediate the transformation of complex residues even under extreme conditions of pH, high salinity, pressure and temperature.43 It is recognized that the nutritional composition and the environment exposed to the microorganism directly influence its growth and, indirectly, the synthesis and ratio of the type of synthesized RLs.39,44,45

The use of biosurfactants from bioprocesses brings benefits to the environment and industrial process, since it is possible to reduce the cost of receiving the substrate, acquiring the agroindustrial residue (unexpensive and available in large quantities), and making use of sources of renewable carbon.42,44,46,47 Among the various agroindustrial residues (effluents from the refining of soybean oil, sunflower oil and palm oil), coconut and cashew residues, which presented promising results for the production of rhamnolipids.44,48,49

Other agroindustrial residues for the production of rhamnolipids targeting substrates of renewable and widely available origin are vegetable oils, sugars and glycerol. The use of glycerol is highlighted among these substrates, since it is highly consumed by yeasts and bacteria, and because of the excessive Brazilian production.39,46

Looking for alternatives to reduce costs of production of rhamnolipids, some authors reported data using lignocellulosic and agricultural residues,50 residues from the dairy industry, molasses and starch residues,51 crude oil,52 soybean sludge, chicken and hydrogenated vegetable fat.53

The use of biosurfactants in advanced oil recovery wells is an ancient technique discovered in the mid-1930s, but it has been improved and better understood in recent decades. This allowed the advances of the research in relation to the factors that favored the emulsification of the oil in the wells from the culture of the microorganisms. However, several difficulties prevent diffusion of biosurfactants, such as low yields, scale up for bioreactors, high production cost, among others.11

Among those agroindustrial residues, the use of soybean oil (37 g / L) in the cultivation of P. aeruginosa E03-40 for the production of RL was promising when compared to the use of glycerol (20 g / L). After optimization, the concentration of 42.1 g / L with an estimated yield of 47.3% was obtained under optimal conditions (10% dissolved oxygen and pH 5.7).54

Recent studies investigated the use of two or more agroindustrial residues for the production of biosurfactants based on RLs. A culture medium containing 5% animal fat and 2.5% corn steep liquor was used in the production of biosurfactants by Candida lipolytica, obtaining satisfactory results in the treatment of sites contaminated by heavy metals and petroleum derivatives.9 Among the emerging biotechnologies with application in the petroleum industry, there are those that make use of biosurfactants with the purpose of coordinating, reducing and treating the effluents generated from the oil processing.55

In light of what has been discussed previously, Table 1 shows the influence of different low-cost substrates and different strains of Pseudomonas aeruginosa on RL production. From this, it can be seen that most studies use SmF in the production of this biosurfactant. Thus, it is interesting that future works may approach SSF as an alternative to the production of RLs.

Table 1: Production of rhamnolipid by low-cost substrates.

Strain Fermentation Substrates Maximum yield (g/L) Ref.
P. aeruginosa PAO1 Submerged (250 mL1) Palm fatty acid distillate (PFDA) 0,43 [48]
P. aeruginosa DR1 Submerged (250 mL) Mango kernel 1,80 [56]
P. aeruginosa #112 Submerged (500 mL) Corn steep liquor (CSL) + molasses 3,20 [19]
P. aeruginosa LBI Submerged (2 L) Soapstock 15,9 [49]
P. aeruginosa PAO1 Submerged (1 L) Olive mill waste (OMW) 0,19 [57]
P. aeruginosa UFPEDA 614 Solid-state (250 mL) Sugarcane bagasse + corn bran 45,0 [25]
P. aeruginosa LBI Submerged (125 mL) Brazilian nut oil 9,90 [58]
P. aeruginosa DS10-129 Submerged (250 mL) Soybean Oil 4,31 [59]
P. aeruginosa UCP092 Submerged (500 mL) Glycerol 3,50 [60]
P. aeruginosa ATCC 10145 Submerged (250 mL) Sugarcane bagasse 9,10 [61]
P. aeruginosa LBI 2A1 Submerged (1 L) Crude glycerol 2,55 [62]
P. aeruginosa #112 Submerged (5 L) CSL + molasses + OMW 5,10 [4]

1Bioreactor volume.

Rhamnolipid Biosynthesis

Microorganisms and Pthways

Pseudomonas aeruginosa is a pathogenic Gram-negative bacterium responsible for producing virulence agents, toxins, alginates and lipopolysaccharides (LPS).63 The biosynthesis of RLs by P. aeruginosa occurs by means of three sequential enzymatic reactions, giving rise to mono-rhamnolipids or di-rhamnolipids, as shown in Figure 1.

In general, microorganisms consume hydrophilic substrates in the synthesis of the polar portion of the biosurfactant molecule, while the hydrophobic substrates are used exclusively in the hydrocarbon moiety.22 The production of RLs occurs by two metabolic pathways, which are responsible for forming the portions of that molecule. The hydrophobic region is formed by a long chain fatty acid, a hydroxy acid or an α-alkyl-β-hydroxy fatty acid, on the other hand the hydrophilic part may be a carbohydrate, a carboxylic acid, an alcohol or an amino acid.64

Hydrophilic substrates such as glucose or glycerol are degraded to form intermediates from glycolytic pathways, such as glucose 6-phosphate, which is one of the main precursors of carbohydrates present in the hydrophilic portion of biosurfactants. For the production of lipids, glucose is oxidized to pyruvate by means of glycolysis, and pyruvate is then converted to acetyl-CoA, which, together with oxaloacetate, produces malonyl-CoA and then fatty acid, one of the precursors for the synthesis of lipids.22

Figure 1 shows that the fatty acid is catalyzed by the enzyme Rh1G, responsible for diverting fatty acid synthesis intermediates into the biosynthetic pathway of RLs in P. aeruginosa. However, more recent studies have indicated that there is no enzyme above the rhamnosyltransferase (RhlA).65 This finding was based on the biochemical properties of the purified RhlA protein and its products when expressed heterologously in an E. coli host.66 Thus, RhlA catalyzes the formation of an ester bond between the fatty acid intermediates, 3-hydroxyalkyl-ACP, forming 3- (3-hydroxyalkanoyloxy) alkanoate (HAA), the lipid component of the RLs.66,67 Then RhlB catalyzes the formation of mono-rhamnolipids using trimethophosphate (dTDP) -L-rhamnose and HAA as precursors.68 For the synthesis of the di-rhamnolipids, a further rhamnosil (Rh1C) group is added to the structure using another molecule of (dTDP) -L-rhamnose, by an α-1,2-glycosidic bond.63

Rhamnose is found in several strains of Pseudomonas spp. as a component of LPS present in the cell wall of several Gram-negative bacteria. It is also present in exopolysaccharides (EPS) [63,65]. The formation of the hydrophilic portion of the RL molecule is initiated by the conversion of glucose-6-phosphate into glucose-1-phosphate through the action of the enzyme phosphoglyceromutase (AlgC). Followed by the action of rmlBDAC genes producing dTDP-L-rhamnose that will be precursor to the synthesis of RLs, as shown in Figure 1.

