agronomy

Review
Biosurfactants: Potential and Eco-Friendly Material for
Sustainable Agriculture and Environmental Safety—A Review
Ekambaram Gayathiri 1 , Palanisamy Prakash 2 , Natchimuthu Karmegam 3 , Sunita Varjani 4 ,
Mukesh Kumar Awasthi 5 and Balasubramani Ravindran 6, *

1 Department of Plant Biology and Plant Biotechnology, Guru Nanak College (Autonomous),
Chennai 600042, Tamil Nadu, India; [email protected]
2 Department of Botany, Periyar University, Periyar Palkalai Nagar, Salem 636011, Tamil Nadu, India;
[email protected]
3 Department of Botany, Government Arts College (Autonomous), Salem 636007, Tamil Nadu, India;
[email protected]
4 Gujarat Pollution Control Board, Gandhinagar 382010, Gujarat, India; [email protected]
5 College of Natural Resources and Environment, Northwest A&F University, Taicheng Road 3#,
Yangling, Xianyang 712100, China; [email protected]
6 Department of Environmental Energy and Engineering, Kyonggi University, Youngtong-gu,
Suwon 16227, Gyeonggi-do, Korea
* Correspondence: [email protected]

Abstract: With the present climate change and increasing world population, there is an urgent
need to discover creative, efficient, and cost-effective natural products for the benefit of humanity.
Biosurfactants are produced by various microorganisms that have several distinct properties com-
pared to other synthetic surfactants, including mild production conditions, multifunctionality, higher





 biodegradability, and lower toxicity of living cells synthesis of active compounds. Due to their surface
Citation: Gayathiri, E.; Prakash, P.; tension reducing, emulsion stabilizing, and biodegrading properties of these in place of chemical sur-
Karmegam, N.; Varjani, S.; Awasthi, factants, they are generating huge demand in terms of research and usage. Biosurfactants are widely
M.K.; Ravindran, B. Biosurfactants: used in the food industry as food-formulation ingredients and antiadhesive agents as emulsifiers,
Potential and Eco-Friendly Material de-emulsifiers, spreading agents, foaming agents, and detergents that find application in various
for Sustainable Agriculture and
fields such as agriculture, industrial sectors, and environmental recreation. Recent research focused
Environmental Safety—A Review.
more on heavy metal bioremediation from compost was achieved using biosurfactants-producing
Agronomy 2022, 12, 662. https://
bacteria, which resulted in an improvement in compost quality. Although a number of studies on
doi.org/10.3390/agronomy12030662
biosurfactants synthesis have been reported, very limited information on its cinematics and the
Academic Editor: Małgorzata consumption of renewable substrates are available. In this review paper, we made an attempt to
Szczepanek critically review biosurfactants, their usage, research related to them, and challenges faced.
Received: 11 February 2022
Accepted: 3 March 2022 Keywords: biosurfactant; microorganisms; bacteria; bioremediation; biodegradation; environment
Published: 9 March 2022

Publisher’s Note: MDPI stays neutral

with regard to jurisdictional claims in
1. Introduction
published maps and institutional affil-
iations. Biosurfactants are valuable, surface active and biologically efficient microbial am-
phiphilic molecules for different industries or processes. Microbes synthesize them and
offer an alternative to chemically prepared conventional surfactants, especially when grow-
ing on water immiscible substrates. These molecules can be widely used in cosmetics,
Copyright: © 2022 by the authors. pharmaceutical and alimentary processes such as emulsifiers, humectants, preserving
Licensee MDPI, Basel, Switzerland. agents and detergents because of their structural diversity (e.g., glycolipids, lipopeptides,
This article is an open access article fatty acids, etc.), their low toxicity and biodegradability. In the fields of bioremediation
distributed under the terms and and waste treatment, they are also ecologically safe. They can be made from different
conditions of the Creative Commons substrates, mainly from renewables, such as vegetable oils, distillery and milk waste; those
Attribution (CC BY) license (https://
are economical, but are not reported in detail. In this review, progress on the use of re-
creativecommons.org/licenses/by/
newable substrates for the production and new applications of biosurfactants is reported.
4.0/).

Agronomy 2022, 12, 662. https://doi.org/10.3390/agronomy12030662 https://www.mdpi.com/journal/agronomy

Agronomy 2022, 12, 662 2 of 35

Living cells synthesize surface-active compounds known a biosurfactants, which are pro-
duced by various microorganisms that have several distinct properties compared to other
synthetic surfactants, including mild production conditions, multifunctionality, higher
biodegradability, and lower toxicity. These compounds are primarily biosynthesized as
secondary metabolites and play important roles in the growth and localization of their
microorganisms.
Based on the chemical structure of their hydrophobic component, BSs are classified
into four types: (1) glycolipid type, (2) fatty acid type, (3) lipopeptide type, and (4) polymer
type. Due to their surface tension reducing, emulsion stabilizing, foam promoting, and
biodegrading properties, use of these in place of chemical surfactants is a highly demanded
area of interest. Thus, in hydrocarbon-contaminated sites, biosurfactants producing mi-
croorganisms accelerate bioremediation. Bacteria and yeasts synthesize most of these.
Glycolipid, phospholipid, rhamnolipid, etc., are the biochemicals with great surface ac-
tivity synthesized by these organisms. The hydrocarbon substrate is emulsified by the
production of these chemicals for the facilitation of transportation into cells. A mechanism
known as swarming motility is identified in the biosurfactants mechanism of action. The
structure of a biosurfactant is composed of a hydrophobic tail and a hydrophilic head [1],
otherwise composed of amphophilic or hydrophobic peptides. Hydrocarbon uptake is
highly related to the spontaneous release and function of biosurfactants. Thus, the max-
imum production is seen in the hydrocarbon-degrading microorganisms. Water-soluble
compounds like glucose, etc., also seemed to produce biosurfactants in rare cases [2,3].
These compounds also have antibiotic properties that disrupt the membrane of the food
competitive microorganisms. These biosurfactants show improved properties compared to
a chemically synthesized surfactant, which enables them to be used in the process of oil
recovery in an environmentally safer way. Their tolerance in extreme conditions, ease of
culturing, high-scale production, eco-friendly nature, and diversified nature makes them
efficient enough in the implementation of various fields, including microbial degradation.
Antiadhesive agents and food-formulation components had long been used in the food
sector. Biosurfactants stabilize emulsions by reducing surface and interfacial tension. Other
activities of biosurfactants in food processing include controlling fat globule agglomeration,
stabilizing aerated systems, enhancing the texture and shelf life of starch-containing goods,
and improving the consistency and texture of fat-based products. Biosurfactants are used
to manage consistency, prolong freshness, and solubilize flavor oils in bread and ice cream
manufacture. They’re also utilized in oil and fat frying as fat stabilizers and anti-spattering
agents. The addition of rhamnolipid biosurfactant to bakery goods increases dough sta-
bility, texture, volume, and preservation, as well as butter cream, croissants, and frozen
confectionary items’ qualities. In light of changing climate circumstances and a growing
global population, it is critical to investigate creative, efficient, and cost-effective natural
products for the betterment of people [4–6].
Biosurfactants have seen a tremendous increase in research and development, as well
as commercialization of biological agents in recent years. Microorganisms create a wide
variety of amphiphilic metabolites, many of which are unique in their structure. Various
ways for categorizing microbial biosurfactants, such as structural similarities, diameters,
moieties, hydrophobicities, degree of change, and other physical and chemical parameters,
are applied in addition to traditional surfactant classification methods. Aside from reducing
surface stress, microbial surfactants may provide a number of other advantages. This
combined effect of bioactivity and interfacial activity, which is highly dependent on the
structure and composition of each molecule, provides the vast majority of biological
structures with unique opportunities for pharmaceutical, agricultural, environmental, and
other applications that have yet to be discovered. Apart from the well-known glycolipids
and lipopeptides, there are several more classes, structures, and structural combinations of
microbial biosurfactants.
Agronomy 2022, 12, 662 3 of 35