Regulation by Quorum Sensing (QS)

The QS corresponds to a bacterial signaling medium that is based on the production, during the cellular growth phase, of mediating molecules called auto-inducers. When a concentration limit is reached, these autoinducers interact with a transcriptional regulator, allowing the specific expression of a group of genes. One of the most studied intraspecies autoinducers is the N-acyl homoserine lactones (AHL) released by Gram-negative bacteria. There are more than 70 species of Gram-negative bacteria known to use AHL as a signaling molecule.69

Pseudomonas aeruginosa are able to grow within the host cells without damaging them until their population density reaches a sufficiently high value, necessary for biofilm formation. From this point, the microorganisms become aggressive to the host’s immune system causing diseases.16 The quantum detection system of P. aeruginosa regulates the production of several essential compounds in the formation of biofilms and acts on the release of extracellular DNA (eDNA). This bacterium has three known systems for the detection of quorum sensing, LasI / LasR, RhI / RhRR and Pseudomonas quinolone signaling system (PQS).70–72

In QS the main systems of regulation are las and rhl. Las and RhlI, catalytic enzymes in the synthesis process, produce the homoserine lactones 3OC12-HSL and C4-HSL signaling molecules, which bind and modulate their corresponding transcriptional regulators LasR and RhRR respectively. The system also requires the RsaL protein, and lasR, which negatively regulates the expression of both genes and indirectly affects the biosynthesis of RLs.73,74

A third signaling system based on 2-heptyl-3-hydroxy-4-quinolone, designated the signal Pseudomonas quinolone (PQS), was shown to be part of the quorum sensing regulatory network in P. aeruginosa.75 The biosynthesis of PQS is promoted by the pqsABCD gene products and binds to the LysR-type regulator PqsR (or MvfR). The expression of PqsR is directed by las and repressed by the rhl QS system, in a typically complex regulatory network. The production of PQS has a profile similar to that of RLs because it reaches its maximum in the stationary phase. Mutant genes of pqsR and pqsE decreased the levels of RLs synthesis, even when supplied with exogenous C4-HSL, indicating a direct relation of PqsR and PQS in the biosynthesis of RLs.76,77

Understanding the biosynthesis of RLs allows us to have conditions to solve everyday problems. For example, due to the high resistance of bacteria against most of the antibiotics on the market, it is necessary to develop techniques that inhibit this resistance. Therefore, recently, some studies have demonstrated anti-QS properties of natural herbal medicinal substances. Inhibition of the QS molecules requires the specific screening of several molecules with different chemical natures69

Figure 1: Rhamnolipid production pathways [45] Figure 1: Rhamnolipid production pathways [45]

Click here to View figure

Rhamnolipid Isoforms

The structural variety of the isoforms of RLs is defined by the presence of rhamnose and / or fatty acids as their chain length may vary from C8 to C14. There are 4 types of isoforms that can be classified as mono (RL1 and RL3) or di-rhamnolipid (RL2 and RL4), as can be seen in Figure 2.12,78

Figure 2: Chemical structure of different isoforms. RL1 (mono-rhamno-di-lipidic), RL2 (di-rhamno-di-lipidic), RL3 (mono-rhamno-mono-lipidic), RL4 (di-rhamno-mono-lipidic) [78,79]. Figure 2: Chemical structure of different isoforms. RL1 (mono-rhamno-di-lipidic), RL2 (di-rhamno-di-lipidic), RL3 (mono-rhamno-mono-lipidic), RL4 (di-rhamno-mono-lipidic) [78,79].

Click here to View figure

The properties presented by the RLs vary according to the composition of the homologues present in the medium and their distribution depends on the culture conditions (composition of the substrates, pH, and temperature), strain used and culture medium.80

Some studies show the formation of different isoforms. For example, when producing RLs from vegetable oils it was found that the mixture had di-rhamnolipid (Rha-Rha-C10-C10) and mono-rhamnolipid (Rha-C10-C10).58 In another investigation, the authors proved that the mixture of RLs synthesized by Pseudomonas aeruginosa MN1 and glycerol was composed of different homologues (Rha-C10, Rha-C8-C10, Rha-C10-C8, Rha-C10-C12: 1, Rha C10-C10, Rha-C10-C10, Rha-C10-C10, Rha-C10-C10, Rha-Rha-C8-C10, Rha-Rha-C10-C8, Rha-Rha-C8-C12: 1, Rha-Rha-C10-C10 , Rha-Rha-C10-C12: 1, Rha-Rha-C12: 1-C10, Rha-Rha-C10 -C12, Rha-Rha-C12 -C10, Rha-Rha-C8 -C8) 35% were mono-rhamnolipids. In addition, the homologue greater quantity had better CMC and surface tension results when compared to the isolated di-rhamnolipid and the mixture of the two homologues.80

Numerous studies have shown that different strains of Pseudomonas aeruginosa when combined with several substrates have fairly variable amounts of homologues.81–85

Biosurfactant Application

Although the biosurfactants have some superior characteristics in relation to the chemical surfactants, they still have very limited applications, mainly because of the cost of production.11 However, there are potential applications where the biological origin promises better biocompatibility and good microbial degradability.86 Therefore, as the cost of production becomes lower, its application should become more generalized, not surprisingly a large number of laboratory researches for the most diverse areas of application. The following topics explore the various uses of RLs proposed in the scientific literature.


Bioremediation can be defined as a process whereby organic wastes are biologically removed or degraded for cleaning of oil spills and treatment of terrestrial and aquatic environments contaminated with xenobiotics.50 These processes appear as an innovative technology in the removal of compounds derived from petroleum, among other pollutants, against chemical surfactants that have high toxicity and non-biodegradable properties.87

It was shown that by adding RLs to pure P. aeruginosa cultures there was an increase in biodegradation of hexadecane, octadecane, n-paraffins and phenanthrene, as well as degradation in soil systems in the presence of hexadecane, tetradecane, pristine, creosote or hydrocarbon mixtures.88,89

Although there are some studies that report the positive effects on the biodegradation of petroleum hydrocarbons in the presence of biosurfactant, there are reports that both in pure addition as in soil systems there was an inhibition of biodegradation by the addition of RLs. This inhibition can occur due to the preference of RLs as carbon source for bacterial metabolism.90

Some previous research suggested that increased degradation by the presence of RLs may occur by increasing the solubility of the hydrocarbon then increasing bioavailability for cell degradation or interaction with the degrading cell, causing the cell surface to become more hydrophobic and interacting more easily with the hydrophobic substrates.91 This second mechanism becomes more economically and environmentally interesting, since a large amount of RLs is required to increase the solubility of the hydrocarbon, whereas to alter the cell surface the required amount is smaller. Indeed, as RLs are the preferred source of carbon, higher concentrations would decrease the degradation of the hydrocarbons.

The decontamination of areas contaminated with heavy metals is a relatively few explored field. The presence of these may inhibit degradation of organic compounds. In a research carried out on systems contaminated with organic compounds, cadmium and naphthalene in the presence of RLs, the cadmium toxicity reduced, which led to the increase of the biodegradation of naphthalene.92 Another study observed a decrease in the inhibition of phenanthrene mineralization in the presence of cadmium by pulsed addition of RLs.90 There are reports of higher copper and nickel removal rates from sediments by adding RLs to a 1% NaOH solution.93

Food Industry

RLs present some properties, such as formulation and stabilization of emulsions, as well as anti-adherence and antimicrobial activity, which makes them interesting for the food industry, particularly in increasing food shelf-life without concerning consumer, eliminating the need for addition of synthetic preservatives. RLs can be used directly to avoid contamination of food, as a food additive, or indirectly, as a detergent formulation to clean surfaces that come into contact with food.94 Other functions performed in the food industry by RLs is acting to improve the stability of the dough, texture, volume and preservation of bakery products obtained by the addition of surfactants.94 It is also possible to use them to improve the properties of frozen butter cream, croissants and confectionery.95 They can also serve as a source of L-rhamnose for the synthesis of food flavors, which has already been successfully obtained by the hydrolysis of surfactants produced by P. aeruginosa.96

In addition to their obvious role as surface and interfacial tension reducing agent, RLs may have other functions in foods aiding in the general blending of ingredients and may also retard the growth of fungi and some bacteria.97 It has also been shown that they can be successfully exploited to break down biofilms from individual and mixed cultures of foodborne pathogens.98


RLs influence nonspecific immunity in plants and induce resistance and are considered potential alternatives to reduce or replace pesticides in agriculture.99 Some studies on the effect of RLs on plants and pests showed that they are capable of stimulating defense genes in tobacco and are potent protectors in monocotyledonous plants against biotrophic fungi Other studies proved that RLs could improve wettability of leaf surfaces.100 Antifitoviral effects were observed for virus / host combinations of tobacco mosaic virus / Nicotiana glutinosa and potato X virus / Nicotiana tabacum.101

Pure mono and di-rhamnolipids were tested in three species representatives of the zoosporic phytopathogen genera, namely, Pythium aphanidermatum, Phytophtora capsici and Plasmopara lactucaeradicis. They showed the ability to control certain pathogens. At concentrations of 5 to 30 mg L-1, both RLs caused a cessation of motility and lysis of the entire zoospora population within 1 min.102 This observation led to the development of a biofungicide formulation containing RLs, used to avoid the contamination of crops by pathogenic fungi.

These molecules are also useful in the removal of polyaromatic hydrocarbons and pentachlorophenol from the soil and may facilitate the uptake of nutrients and fertilizers through the roots.103 Due to their anionic nature, they are able to remove toxic metals from agriculture. However, the success in increasing the recovery of heavy metals will also depend on the amount of RLs present in the aqueous phase.104

Agricultural lands containing petroleum hydrocarbons that are important contaminants can be remedied by an introduction of RLs in the soil due to their high solubilization and increase of the bioavailability properties in some inaccessible compounds.105 Other applications of RLs found in the literature in the field involve eradication of the disease caused by P. capsici in pepper plants (Capsicum annuum) and control of a serious pathogen for certain tomato crops. In addition, it has been suggested the application in the treatment and prevention of overwatering during irrigation due to its wettability properties facilitating the breaking of impenetrable barriers to the water and allowing the water to easily reach the soil spreading more evenly99

Pharmaceutical and Cosmetic Industry

Several cosmetic creams exhibit, in their formula, essential oils from plants due to their occlusive, emollient and moisturizing properties in the skin. Many of these oil-based substances requires the presence of a stabilizing agent as emulsifiers and / or surfactants in order to obtain good emulsions. Other functions of cosmetic surfactants are detergency, wetting, solubilization, dispersion and foaming effects.106 Natural surfactants besides performing these functions have benefits such as biodegradability, low toxicity and acceptability, making them in high demand.

The application of RLs in cosmetics and pharmaceuticals as emulsifiers, penetrating agents and drug delivery systems is still an emerging area of research. Surfactants as emulsifiers, foaming agents, solubilizers, wetting agents, cleaning agents, antimicrobial agents and enzymatic mediators in various dosage forms such as creams, lotions, liquids, pastes, powders, sticks, gels, films and sprays can be replaced by biosurfactants. Patents were granted for cosmetics containing RLs for anti-wrinkle and anti-aging products,98 which were released as commercial cosmetics for skin care because of their skin compatibility and extremely low skin irritation.97 Briefly, some applications of RLs produced by different strains of Pseudomonas aeruginosa are presented in Table 2.

Table 2: Rhamnolipid Application.

Strain Carbon Source Application References
Pseudomonas aeruginosa AP 029/GLVIIA Glucose Enzymatic hydrolysis [108]
Pseudomonas aeruginosa L2-1 Casssava wastewater Crude oil removal [109]
Pseudomonas aeruginosa 1501 Water-soluble diesel Antimicrobial agent [110]
Pseudomonas aeruginosa PA1 Glycerol Nano/micropheres formulations [111]


The synthesis of rhamnolipids occurs mainly by submerged fermentation, however, solid-state fermentation presents advantages according to some studies. In addition, the use of renewable and low-cost substrates, such as agroindustrial waste, make the production process more interesting, economically. Another point corresponding to the diversity of chemical structures formed when working with different substrates and strains of Pseudomonas aeruginosa, the four existing isoforms constitute numerous homologs of rhamnolipids that vary according to the amount of rhamnose and extension of the carbonic chain. Finally, researches on applications of this bioproduct makes the study even more interesting, since it is possible to analyze its future commercial outcomes and, mainly, to substitute chemical surfactants by biological eco-friendly surfactants.

Conflicts of Interest

There is no conflict of interest.