Objectives:
1. Biosurfactants have emerged as potential molecules for drug delivery vehicles, medic-
inal applications, agricultural applications, and environmental safety, all of which
provide economically appealing and scientifically novel applications.
2. The current study discusses biosurfactants and their production by bacteria, with an
emphasis on their involvement in oil cleanup.
3. To explore novel biosurfactants those are commonly used for soil remediation.
4. To identify the notable biomolecules that potentially replace harsh surfactants now
employed in pesticide manufacturing.
5. To determining the significance of environmental biosurfactants in plant growth
promotion and other agricultural uses requires detailed investigation.
6. Emphasize the use of biosurfactants as eco-friendly and alternatives to synthetic
surfactants.
This review offers an overview of microbial biosurfactants’ diversity and classification
by presenting both well-known and well-investigated examples, and covers the present
reality of biosurfactant research, pointing the way toward the discovery and development
of molecules with innovative structures and different functionalities for modern techniques.

2. Classification of Biosurfactants
Surfactants are one of the most diverse chemical groups used in different industrial
processes. They have a competitive market, and producers will have to expand the pro-
duction of surfactants in an environmentally friendly way. Incentives for environmentally
friendly and cost-efficient biosurfactants have led to an increased interest in biological
agents. The structural variety and functional features of biosurfactants make them an
enticing compound class that may be employed for a broad range of industrial, environ-
mental, and biotechnological applications. Screening methods make it simpler to find
prospective bacteria that produce biosurfactants. A variety of purifying and analytical
approaches are available for the characterization of biosurfactants. Biosurfactants are
classified largely based on their chemical composition and source of generation. Based on
their molecular mass, another classification with two major classes was also suggested [7].
Glycolipids and lipopeptides constitute the low molecular biosurfactants, whereas lipopro-
teins, lipopolysaccharides, and amphipathic polysaccharides constitute the high molecular
mass biosurfactants. The low molecular biosurfactants lower the surface and interfacial ten-
sions, whereas the other is highly efficient in stabilizing emulsions [8]. The other classes of
biosurfactants include phospholipids, polymeric surfactants, and particulate surfactants [9].
The detailed list of classification is given in Table 1.

Table 1. Types of biosurfactants—according to the structure.

Surface-Active Fatty Acids/ Particulate

Glycolipids Lipo–Peptides Polymeric Surfactants
Antibiotics Neutral Lipids Biosurfactants

(i) Rhamnolipids (i) Surfactin/Iturin/

(ii) Trehalose lipids Fengycin (i) Emulsan
(i) Gramicidin (i) Vesicles
(iii) Sophorolipids (ii) Viscosin (i) Corynomicolic (ii) Alasan
(ii) Polymixin (ii) Whole Microbial
(iv) Manno- (iii) Lichenysin Acids (iii) Lipan
(iii) Antibiotics TA Cells
sylerythritol (iv) Serrawettin (iv) Lipomanan
lipids (v) Phospholipids

2.1. Glycolipid
Long-chain aliphatic acids combined with carbohydrates make up the glycolipids
orhydroxy aliphaticacids. Rhamnolipids, sophorolipids, trehalolipids, and fructose-lipids
are some of the glycolipids β-hydroxy fatty acids attached with different sugars are found in
glycolipids whereas cycloheptapeptides with amino acids linked to fatty acids of different
chain lengths are found in lipopeptides. Their solubility is seen in both polar and non-polar
solvents [10–12].
Agronomy 2022, 12, 662 4 of 35

2.1.1. Rhamnolipid
One or two molecules of rhamnose attached to one or two molecules of b-hydroxy
decanoic acid make up the rhamnolipid. Thus far, seven homologs of rhamnolipids
have been reported [13]. P. aeruginosa produces the following prominent substances:
L-Rhamnosyl-L-rhamnosyl-b-hydroxy decanoyl-b-hydroxy decanoate and L-rhamnosyl-
b-hydroxy decanoyl-b-hydroxy decanoate, which were referred to as rhamnolipid 1 and
2, respectively [9]. Against n-hexadecane the surface tension and the interfacial tension
have been reduced to 30 mN/m and 1 mN/m, respectively, by the rhamnolipids from
P. aeruginosa [14]. The rhamnolipids synthesized from Pseudomonas aeruginosa L2-1 showed
100% emulsification against soybean oil, and 69% of crude oil [15]. Rhamnolipid also
showed significant antimicrobial activities against several microbes, thus playing a cru-
cial role in the field of pharmaceuticals [16]. In few cases, heterogeneous mixtures of
rhamnolipids are isolated from Pseudomonas sp. [17], whereas rhamnolipids produced
from Marinobacter sp. have prospects for industrial applications [18]. Rhamnolipids have
antimicrobial potential against L. monocytogenes [19]. Rhamnolipids in combinations of
enzymes for improved cleaning wherein the weight percent of the rhamnolipid made up
by mono-rhamnolipids can be a good detergent composition [20]. Rhamnolipids can be
produced using sugars as carbon sources by P. aeruginosa [21]. DYNA270 produced by
Pseudomonas sp. showed to be a promising factor in displacing heavy oily sludges from
polypropylene coupons [22]. The desorption efficiency of phenanthrene is found to be
increased by rhamnolipid, thus proving their application in the remediation of PAH con-
taminated soils in cold region [23]. The structures of rhamnolipid are given in Figure 1A–H.

2.1.2. Sophorolipids
Extracellular sophorolipids consisting of a mixture of different hydrophobic sophoro-
sides constitute the sophorolipids. Surface-active glycolipids like sophorolipids are made
up of a disaccharide sophorose unit glycosidically attached to hydroxylated fatty acid
surface-active glycolipids [24]. They exist naturally in both open (acidic sophorolipids)
and closed (lactonic sophorolipids) types. Furthermore, novel sophorolipids, sophoro-
sides (SSs), and glucosides with promising metal coordination properties, have been
formed [25], including novel bolaform sophorolipids and sophorosides [26,27]. The sys-
temic and structural disparities among the sophorolipid substances have an impact on
their physicochemical activities. Lactonic acetylated forms, for example, have the greatest
biological activity, while acidic forms provide greater foam-forming abilities and water
solubility [28]. Sophorolipids are manufactured across a number of companies in the United
States, Europe, and Asia. It is used in sustainable safe cleaning goods and cosmetics [29].
This market success is mostly due to their high productivity. A fed-batch fermentation
method will produce over 300–400 g/L of liquid [30].
Certain species of Torulopsis has been reported to produce dimeric carbohydrate
sophorose linked to 1,2 long chain hydrocarboxylic acids [8]. Though the emulsifying
properties were not seen, surface and interfacial tension, and reduction capability has
been reported [31,32]. The surface tension and the interfacial tension against n-hexadecane
and water have been reduced to 33 mN/m and 5 mN/m by both lactonic and acidic
sophorolipids with 10 mg/L of pure sophorolipid. Though the temperature and pH are
changed, notable stability is seen. In the pH values of 6–9, and at temperatures 20–90 ◦ C
stable surface-active properties have been noted. Antibiotic resistance problem was tried to
address the antimicrobial properties of Sophorolipids (SLs). Staphylococcus aureus growth
was completely inhibited by the SL-tetracycline combination [33]. The growth of organisms
like Bacillus subtilis, Staphylococcus xylosus, Streptococcus mutans, and Propionibacterium acne
were completely inhibited by the sophorolipids produced by Candida bombicola ATCC
22214 [34,35]. Biofilm formation was also being disturbed by these sophorolipids [36].
Acidic sophorolipids are reported have higher antimicrobial properties against the noso-
comial infective agents thus contributing a major part in the production of antimicrobial
cream [37]. Sophorolipid produced by Rhodotorula babjevae YS3 shows a greater antifungal
Agronomy 2022, 12, 662 5 of 35

activity when compared to others [38]. Bolaform sophorolipids may be an intriguing

biological option, with potentially different characteristics. However, apart from certain
“high-end” uses, the existing demand for industrially manufactured SLs is controlled by its
usage in detergent applications by more than 90% [39]. The structure of sophorolipid is
given in Figure 2A–D.

Figure 1. Cont.
Agronomy 2022, 12, 662 6 of 35

Figure 1. (A–H) The structures of rhamnolipid.

Agronomy 2022, 12, 662 7 of 35

Figure 2. (A–D) The structure of sophorolipid.

2.2. Lipoproteins and Lipopeptides

Cyclic lipopeptides with 8–17 amino acids and a lipid portion constitutes the lipopeptides.
They show heavy variation in their amino acid and lipid composition [9]. Streptomyces sp.
DPUA1566 has potential applications either in bioremediation processes or in pharmaceutical
and cosmetic industries, and it is from a low-cost waste source [40]. Lipoprotein synthesized
from Pseudomonas gessardii shows prominent metal ion removing properties [41], similarly a
Agronomy 2022, 12, 662 8 of 35

lipopeptide produced by Bacillus atrophaeus 5-2a showed high efficiency in crude oil removal [42].
The anthracnose disease caused by Colletotrichum gloeosporioides has been greatly inhibited by
the lipoprotein produced by Bacillus subtilis CMB32 [43]. The structure of lipopeptides is given
in Figure 3A,B.

Figure 3. (A,B) The structure of lipopeptides.

Surfactin
Cyclic acidic lipopeptides constitute the surfactin. Out of all known biosurfactants, the
surfactin produced by B. subtilis, is the most effective one. [44,45] Surfactin is attributed with
an important property of red blood lysing. Additionally, they act as good antibiotic [44,46].
A hydrophobic ball-like structure of surfactin is formed in water and air [47,48]. Surfactin
is produced from Bacillus subtilis ATCC21, 332 is used in the enhancement of the iron-
remediation [49,50]. Anti-inflammatory activity is also reported from these surfactin. It
also inhibits the expression of IFN-γ, IL-6, iNOS, and nitric oxide (NO) [51]. Cancer can
be inhibited by the application of surfactin. Anticancer therapy can be improved by the
optimization of nano-particle delivery system of surfactin [52–54]. The surfactin isolated
from Bacillus subtilis HSO121 contribute maximum in daily life and industrial applications
due to their diversified and effective characteristic features [55]. The structure of surfactin
is given in Figure 4A–F and the types of surfactin is given in Figure 4G.
Agronomy 2022, 12, x FOR PEER REVIEW 10 of 38