  1. Otzen D. E. Biosurfactants and surfactants interacting with membranes and proteins: Same but different? Biochim. Biophys. Acta – Biomembr. 2017;1859:639–649. doi:10.1016/j.bbamem. 2016.09.024.
  2. Varjani S. J., Upasani V. N. Critical review on biosurfactant analysis, purification and characterization using rhamnolipid as a model biosurfactant. Bioresour. Technol. 2017;232:389–397. doi:10.1016/j.biortech.2017.02.047.
  3. de Silva R. C. F. S.,Almeida D. G.,Brasileiro P. P. F., Rufino R. D., Luna J. M., Sarubbo L. A. Biosurfactant Formulation of Pseudomonas cepacia and Application in the Removal of Oil from Coral Reef. Chem. Eng. Trans. 2017;57:649–654. doi:10.3303/CET 1757109.
  4. Gudiña E. J., Rodrigues A. I., de Freitas V., Azevedo Z., Teixeira J. A., Rodrigues L. R. Valorization of agro-industrial wastes towards the production of rhamnolipids. Bioresour. Technol. 2016;212:144–150. doi:10.1016/j.biortech. 2016.04.027.
  5. Araújo L. V., Freire D. M. G.,Nitschke M. Biossurfactantes: propriedades anticorrosivas, antibiofilmes e antimicrobianas. Quim. Nova. 2013;36:848–858. doi:10.1590/S0100-40422013000600019.
  6. Khan A. H. A., Tanveer S., Alia S.,Anees M.,Sultan A.,Iqbal M.,Yousaf S. Role of nutrients in bacterial biosurfactant production and effect of biosurfactant production on petroleum hydrocarbon biodegradation. Ecol. Eng. 2017;104:158–164. doi:10.1016/j.ecoleng. 2017.04.023.
  7. Souza K. S. T., Gudiña E. J., Schwan R. F.,Rodrigues L. R., Dias D. R., Teixeira J. A. Improvement of biosurfactant production by Wickerhamomyces anomalus CCMA 0358 and its potential application in bioremediation. J. Hazard. Mater. 2018:346;152–158. doi:10.1016/j.jhazmat. 2017.12.021.
  8. Banat I. M., Satpute S. K.,Cameotra S. S.,Patil R.,Nyayanit N. V. Cost effective technologies and renewable substrates for biosurfactants’ production. Front. Microbiol. 2014;5:1–18. doi:10.3389/fmicb. 2014.00697.
  9. Santos D. K. F., Resende A. H. M., de Almeida D. G., da Silva R. de C. F. S. Rufino R. D.,Luna J. M., Banat I. M., Sarubbo L. A. Candida lipolytica UCP 0988 biosurfactant: Potential as a bioremediation agent and in formulating a commercial related product. Front. Microbiol. 2017;8:1–11. doi:10.3389/fmicb. 2017.00767.
  10. Souza K. S. T., Gudiña E. J., Schwan R. F., Rodrigues L. R.,Dias D. R.,Teixeira J. A. Improvement of biosurfactant production by Wickerhamomyces anomalus CCMA 0358 and its potential application in bioremediation. J. Hazard. Mater. 2018;346:152–158. doi:10.1016/j.jhazmat. 2017.12.021.
  11. Geetha S. J., Banat I. M.,Joshi S. J. Biosurfactants: Production and Potential applications in Microbial Enhanced Oil Recovery (MEOR). Biocatal. Agric. Biotechnol. 2018;1–30. doi:10.1016/j.jpcs. 2015.12.001.
  12. Gogoi D., Bhagowati P.,Gogoi P., Bordoloi N. K., Rafay A., Dolui S. K., Mukherjee A. K. Structural and physico-chemical characterization of a dirhamnolipid biosurfactant purified from Pseudomonas aeruginosa: application of crude biosurfactant in enhanced oil recovery. RSC Adv. 2016;6:70669–70681. doi:10.1039/C6RA 11979D.
  13. Randhawa K. K. S., Rahman P. K. S. M. Rhamnolipid biosurfactants-past, present and future scenario of global market. Front. Microbiol. 2014;5:1–7. doi:10.3389/fmicb. 2014.00454.
  14. Nalini S., Parthasarathi R. Optimization of rhamnolipid biosurfactant production from Serratia rubidaea SNAU02 under solid-state fermentation and its biocontrol efficacy against Fusarium wilt of eggplant. Ann. Agrar. Sci. 2018;16:108–115. doi:10.1016/j.aasci. 2017.11.002.
  15. Dobler L., Vilela L. F., Almeida  R. V., Neves B. C. Rhamnolipids in perspective: Gene regulatory pathways, metabolic engineering, production and technological forecasting. N. Biotechnol. 2016;33:123–135. doi:10.1016/j.nbt. 2015.09.005.
  16. Kariminik A., Baseri-Salehi M., Kheirkhah B. Pseudomonas aeruginosa quorum sensing modulates immune responses: An updated review article. Immunol. Lett. 2017;190:1–6. doi:10.1016/j.imlet. 2017.07.002.
  17. Das P.,Yang X. P., Ma L. Z. Analysis of biosurfactants from industrially viable Pseudomonas strain isolated from crude oil suggests how rhamnolipids congeners affect emulsification property and antimicrobial activity. Front. Microbiol. 2014;5:1–8. doi:10.3389/fmicb. 2014.00696.
  18. . De França Í. W. L., Lima A. P., Lemos J. A. M.,Lemos C. G. F.,Melo V. M. M., De Sant’ana H. B.,Gonçalves L. R. B. Production of a biosurfactant by Bacillus subtilis ICA56 aiming bioremediation of impacted soils. Catal. Today. 2015;255:10–15. doi:10.1016/j.cattod. 2015.01.046.
  19. Gudiña E. J., Rodrigues A. I.,Alves E., Domingues M. R., Teixeira J. A., Rodrigues L. R. Bioconversion of agro-industrial by-products in rhamnolipids toward applications in enhanced oil recovery and bioremediation. Bioresour. Technol. 2015;177:87–93. doi:10.1016/j.biortech. 2014.11.069.
  20. Magalhães L.,Nitschke M. Antimicrobial activity of rhamnolipids against Listeria monocytogenes and their synergistic interaction with nisin. Food Control 2013;29:138–142. doi:10.1016/j.foodcont. 2012.06.009.