Agronomy 2022, 12, 662 9 of 35

(A) Linear surfactin (B) Surfactin peptide

(C) Surfactin-C13 (D) Surfactin A

(E) Surfactin B (F) Surfactin-C13

Figure 4. Cont.
Agronomy
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(G)structure
Figure 4. (A–G) The The types of Surfactant
of surfactin; and (G) the types of surfactant.
Figure 4. (A–G) The structure of surfactin; and (G) the types of surfactant.
2.3. Fatty Acids, Phospholipids, and Neutral Lipids
Several
2.3. Fatty microorganisms
Acids, Phospholipids, and such as bacteria
Neutral Lipids and yeast produce fatty acid and phos-
pholipid, which have received increasing demand in the current situation due to their
Several microorganisms such as bacteria and yeast produce fatty acid and phospho-
highly diversified and effective properties as biosurfactants [8]. Microemulsions of alka-
lipid, which have received increasing demand in the current situation due to their highly
nes were produced by these biosurfactants. [56] Lipoamino acid is produced by pro-
diversified and effective properties as biosurfactants [8]. Microemulsions of alkanes were
teinogenic or non-proteinogenic amino acids. [57,58] Sulfur-reducing bacteria such as
produced by these biosurfactants. [56] Lipoamino acid is produced by proteinogenic or
Thiobacillus thiooxidans produces this biosurfactant in higher amounts [8,9,59,60]. Phos-
non-proteinogenic amino acids. [57,58] Sulfur-reducing bacteria such as Thiobacillus thi-
phatidylethanolamine produced by Rhodococcus erythropolis reported to show a lowered
ooxidans produces this biosurfactant in higher amounts [8,9,59,60]. Phosphatidylethanola-
interfacial tension against hexadecane to less than 1 mN/m and a CMC of 30 mg/L [8,61].
mine produced by Rhodococcus erythropolis reported to show a lowered interfacial tension
When grown on n-alkanes. Acinetobacter spp. produces phosphatidyl ethanolamine-rich
against hexadecane to less than 1 mN/m and a CMC of 30 mg/L [8,61]. When grown on n-
vesicles and forms medically essential micro emulsions. According to Gautamand Tyagi,
alkanes.
the majorAcinetobacter spp. produces phosphatidyl
cause of respiratory-related ethanolamine-rich
problems is phospholipid protein vesicles
complexand forms
deficiency.
medically essential micro emulsions. According to Gautamand Tyagi, the
The gene from selected bacteria and yeast can be isolated and cloned, and by use of fer- major cause of
respiratory-related problems is phospholipid protein complex deficiency.
mentative processes, surfactants can be produced. Biosurfactants produced by P. putida The gene from
selected
inhibitedbacteria and yeast
the growth can be isolated
of C. albicans [62,63]. and cloned,
Neutral andexpected
lipids by use of tofermentative
be wax ester-likepro-
cesses, surfactants can be produced. Biosurfactants produced by P. putida
lipids produced by the marine hydrocarbonoclastic bacteria. Trehalose lipids synthesized inhibited the
growth of C. albicans
from Rhodococcus [62,63].
fascians BD8Neutral
makes alipids expected
greater to be wax
contribution in theester-like lipidsdue
medical field produced
to their
by the marine hydrocarbonoclastic bacteria. Trehalose lipids synthesized
antimicrobial and other properties [64]. Biosurfactants produced by Lactobacillus pentosus from Rhodococ-
cus
(PEB)fascians
with BD8 makes a greater
antimicrobial contribution
and anti- adhesive in the medical
activities fieldgreater
showed due to inhibition
their antimicro-
effect
bial and other properties [64]. Biosurfactants produced by Lactobacillus
towards the adverse effects and growth of the skin microflora, thus finding a greater pentosus (PEB) place
with
antimicrobial
in the cosmetic industry [65]. The structures of fatty acids are given in Figure 5A–D. the
and anti- adhesive activities showed greater inhibition effect towards
adverse effects and growth of the skin microflora, thus finding a greater place in the cos-
2.4. Polymeric
metic industryBiosurfactants
[65]. The structures of fatty acids are given in Figure 5A–D.
A complex mixture of biopolymers such as proteins, polysaccharides, lipopolysaccha-
rides were the base components of the exocellular polymeric surfactants obtained from
many bacterial species of different genera [8]. Through o-ester linkages the polysaccharides
are covalently linked to fatty acids [9,66]. Emulsan, liposan, and mannoprotein are the best
examples of polymeric biosurfactants [9,67,68].

2.4.1. Emulsan
Pure form of emulsan shows emulsifying activities under low concentrations. Emulsan
produced by Acinetobacter calcoaceticus PTCC1318 showed crude oil degradation proper-
ties [69]. By coating the hydrophobic substrate, they are made readily available for microbial
(A) (1S,2S)-2-(anthracene-2,3-dicarboximido)cyclohex-
access. Mixtures of aliphatic and aromatic(B)
anecarboxylicacid
16-methylheptadecanoic
hydrocarbons acid efficiently in
can be emulsified
balanced proportions except their pure forms. Fatty acid components determine the emul-
sifying activities of emulsan [70,71]. Acinetobacter calcoaceticus BD4 emulsan showed the
optimal emulsification activity when polysaccharides and proteins were mixed and when
they are separate no activity is reported [72].
Agronomy 2022, 12, 662 11 of 35

Figure 5. (A–D) shows the structures of fatty acids.

2.4.2. Liposan
This emulsifier is synthesized by C. lipolytica consisting of carbohydrates and proteins.
It finds a prominent place in the food and cosmetic industries [73]. It forms stable emulsions
with edible oils [74].

2.4.3. Mannoprotein
The major component of the cell wall of Saccharomyces cerevisiae, the mannoprotein
is an effective bioemulsifier [75]. They were capable of forming stable emulsions with
many hydrocarbons and other substances, thus suggesting their applications as cleaning
agents. The structural and enzymatic classification of mannoproteins is dependent on
its chemical composition and specific function. The most abundant ones are the struc-
tural mannoproteins, which are made up of mannopyranosyl attached to a small protein,
but the enzymatic mannoproteins are the most effective emulsifiers and have more pro-
tein moieties. They activate the immune cells for the production of antibodies [76,77].
Kluyveromyces marxianus produced a mannoprotein that formed a stable emulsion in corn
oil [78]. Mannoprotein bioemulsifier isolated from Saccharomyces cerevisiae 2031 showed
similarity with the mannoprotein obtained from Acinetobacter sp. in 77% carbohydrate and
23% protein [79] 53% protein, 42% polysaccharide, and 2% lipid [80].