  21. Semedo-Lemsaddek T.,Pedroso N. M.,Freire D., Nunes T.,Tavares L.,Verdade L. M., Oliveira M. Otter fecal enterococci as general indicators of antimicrobial resistance dissemination in aquatic environments. Ecol. Indic. 2018;85:1113–1120. doi:10.1016/j.ecolind. 2017.11.029.
  22. Fontes G. C.,Amaral P. F. F., Coelho M. A. Z. Produção de biossurfactante por levedura. Quim. Nov. 2008;31:2091–2099.
  23. Banat I. M.,Franzetti A.,Gandolfi I.,Bestetti G.,Martinotti M. G., Fracchia L., Smyth T. J., Marchant R. Microbial biosurfactants production, applications and future potential. Appl. Microbiol. Biotechnol. 2010;87:427–444. doi:10.1007/s00253-010-2589-0.
  24. Gibbs P. A.,Seviour R. J., Schmid F. Growth of filamentous fungi in submerged culture: problems and possible solutions. Crit. Rev. Biotechnol. 2000;20:17–48. doi:10.1080/07388550091144177.
  25. Camilios-Neto D., Bugay C.,de Santana-Filho A. P.,Joslin T.,de Souza L. M.,Sassaki G. L.,Mitchell D. A., Krieger N. Production of rhamnolipids in solid-state cultivation using a mixture of sugarcane bagasse and corn bran supplemented with glycerol and soybean oil. Appl. Microbiol. Biotechnol. 2011;89:1395–1403. doi:10.1007/s00253-010-2987-3.
  26. Lotfabad T. B.,Ebadipour N., Roostaazad R. Evaluation of a recycling bioreactor for biosurfactant production by Pseudomonas aeruginosa MR01 using soybean oil waste. J. Chem. Technol. Biotechnol. 2015;91:1368–1377. doi:10.1002/jctb.4733.
  27. Lee B. S., Kim E. K. Lipopeptide production from Bacillus sp. GB16 using a novel oxygenation method. Enzyme Microb. Technol. 2004;35:639–647. doi:10.1016/j.enzmictec. 2004.08.017.
  28. Yeh M. S., Wei Y. H., Chang J. S. Bioreactor design for enhanced carrier-assisted surfactin production with Bacillus subtilis. Process Biochem. 2006;41:1799–1805. doi:10.1016/j.procbio. 2006.03.027.
  29. Pandey A. Solid-state fermentation. Biochem. Eng. J. 2003;13:81–84. doi:10.1016/S1369-703X(02)00121-3.
  30. Ortiz G. E.,Ponce-Mora M. C., Noseda D. G.,Cazabat G.,Saravalli C., López M. C., Gil G. P., Blasco M.,Albertó E. O. Pectinase production by Aspergillus giganteus in solid-state fermentation: optimization, scale-up, biochemical characterization and its application in olive-oil extraction. J. Ind. Microbiol. Biotechnol. 2016;44:197–211. doi:10.1007/s10295-016-1873-0.
  31. Vandenberghe L. P. S.,Soccol C. R., Pandey A., Lebeault J. M. Solid-state fermentation for the synthesis of citric acid by Aspergillus niger. Bioresour. Technol. 2000;74:175–178. doi:10.1016/S0960-8524(99)00107-8.
  32. Ramos-Sanchez L. B., Cujilema-Quitio M. C., Julian-Ricardo M. C.,Cordova J., Fickers P. Fungal Lipase Production by Solid-State Fermentation. J. Bioprocess. Biotech. 2015;05:1–9. doi:10.4172/2155-9821.1000203.
  33. Hosseinpour M. N., Najafpour G. D., Younesi H.,Khorrami M., Vaseghi Z. Lipase production in solid state fermentation using aspergillus niger: Response surface methodology. Int. J. Eng. Trans. B Appl. 2012;25:151–159. doi:10.5829/idosi.ije. 2012.25.03b.01.
  34. Hölker U., Lenz J. Solid-state fermentation – Are there any biotechnological advantages? Curr. Opin. Microbiol. 2005;8:301–306. doi:10.1016/j.mib. 2005.04.006.
  35. Ohno A., Ano T.,Shoda M. Production of a lipopeptide antibiotic, surfactin, by recombinant Bacillus subtilis in solid state fermentation. Biotechnol. Bioeng. 1995;47:209–214. doi:10.1002/bit.260470212.
  36. Das K., Mukherjee A. K. Comparison of lipopeptide biosurfactants production by Bacillus subtilis strains in submerged and solid state fermentation systems using a cheap carbon source: Some industrial applications of biosurfactants. Process Biochem. 2007;42:1191–1199. doi:10.1016/j.procbio. 2007.05.011.
  37. Durand A. Bioreactor design for solid state fermentation. 2003;13:113–125.
  38. Bahry H., Pons A., Abdallah R., Pierre G., Delattre C., Fayad N., Taha S., Vial C. Valorization of carob waste: Definition of a second-generation bioethanol production process. Bioresour. Technol. 2017;235:25–34. doi:10.1016/j.biortech.2017.03.056.
  39. Sousa J. R., Correia J. A. C., Melo V. M. M., Gonçalves L. R. B., Cruz A. J. G. Cinética e caracterização de ramnolipídeos produzidos por pseudomonas aeruginosa msic02 utilizando glicerol como fonte de carbono. Quim. Nova. 2014;37:431–441. doi:10.5935/0100-4042.20140064.
  40. Müller M. M., Kügler J. H., Henkel M.,Gerlitzki M.,Hörmann B.,Pöhnlein M.; Syldatk C., Hausmann R. Rhamnolipids-Next generation surfactants? J. Biotechnol. 2012;162:366–380. doi:10.1016/j.jbiotec. 2012.05.022.
  41. Almeida D. G. d., de Soares R. C. F.,   da S., Luna J. M., Rufino R. D., Santos V. A., Banat I. M.,Sarubbo L. A. Biosurfactants: Promising molecules for petroleum biotechnology advances. Front. Microbiol. 2016;7:1–14. doi:10.3389/fmicb. 2016.01718.
  42. Rufino R. D., de Luna J. M.,de Takaki G. M.C.,Sarubbo L. A. Characterization and properties of the biosurfactant produced by Candida lipolytica UCP 0988. Electron. J. Biotechnol. 2014;17:34–38. doi:10.1016/j.ejbt. 2013.12.006.
  43. Montiel C.,Quintero R.,Aburto J. Petroleum biotechnology : Technology trends for the future. J. Biotechnol. 2009;8:2653–2666.
  44. Magalhães E. R. B., Silva F. L., dos Sousa M. A. S. B.,Santos E. S. dos Use of Different Agroindustrial Waste and Produced Water for Biosurfactant Production. Biosci. Biotechnol. Res. Asia. 2018;15:17–26. doi:10.13005/bbra/2604.