2.5. Particulate Biosurfactant

The vesicles and fimbriae synthesized by Acinetobacter sp. are the constituents of
this surfactant. Protein, phospholipid, and lipopolysaccharide are the components of
the purified vesicles. Microemulsion has been formed by these [9,81]. Hydrophilic and
hydrophobic moieties of biosurfactants enable them in the accumulation in different phases
of interfaces [82]. The biosurfactant consists of a simple polar head made up of simple
or a mixture of compounds like amino acids, peptides, etc., whereas the nonpolar tail is
made of fatty acids and rarely cyclic structures were also reported [8,83,84]. Spherical
biosurfactant vesicles were formed at concentrations greater than their critical micelle
concentration (CMC) [85,86]. The attachment of biofilm formers have been effectively
Agronomy 2022, 12, 662 12 of 35

reduced by biosurfactant. The adherence of L. monocytogenes is strongly impeded by RHL.

This shows the inhibition of bacterial populations during biofilm formation [87].

3. Biosurfactant Producing Mechanisms

Surfactants are a structurally varied and heterogeneous class of microorganism-
produced surface-active chemicals. Surfactants are synthesized in both the synthetic
and biological realms [88]. Biosurfactants (BSs) are surfactants obtained from biological
entities, particularly bacteria and fungi. These compounds are synthesized as metabolic
products or as a result of the surface chemistry of the cells. BSs are mostly created by
aerophilic microorganisms in aqueous media using carbon source feedstocks such as hy-
drocarbon, polysaccharides, lipids, and oil derived from bacteria (Pseudomonas, Bacillus,
and Acinetobacter), fungus (Aspergillus and Fusarium), and yeast (Candida and Pseudozyma).
Rhamnolipids, surfactins, sophorolipids, emulsans, and mannosylerythritol lipids are the
most often seen BSs [89]. These surface-active chemicals serve a physiologic purpose by
allowing BSs generating microbes to grow on insoluble substrates, assure exponential
biomass growth, demonstrate antimicrobial activity against potential predators, and enable
them to withstand harsh environmental conditions, pathogenicity, and cell desorption. The
mechanisms of biosurfactants vary according to the class to which they belong [88].

Mechanism of Action of Surfactants

When a decent number of surfactant molecules are accumulated in a solution, they
mix to create micelles. As the micelle develops, the surfactant heads align themselves to
be exposed to water, while the tails cluster together in the micelle’s core, sheltered from
water. Micelles cooperate together to reduce contaminants. The hydrophobic tails cling to
and surround dirt, while the hydrophilic heads lift the clinging soils off the surface and
into the cleaning solution. The micelles then reform, with the tails hanging the dirt in the
structure’s core [90,91].

4. Isolation of Biosurfactant
Microbes nearly usually exist in mixed populations in natural habitats, comprising a
diverse range of strains and species. A pure culture is necessary to analyze the qualities
of a specified organism isolated from such a mixed population. Apart from direct strain
separation by dilution and plating, enrichment cultures using hydrophobic surfaces are
very promising for the isolation of biosurfactant producing microorganisms [92]. Minimal
salt medium is a widely used media to isolate microbes from hydrocarbon contaminated
regions. Enrichment procedures were used to extract 130 oil-degrading isolates from
hydrocarbon-polluted sites. The researchers used a mineral salts medium with crude oil as
the only carbon source [93].
To get crude biosurfactant, cell-free supernatant was collected by centrifuging culture
broth for 20 min at 10,000 rpm and 4◦ C. Then, 6 N HCl was added to the clear supernatant
to adjust the pH to 2 [94]. Additionally, hydrophobic interaction chromatography and the
replica plate technique are successful approaches [92]. To generalize, surveying polluted
locations in combination with direct isolation or enrichment culture is a proven technique
for finding novel biosurfactant producing bacteria. However, since the percentage of
positives is just a few percent, many hundred isolates must be screened for each hit [94].
Isolation, screening and characterization of biosurfactants from natural habitat is shown in
Figure 6.
Agronomy 2022, 12, 662 13 of 35

Figure 6. Isolation, screening and characterization of biosurfactants.

5. Biosurfactant Produced by Microorganisms

Various microorganisms contribute to the production of various biosurfactants. The
detailed list is given in Table 2.

Table 2. Various biosurfactants produced by microorganisms.

Microorganism Biosurfactant Property Technology/Application References

A. calcoaceticus RAG-1 A very good bioemulsifier, which
Heteropolysaccharides Biostimulation [95]
(Arthrobacter RAG) makes heavy crude oil less viscous.
Lower the interfacial tension and
Arthrobacter. sp., Trehalose, sucrose, Biodegradation and
make hydrophobic compounds [96]
Rhodococcus aurantiacus and fructose, lipids Bioaugmentation
more “pseudosoluble.”
Mycobacterium sp., Used more in cosmetic industries
Trehalose dimycolates Bioaugmentation [97]
Nocardia sp. because of its lower irritability.
Arthrobacter MIS 38
Low interphase surface tension due Bioaugmentation and
Bacillus atrophaeus 5-2a Lipopeptide [51]
to emulsifying action. biostimulation
Pseudomonas fluorescence
Used in enhancement of the
Bacillus subtilis ATCC Bioaugmentation and
Surfactin iron-remediation, [49,50]
21332 biodegradation
anti-inflammatory activity.
Pseudomonas aeruginosa Playing a crucial role in the field of
Rhamnolipids Biodegradation [16]
L2-1, Bacillus sp. AB-2 pharmaceuticals.
Substantial % E24 against diverse
hydrocarbons, including light and
Candida bombicola ATCC
Sophorose lipids heavy crude oils, and high stability Bioaugmentation [98]
22214
under salinity, pH,
and extreme heat.
Recognized as key
Candida tropicalis Mannan-fatty acid Biostimulation [99]
antigenic determinants.
Candida lipolytica Y-917, Produces hydrocarbon and oil
Sophoros lipid Biostimulation [100]
Torulopsis bombicola emulsions in a liquid like water.
Agronomy 2022, 12, 662 14 of 35

Table 2. Cont.