  45. Soberón-Chávez G., Lépine F., Déziel E. Production of rhamnolipids by Pseudomonas aeruginosa. Appl. Microbiol. Biotechnol. 2005;68:718–725. doi:10.1007/s00253-005-0150-3.
  46. Henkel M., Müller M. M.,Kügler J. H., Lovaglio R. B.,Contiero J.,Syldatk C., Hausmann R. Rhamnolipids as biosurfactants from renewable resources: Concepts for next-generation rhamnolipid production. Process Biochem. 2012;47:1207–1219. doi:10.1016/j.procbio. 2012.04.018.
  47. Makkar R. S.,Cameotra S. S. An update on the use of unconventional substrates for biosurfactant production and their new applications Received: J. Surfactants Deterg. 2002;5:11–17. doi:10.1007/s11743-002-0199-8.
  48. Radzuan M. N.,Banat I. M., Winterburn J. Production and characterization of rhamnolipid using palm oil agricultural refinery waste. Bioresour. Technol. 2017;225:99–105. doi:10.1016/j.biortech. 2016.11.052.
  49. Benincasa M., Contiero J.,Manresa M. A., Moraes I. O. Rhamnolipid production by Pseudomonas aeruginosa LBI growing on soapstock as the sole carbon source. J. Food Eng. 2002;54:283–288. doi:10.1016/S0260-8774(01)00214-X.
  50. Mukherjee S., Das P.,Sen R. Towards commercial production of microbial surfactants. Trends Biotechnol. 2006;24:509–515. doi:10.1016/j.tibtech. 2006.09.005.
  51. Joshi S.,Bharucha C., Jha S., Yadav S., Nerurkar A.,Desai A. J. Biosurfactant production using molasses and whey under thermophilic conditions. Bioresour. Technol. 2008;99:195–199. doi:10.1016/j.biortech. 2006.12.010.
  52. Shah M. U. H., Sivapragasam M., Moniruzzaman M., Yusup S. B. A comparison of Recovery Methods of Rhamnolipids Produced by Pseudomonas Aeruginosa. Procedia Eng. 2016;148:494–500. doi:10.1016/j.proeng. 2016.06.538.
  53. Costa S. G. V. A., Nitschke M.,Contiero J. Produção de biotensoativos a partir de resíduos de óleos e gorduras. Ciência e Tecnol. Aliment. 2008;28:34–38. doi:10.1590/S0101-20612008000100007.
  54. Sodagari M., Invally K., Ju L. K. Maximize rhamnolipid production with low foaming and high yield. Enzyme Microb. Technol. 2018;110:79–86. doi:10.1016/j.enzmictec. 2017.10.004.
  55. de  Silva R. C. F. S.,Almeida D. G., Rufino R. D., Luna J. M., Santos V. A.,Sarubbo L. A. Applications of biosurfactants in the petroleum industry and the remediation of oil spills. Int. J. Mol. Sci. 2014;15:12523–12542. doi:10.3390/ijms150712523.
  56. Reddy K. S., Khan M. Y.,Archana K.,Reddy M. G., Hameeda B. Utilization of mango kernel oil for the rhamnolipid production by Pseudomonas aeruginosa DR1 towards its application as biocontrol agent. Bioresour. Technol. 2016;221:291–299. doi:10.1016/j.biortech.2016.09.041.
  57. Ramírez I. M., Tsaousi K., Rudden  M., Marchant R.,  Alameda E. J.,  Román M. G., Banat I. M. Rhamnolipid and surfactin production from olive oil mill waste as sole carbon source. Bioresour. Technol. 2015;198:231–236. doi:10.1016/j.biortech. 2015.09.012.
  58. Costa S. G. V. A. O., Nitschke M., Haddad R., Eberlin M. N.,Contiero J. Production of Pseudomonas aeruginosa LBI rhamnolipids following growth on Brazilian native oils. Process Biochem. 2006;41:483–488. doi:10.1016/j.procbio. 2005.07.002.
  59. Rahman K. S. M., Rahman T. J., McClean S.,Marchant R.,Banat I. M. Rhamnolipid Biosurfactant Production by Strains of Pseudomonas aeruginosa Using Low-Cost Raw Materials. Biotechnol. Prog. 2002;18:1277–1281. doi:10.1021/bp020071x.
  60. Silva S. N. R. L., Farias C. B. B.,Rufino R. D., Luna J. M., Sarubbo L. A. Glycerol as substrate for the production of biosurfactant by Pseudomonas aeruginosa UCP0992. Colloids Surfaces B Biointerfaces 2010;79:174–183. doi:10.1016/j.colsurfb. 2010.03.050.
  61. Lopes V.,dos S., Fischer J., Pinheiro T. M. A.,Cabral B. V., Cardoso V. L.,Filho U. C. Biosurfactant and ethanol co-production using Pseudomonas aeruginosa and Saccharomyces cerevisiae co-cultures and exploded sugarcane bagasse. Renew. Energy. 2017;109:305–310. doi:10.1016/j.renene. 2017.03.047.
  62. Salazar-Bryam A. M.,Lovaglio R. B., Contiero J. Biodiesel byproduct bioconversion to rhamnolipids: Upstream aspects. Heliyon. 2017;3:e00337.doi:10.1016/j.heliyon. 2017.e00337.
  63. Rahim R.,Burrows L. L., Monteiro M. A.,Perry M. B., Lam J. S. Involvement of the rml locus in core oligosaccharide and O polysaccharide assembly in Pseudomonas aeruginosa. Microbiology. 2000;146:2803–2814. doi:10.1099/00221287-146-11-2803.
  64. Banat I. M., Desai J. D. Microbial production of surfactants and their commercial potential. Microbiol. Mol. Biol. Rev. 1997;61:47–64. doi:10.1016/S0140-6701(97)84559-6.
  65. Abdel-Mawgoud A. M.,Hausmann R., Lépine F., Muller M. M.,Déziel E. Rhamnolipids: detection, analysis, biosynthesis, genetic regulation and bioengineering of production Biosurfactants: from genes to applications. 2011;20. ISBN 978-3-642-14489-9.
  66. Zhu K., Rock C. O. RhlA converts ␤-hydroxyacyl-acyl carrier protein intermediates in fatty acid synthesis to the ??-hydroxydecanoyl-??-hydroxydecanoate component of rhamnolipids in Pseudomonas aeruginosa. J. Bacteriol. 2008;190:3147–3154. doi:10.1128/JB. 00080-08.