Microorganism Biosurfactant Property Technology/Application References

Have novel organic pollutant
catabolism pathways and potential
Clostridium pasteurianum;
Neutral lipids soil bioremediation capabilities for Biodegradation [101]
Nocardia erythropolis
hydrocarbons and
aromatic chemicals.
Corynebacterium
hydrocarbolastus, Fluids viscosified with viscoelastic
Protein-lipid
Corynebacterium lepus surfactants (VESs) used in Biodegradation [102]
-carbohydrate
Strain MM1, Phaffta hydrocarbon recovery procedures.
rhodozyma
Myroides sp., Pseudomonas
sp., Thibacillus sp., A potential biorestoration technique Bioaccumulation and
Ornithine lipids [103]
Agrobacterium sp., for contaminated sediments. biodegradation
Cluconobacter sp.
Corynebacterium
Phospholipids Multiantibiotic resistant. Biostimulation [104]
insidiosum
Reduces the viscosity of heavy oil,
Ochrobactrum
cleans oil storage tanks, and
anthropiHM-1 and Proteins Biostimulation [104]
increases the flow of oil
Citrobacter freundiiHM-2
through pipelines.
Used as a bioactive compound to
Biostimulation and
Penicillium spiculisporum Spiculosporic acid remove heavy metal cations [105]
bioaugmentation
from water.
Trehalose- A potential biorestoration technique Bioremediation and
Rhodococcus erythropolis [106]
dicarynomycolate for contaminated sediments. biotransformation
Responsiveness to hazardous and
refractory chemicals such
Rhodococcus sp. ST-5
chlorinated aliphatic and aromatic Bioremediation and
Rhodococcus sp. H13-A Glycolipid [106]
hydrocarbons, N- and S-heterocyclic biotransformation
Rhodococcus sp. 33
compounds, and
synthetic polymers.

6. Properties of Biosurfactants
Biosurfactants such as rhamnolipid and greenzyme were attributed with a special
property of reducing the surface tension of water and interfacial tension of n-hexadecane
considerably [9,107,108]. Temperature stability, critical micelle concentration (CMC), and
low interfacial tensions were the effective physicochemical properties of these biosur-
factants. These properties enable the formation of microemulsions, which help in the
solubilization of hydrocarbons in water [83] (Figure 7). Thus, micelle formation is one of
the most important properties of a surfactant. With an increase in the surfactant concentra-
tions, there is a gradual decrease in the surface tension [109]. At concentrations above CMC,
surfactants such as rhamnolipids stabilize the micelle formation [110]. As a substitute
for synthetic biosurfactants, the synergistic impact of antioxidants has led to the use of
several biosurfactant extracts in the cosmetic sector. [111]. When different concentrations of
biosurfactants were applied to dyed hair, the results revealed that the adsorption of dyed
hair is high above the CMC of the biosurfactant, though maintaining the hair in a good
state. This reveals their best application in the cosmetic industry [112]. A biosurfactant
improves water loss, thus wetting the solid surfaces [107]. It is reported that many emulsi-
fiers were attributed with only minimal surface tension reducing property [107]. Based on
the concentration of the biosurfactant the clear zone varies [113]. Figure 8 shows types of
biosurfactant used in various industries.
Agronomy 2022, 12, 662 15 of 35

6.1. Surface and Interface Activity

Surfactin from B. subtilis, rhamnolipids from P. aeruginosa, and sophorolipids from
T. bombicola were reported with high surface tension reduction and interfacial tension reduc-
tion properties of water and n-hexadecane, respectively. Comparatively, the biosurfactants
are 10–40 times more efficient than the synthetic surfactants. The biosurfactant produced
by the bacteria Bacillus mojavensis JF-2 was reported to reduce the interfacial tension of
crude oil while lowering the surfactant concentrations [114].

Figure 7. Properties of biosurfactant.

6.2. Temperature, PH, Ionic Strength Tolerance

Many biosurfactants and their surface activities remain unaffected by the environmen-
tal conditions like pH and temperature. The biosurfactant lichenysin from B. licheniformis
JF-2 remains unaffected at temperatures up to 50 ◦ C with pH range of 4.5 and 9.0. Similarly,
a lipopeptide synthesized by B. subtilis LB5a sustained in a high temperature of 121 ◦ C and
at high salt conditions even for 6 months without losing its surface activities. At pH 4–10,
high temperatures up to 120 ◦ C, and a NaCl content of up to 10% (w/v), the biosurfactant
maintains emulsification activity [115]. Even after high temperature treatment surfactants
like surfactin and rhamnolipids showed anti-adhesive properties to various microorganisms
such as Staphylococcus aureus, etc., thus protecting the surface from these pathogens [116].
Similar effects were produced by the biosurfactant synthesized by C. lipolytica UCP0988
but under extreme pH conditions. [117,118] The biosurfactants produced by B. licheniformis
STK01, B. subtilis STK02, and P. aeruginosa STK03 showed emulsifying properties even after
treating at temperatures up to 50 ◦ C [119]. The surface activity and emulsification capacity
is reported to be stable even at pH of 7–8 for the biosurfactant produced by Bacillus subtilis
strain JA-1. The biosurfactant produced by Pseudomonas aeruginosa RS29 showed stable
temperature stability, pH and saline stability. It was also attributed with high foaming and
emulsifying activities even after these extreme conditions [120]. A similar condition has
Agronomy 2022, 12, 662 16 of 35

been reported from the surfactant produced by Virgibacillus salaries, thus finding its effi-
cient application in various industries, that to particularly in marine bioremediation [121].
Biosurfactant synthesized by Streptomyces sp. DPUA 1566 also showed stability with slight
variations when exposed to various temperatures and pH [40]. The same conditions were
also reported from the surfactants of B. subtilis and B. tequilensis.

Figure 8. Types of biosurfactant used in various industries.