  67. Déziel E.,Lépine F.,Milot S., Villemur R. rhlA is required for the production of a novel biosurfactant promoting swarming motility in Pseudomonas aeruginosa: 3-(3-hydroxyalkanoyloxy)alkanoic acids (HAAs), the precursors of rhamnolipids. Microbiology. 2003;149:2005–2013. doi:10.1099/mic.0.26154-0.
  68. Miller D. J.,Zhang Y. M.,Rock C. O.,White S. W. Structure of RhlG, an essential ??-ketoacyl reductase in the rhamnolipid biosynthetic pathway of Pseudomonas aeruginosa. J. Biol. Chem. 2006;281:18025–18032. doi:10.1074/jbc.M601687200.
  69. Bouyahya A.,Dakka N., Et-touys A.,Abrini J.,Bakri Y. Medicinal plant products targeting quorum sensing for combating bacterial infections. Asian Pac. J. Trop. Med. 2017;10:729–743. doi:10.1016/j.apjtm.2017.07.021.
  70. Tielker D.,Hacker S., Loris R., Strathmann M.,Wingender J.,Wilhelm S.,Rosenau F., Jaeger K. E. Pseudomonas aeruginosa lectin LecB is located in the outer membrane and is involved in biofilm formation. Microbiology. 2005;151:1313–1323. doi:10.1099/mic.0.27701-0.
  71. Rasamiravaka T., Labtani Q., Duez P., El Jaziri M. The formation of biofilms by pseudomonas aeruginosa: A review of the natural and synthetic compounds interfering with control mechanisms. Biomed Res. Int. 2015;2015. doi:10.1155/2015/759348.
  72. Pham T. H.,Webb J. S.,Rehm B. H. A. The role of polyhydroxyalkanoate biosynthesis by Pseudomonas aeruginosa in rhamnolipid and alginate production as well as stress tolerance and biofilm formation. Microbiology.  2004;150:3405–3413. doi:10.1099/mic.0.27357-0.
  73. Dekimpe V.,Déziel E. Revisiting the quorum-sensing hierarchy in Pseudomonas aeruginosa: The transcriptional regulator RhlR regulates LasR-specific factors. Microbiology. 2009;155:712–723. doi:10.1099/mic.0.022764-0.
  74. Rampioni G.,Polticelli F., Bertani I.,Righetti K.,Venturi V., Zennaro E., Leoni L. The Pseudomonas quorum-sensing regulator RsaL belongs to the tetrahelical superclass of H-T-H proteins. J. Bacteriol. 2007;189:1922–1930. doi:10.1128/JB.01552-06.
  75. Gallagher L. A., McKnight S. L., Kuznetsova M. S.,Pesci E. C., Manoil C. Functions required for extracellular quinolone signaling by Pseudomonas aeruginosa. J. Bacteriol. 2002;184:6472–6480. doi:10.1128/JB.184.23.6472-6480.2002.
  76. Medina G.,Juárez K.,Díaz R., Soberón-Chávez G. Transcriptional regulation of Pseudomonas aeruginosa rhlR, encoding a quorum-sensing regulatory protein. Microbiology. 2003;149:3073–3081. doi:10.1099/mic.0.26282-0.
  77. Diggle S. P.,Winzer K.,Chhabra S. R.,Worrall K. E.,Cámara M., Williams P. The Pseudomonas aeruginosa quinolone signal molecule overcomes the cell density-dependency of the quorum sensing hierarchy, regulates rhl-dependent genes at the onset of stationary phase and can be produced in the absence of LasR. Mol. Microbiol. 2003;50:29–43. doi:10.1046/j.1365-2958.2003.03672.x.
  78. Mulligan C. N. Environmental applications for biosurfactants. Environ. Pollut. 2005;133:183–198. doi:10.1016/j.envpol.2004.06.009.
  79. Abdel-Mawgoud A. M.,Lépine F., Déziel E. Rhamnolipids: Diversity of structures, microbial origins and roles. Appl. Microbiol. Biotechnol. 2010;86:1323–1336. doi:10.1007/s00253-010-2498-2.
  80. Samadi N., Abadian N., Ahmadkhaniha R.,Amini F.,Dalili D.,Rastkari N.,Safaripour E.,Mohseni F. A. Structural characterization and surface activities of biogenic rhamnolipid surfactants from Pseudomonas aeruginosa isolate MN1 and synergistic effects against methicillin-resistant Staphylococcus aureus. Folia Microbiol. (Praha). 2012;57:501–508. doi:10.1007/s12223-012-0164-z.
  81. Mondal M. H., Sarkar A., Maiti T. K., Saha B. Microbial assisted (pseudomonas sp.) production of novel bio-surfactant rhamnolipids and its characterisation by different spectral studies. J. Mol. Liq. 2017;242:873–878. doi:10.1016/j.molliq.2017.07.089.
  82. Pereira J., Gudiña E., Dória M.,Domingues M., Rodrigues L., Teoxeira J., Coutinho J. Characterization by electrospray ionization and tandem mass spectrometry of rhamnolipids produced by two Pseudomonas aeruginosa strains isolated from Brazilian crude oil. Eur. J. Mass Spectrom. 2012;18:399–406. doi:10.1255/ejms.1194.
  83. Ndlovu T.,Rautenbach M.,Vosloo J. A.,Khan S.,Khan W. Characterisation and antimicrobial activity of biosurfactant extracts produced by Bacillus amyloliquefaciens and Pseudomonas aeruginosa isolated from a wastewater treatment plant. AMB Express. 2017;7. doi:10.1186/s13568-017-0363-8.
  84. Moussa T. A. A.,Mohamed M. S.,Samak N. Production and characterization of di-rhamnolipid produced by Pseudomonas aeruginosa TMN. Brazilian J. Chem. Eng. 2014;31:867–880. doi:10.1590/0104-6632.20140314s00002473.
  85. Déziel E., Lépine F.,Dennie D., Boismenu D. Liquid chromatography/mass spectrometry analysis of mixtures of rhamnolipids produced by{&}lt; i{&}gt; Pseudomonas aeruginosa{&}lt;/i{&}gt; strain 57RP grown on mannitol or {&}hellip; Biophys. Acta BBA. 1999;1440:1–9. doi:10.1016/S1388-1981(99)00129-8.
  86. Tummuscheit M.,Schacht H.,Schacht W. K. M. T. H. Ecological washing of textiles with microbial surfactants. 1996, 125–130.
  87. Luna, J. M.; Santos Filho, A. S.; Rufino, R. D.; Sarubbo, L. A. Production of Biosurfactant from Candida bombicola URM 3718 for Environmental Applications. Chem. Eng. Trans. 2016, 49, 583–588, doi:10.3303/CET1649098.