6.3. Biodegradability and Low Toxicity

Biosurfactants were the most appropriate tool for bioremediation without producing
any harmful effects to the environment. They are highly eco-friendly and safer than
the synthetic surfactants. Only a very few data have reported the harmful effects of
biosurfactants. Rhamnolipids have been reported with a lethal concentration (LC) of
50% against Photobacterium phosphoreum, which is comparatively 10 times higher than
that of a synthetic surfactant. On analyzing the toxicity of biosurfactants it has been
reported that synthetic surfactant sucrose-stearate is similar in homology to biologically
synthesized glycolipid, but it degraded faster than that of this biologically synthesized one.
A comparative study between Marlon A-350, a synthetic surfactant and a biosurfactant
synthesized by P. aeruginosa revealed that the synthetic surfactant was highly toxic in all
the assays and its properties, whereas the biosurfactant was non-toxic [122]. An analysis
revealed that the biosurfactant produced by Candida sphaerica reduced the surface tension
when applied over the seeds of Brassica oleracea, Cichorium intybus, and Solanum gilo [102].
Rhamnolipid produced by P. aeruginosa PG1 showed a cytotoxic effect against the cell line
L292, yet serves to be a promising class of biosurfactants with antimicrobial properties,
thus being a very good option to be applied for the bioremediation of soil and crude oil
processes [123]. When the application of surfactin produced by Bacillus subtilis HSO121
Agronomy 2022, 12, 662 17 of 35

was analyzed, it revealed that it is a non-toxic, nonirritant compound, thus a safer one to
be used in the detergent formulations [55].

6.4. Emulsion Forming and Emulsion Breaking

Produced emulsions have a life span of few months to years. Biosurfactants can
either stabilize the emulsion as emulsifiers or de-stabilize the emulsion as de-emulsifiers.
Comparatively, high molecular-mass biosurfactants are better emulsifiers than the low-
molecular ones. Few surfactants might act as a good surface tension reducer, whereas
they might not be good emulsifiers, whereas few others might not be good in reduc-
ing the surface tension, but they were good emulsifiers indeed. Polymeric surfactants
coat the oil, thus making stable emulsions, thereby being applied in various cosmetic
and food industries. When the agro waste Beta vulgaris (Beetroot) was used as a sub-
strate, the bacterial isolates B. licheniformis STK01, B. subtilis STK02, and P. aeruginosa
STK03 produced the biosurfactants, which showed very high emulsification rate for heavy
hydrocarbons such as anthracene, and lubricant oil [119]. For the discovery of novel
microbial emulsifiers, novel screening methods are required, due to the great potential
that these molecules possess for green technology [124]. Few emulsifiers with higher
emulsifying ratio have been reported from the species of Saccharomyces cerevisiae and
Acinetobacter sp. [79,80]. Using the emulsification index E24 test, six isolates were identified
which showed significant emulsification against heavy hydrocarbons without reducing
the interfacial tension [125–127]. It was reported that exopolymers of Bacillus subtilis 28,
Alcaligenes faecalis 212, and Enterobacter sp. 214 showed high emulsification activity [128].
Mannoproteins formed stable emulsions with almost all the compounds such as hydrocar-
bons, oils, etc., thus serves to be a promising class of biosurfactants to be applied in various
industries [129]. Bioemulsifiers also serve to take a better place in the bioremediation of oil
spills, heavy metal removal, food and agro industries, cosmetic industries, and pharmaceu-
tical industries [127,130–133]. Thus, these serve to be better solutions in these industries
rather than the application of toxic synthetic surfactants [134].

7. Application of Biosurfactants
In comparison to chemically synthesized surfactants, biosurfactants offer a number of
advantages (Figure 9).

7.1. Agriculture
Agriculture productivity is a major challenge for all countries as it relates to meeting
the expanding needs of the human population. It is now necessary to use green substances
in order to achieve sustainable agriculture. This review demonstrates the widespread usage
of caustic surfactants in agriculture and the agrochemical industry. Green surfactants may
be biosurfactants generated by bacteria, yeasts, and fungus. Biosurfactants are used in
agriculture to eliminate plant pathogens and increase nutrient bioavailability for beneficial
plant microorganisms. Agricultural soil remediation with biosurfactants may greatly
improve agricultural soil quality. Surfactants are employed in crop protection and pesticide
formulations in an estimated 0.2 million tonnes per year [115].
Good wettability, inhibition of toxicants of pesticides and even distribution of fertiliz-
ers in the soil are achieved by the process of hydrophilization using biosurfactants [135].
Removal of plant pathogens and the advancement of the bioavailability of plant nutri-
ents have been increased considerably upon the application of biosurfactants. In several
ways, growth of rhizobacteria in the rhizosphere is increased, which helps in the plant
growth promotion [136,137]. These biosurfactants improves the quality of soil, and increase
the interaction between the plants and the beneficial microbes [138]. These have more
advantages over the synthetic surfactants, which are currently used in the pesticides, as
these biosurfactants are eco-friendly in nature, economically cheap and also increase the
beneficial microbes in the soil [139–143].
Agronomy 2022, 12, 662 18 of 35

Figure 9. Application of biosurfactant in various fields.

7.2. Biosurfactants in Agriculture and Agrochemicals

A growing population demands more agricultural output in order to fulfil the needs of
the world’s growing population. It is imperative that green technology be used in agricul-
ture to ensure long-term viability. It is possible to utilize biosurfactants as plant-protection
measures in a variety of ways. It has the potential to increase the supply of nutrients for
beneficial microorganisms associated with plants. Soil health may be improved by the
use of biosurfactants, according to a variety of studies. For contemporary agriculture, bio-
surfactants are a necessary component. Biosurfactants derived from Pseudomonas sp. and
Burkholderia sp. bacteria might be used as safe biopesticides. Cationic, anionic, anionic, and
amphoteric surfactants may all be used to create these pesticides. Some microbe-derived
biosurfactants exhibit anti-microbial efficacy against plant pathogens. They are a potential
biocontrol agent for sustainable farming. A number of microbe-derived biosurfactants
exhibit antibacterial properties against plant diseases, making it the preferred approach for
biocontrol in agriculture. Antagonizing rhizobacterial bio surfactants. Pesticides, fungi-
cides, and biosurfactants are often used in agriculture to suppress plant growth-promoting
microorganisms. Agronomists utilize surfactants to boost bacteria’s antimicrobial activity.
Several in vitro and in situ investigations have shown the importance of surfactants in
boosting insecticidal abilities [144].
Agronomy 2022, 12, 662 19 of 35