  88. Zhang, Y.; Miller, R. M. Effect of a Pseudomonas rhamnolipid biosurfactant on cell hydrophobicity and biodegradation of octadecane. Appl. Environ. Microbiol. 1994, 60, 2101–2106.
  89. Zhang, Y.; Miller, R. M. Effect of rhamnolipid (biosurfactant) structure on solubilization and biodegradation of n-alkanes. Appl. Environ. Microbiol. 1995, 61, 2247–2251.
  90. Maslin, P.; Maier, R. M. Rhamnolipid-enhanced mineralization of phenanthrene in organic-metal co-contaminated soils. Bioremediat. J. 2000, 4, 295–308, doi:10.1080/10889860091114266.
  91. Zhang, Y.; Miller, R. M. Enhanced octadecane dispersion and biodegradation by a Pseudomonas rhamnolipid surfactant (biosurfactant). Appl. Environ. Microbiol. 1992, 58, 3276–3282.
  92. Sandrin, T. R.; Chech, A. M.; Maier, R. M. A rhamnolipid biosurfactant reduces cadmium toxicity during naphthalene biodegradation. Appl. Environ. Microbiol. 2000, 66, 4585–4588, doi:10.1128/AEM.66.10.4585-4588.2000.
  93. Dahrazma, B.; Mulligan, C. N. Investigation of the removal of heavy metals from sediments using rhamnolipid in a continuous flow configuration. Chemosphere 2007, 69, 705–711, doi:10.1016/j.chemosphere.2007.05.037.
  94. Evans, T.C., Gavrilovich, E., Mihai, R.C. and Isbasescu, I., E. L.; Thelen, D.; Martin, J. A.; Allen, S. M.; SA, S. ( 12 ) Patent Application Publication ( 10 ) Pub . No .: US 2006 / 0222585 A1 Figure 1. 2015, 002, 354, doi:10.1037/t24245-000.
  95. Kosaric, N. Biosurfactants and their applications for soil bioremediation. Food Technol. Biotechnol. 2001, 39, 295–304, doi:10.1002/chin.199112362.
  96. Linhardt, R. J.; Bakhit, R.; Daniels, L. Communications to the Editor Microbially Produced Rhamnolipid as a. 1989, 33, 365–368.
  97. Haba, E.; Espuny, M. J.; Busquets, M.; Manresa, A. Screening and production of rhamnolipids by Pseudomonas aeruginosa 47T2 NCIB 40044 from waste frying oils. J. Appl. Microbiol. 2000, 88, 379–387, doi:10.1046/j.1365-2672.2000.00961.x.
  98. Rikalović, M. G.; Vrvić, M. M.; Karadžić, I. M. Rhamnolipid biosurfactant from Pseudomonas aeruginosa – From discovery to application in contemporary technology. J. Serbian Chem. Soc. 2015, 80, 279–304, doi:10.2298/JSC140627096R.
  99. Sinumvayo, J. P.; Ishimwe, N. Agriculture and Food Applications of Rhamnolipids and its Production by Pseudomonas Aeruginosa. J. Chem. Eng. Process Technol. 2015, 06, 2–9, doi:10.4172/2157-7048.1000223.
  100. Bunster, L.; Fokkema, N. J.; Schippers, B. Effect of surface-active Pseudomonas spp. on leaf wettability. Appl. Environ. Microbiol. 1989, 55, 1340–5.
  101. Leipzig, K.; Biotechnologie, H. Antiphytovirale Aktivitat yon Rhamnolipid aus Pseudomona,s aeruginosa. 1987, 7, 353–356.
  102. Stanghellini, M. E.; Miller, R. M. Their Identity and potencial Efficacy in the Biological Control of Zoosporic Plant Pathogens. Plant Dis. 1997, 81, 4–12.
  103. Sachdev, D. P.; Cameotra, S. S. Biosurfactants in agriculture. Appl. Microbiol. Biotechnol. 2013, 97, 1005–1016, doi:10.1007/s00253-012-4641-8.
  104. Hashim, M. A.; Mukhopadhyay, S.; Sahu, J. N.; Sengupta, B. Remediation technologies for heavy metal contaminated groundwater. J. Environ. Manage. 2011, 92, 2355–2388, doi:10.1016/j.jenvman.2011.06.009.
  105. Urum, K.; Grigson, S.; Pekdemir, T.; McMenamy, S. A comparison of the efficiency of different surfactants for removal of crude oil from contaminated soils. Chemosphere 2006, 62, 1403–1410, doi:10.1016/j.chemosphere.2005.05.016.
  106. Lourith, N.; Kanlayavattanakul, M. Natural surfactants used in cosmetics: Glycolipids. Int. J. Cosmet. Sci. 2009, 31, 255–261, doi:10.1111/j.1468-2494.2009.00493.x.
  107. Ortiz, A.; Teruel, J. A.; Espuny, M. J.; Marqués, A.; Manresa, Á.; Aranda, F. J. Effects of dirhamnolipid on the structural properties of phosphatidylcholine membranes. Int. J. Pharm. 2006, 325, 99–107, doi:10.1016/j.ijpharm.2006.06.028.
  108. Araújo, C. K. C. de; Campos, A. de O.; Padilha, C. E. de A.; Sousa Júnior, F. C. de; Nascimento, R. J. A. do; Macedo, G. R. de; Santos, E. S. dos Enhancing enzymatic hydrolysis of coconut husk through Pseudomonas aeruginosa AP 029/GLVIIA rhamnolipid preparation. Bioresour. Technol. 2017, 237, 20–26, doi:10.1016/j.biortech.2017.03.178.
  109. Costa, S. G. V. A. O.; Nitschke, M.; Lépine, F.; Déziel, E.; Contiero, J. Structure, properties and applications of rhamnolipids produced by Pseudomonas aeruginosa L2-1 from cassava wastewater. Process Biochem. 2010, 45, 1511–1516, doi:10.1016/j.procbio.2010.05.033.
  110. Oluwaseun, A. C.; Kola, O. J.; Mishra, P.; Ravinder Singh, J.; Kumar Singh, A.; Singh Cameotra, S.; Oluwasesan Micheal, B. Characterization and optimization of a rhamnolipid from Pseudomonas aeruginosa C1501 with novel biosurfactant activities. Sustain. Chem. Pharm. 2017, 6, 26–36, doi:10.1016/j.scp.2017.07.001.
  111. Mendes, A. N.; Filgueiras, L. A.; Pinto, J. C.; Nele, M. Physicochemical Properties of Rhamnolipid Biosurfactant from Pseudomonas aeruginosa PA1 to Applications in Microemulsions. J. Biomater. Nanobiotechnol. 2015, 06, 64–79, doi:10.4236/jbnb.2015.61007.
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