7.3. Biosurfactant Mediated Plant Growth Promotion in Soils

When a microbial consortium is utilized for bioremediation, biosurfactants accelerate
pesticide breakdown owing to the synergistic effect of microbial communities. Biosurfac-
tants enhance soil quality, participate in PGP activity, and degrade/solubilize pesticides,
according to reports. Pseudomonas sp. has been shown to have significant communication
with their host plant, displaying a variety of plant growth promoting (PGP) characteristics,
and is one of the most important biocontrol agents against pathogenic microorganisms.
Rhamnolipids derived from Pseudomonas aeruginosa promote herbicide trifluralin and pes-
ticide chlorpyrifos degradation in polluted soil [145]. PGP Pseudomonas sp. producing
BS was recently identified from pesticide-contaminated fields and common reed roots
with hydrocarbon-degrading capability [146]. Biosurfactant from Pseudomonas cepa-
cian degrade hydrophobic herbicide 2,4,5-trichlorophenoxyacetic acid [147]. Pseudomonas
aeruginosa possesses a biosurfactant that has the potential to breakdown huge amounts of
quinalphos (a pesticide) in a shorter period of time [148]. Similarly, the glycolipid biosur-
factant produced from pseudomonas species has been shown to improve methyl parathion
and endosulfan solubilization [145]. Moreover, while growing crops, biosurfactants may
be used to increase the availability of micronutrients that are often present naturally or
in wastes to plants. Additionally, it has been shown that rhamnolipids and lipopeptides
enhance plant tolerance to phytopathogens by boosting plant immune systems. Addi-
tional rhamnolipid uses as biopesticides, fungicides, and anti-zoospore agents have been
investigated in recent decades [149].

7.4. Food and Agricultural Waste as Substrates for Biosurfactant Production

Waste from the food and agricultural industry, as well as their related items, is a
significant source of carbon. Numerous studies have demonstrated that agricultural waste
lignocellulosic biomass can be converted into a variety of products [150], one of which is
supplements for biosurfactant production [151]. As a result, this lignocellulose biomass
offers microorganisms with a nutrient-rich habitat [152]. Sugar beets bagasse, banana,
citrus, residual cooking oil, fried palm oil, moringa, and yam residues have all been studied
for their propensity to create biosurfactants [152]. Glycolipids are extracted from low-cost
raw materials such as distillery waste, fruit and vegetable byproducts, bagasse, rapeseed,
sweet potato flour wastewater, bird’s feather protein, and beverage effluent [153]. The type
of biosurfactant generated is determined by the substrate utilized, since the content of the
substrate aids microbial growth throughout the fermentation [153]. Emulsifiers are used in
baking to enhance dough rheology, water retention, ingredient mixing and handling. They
are good additives and preservatives in food manufacturing [154].
Prior to biosurfactant production, agricultural waste was hydrolyzed to yield hy-
drolysate sugar that could be utilized as a substrate for biosurfactant production [142].
The enzymes and live cells that can be used to hydrolyze agricultural waste [155]. It has
been shown that Penicillium citrinum H9, a lignocellulolytic mold, is capable of hydrolyzing
agricultural wastes, such as straw and hay [155,156]. Penicillium citrinum H9 hydrolyzed
rice straw to produce as much as 209.25 g/mL sugar [155]. Achromobacter sp. BP(1)5, a
hydrocarbonoclastic bacterium isolated from Balongan oil sludge, was shown to be more
capable of making biosurfactant from rice straw hydrolysate [156]. Achromobacter sp. BP(1)5
can convert the hydrolysate sugar of rice straw and maize cobs by Penicillium citrinum H9.
Achromobacter sp. BP(1)5 showed 98 percent query cover with Achromobacter xylosoxidans.
The emulsifying efficiency of the crude extract of Achromobacter sp. BP(1)5 biosurfactant
on kerosene was 27.22 percent and 36.84 percent, significantly [157].
The generation of biosurfactants by different Lactobacillus bacteria utilizing sugar
beets molasses (cellulose fibers) and glycerol as substrate resulted in the formation of
glycolipids and glycolipopetides as multi-component mixture. Rodrigues et al. reported
same approach with minor modifications [158]. The fermentation was carried out in a
laboratory-scale bioreactor using De Man, Rogosa and Sharpe medium supplemented
with different quantities of lactose as substrate. The findings indicate that sugar beets
Agronomy 2022, 12, 662 20 of 35

molasses and glycerol provide a higher yield than typical synthetic broth. The solvent
approach might be used to extract the product from the broth [159]. Overall production
was 2.43–3.03 g/L when sugar beets molasses was used as the substrate. The yield obtained
in this research is much more than that obtained in earlier experiments, indicating that
supplementing broth with yeast and peptone extracts improves the yield of biosurfactant
synthesis [160].

7.5. Biosurfactant for Cosmetic Application

The natural moisturizing factor has been reported from the chemically modified
sophorolipid synthesized by Torulopsis bombicola. The highest application in cosmetic in-
dustry has been reported from the sophorolipids, rhamnolipids, and mannosylerythritol
lipids [32]. In the treatment of acne, dandruff, and body odors, sophorolipids play a crucial
role due to their anti-microbial properties. Similarly, in the formulations of deodorants,
nail care products, and toothpaste, and as an anti-wrinkle agent, rhamnolipids play a
crucial role [161]. Mannosylerythritol lipids are used as anti-aging skincare products and
are comparatively more effective than arbutin [162]. Application of lipopeptides as anti-
wrinkle agents have also been reported [163]. Biosurfactants are highly eco-friendly and
safer enough to be applied over the skin, and their non-toxic, non-irritant properties find
them a best place in the cosmetic industry [112]. Biocompatible and low toxic alternative
surfactant, bacterial biosurfactant are in increasing demand. The biosurfactant extracted
from Pseudomonas sp. was used in the formulation of toothpaste due to their foaming,
non-toxic and biodegradable properties thus of a great use in the cosmetics industry [151].
Mildness over the skin and superior interfacial activity of the newly synthesized biosurfac-
tants phospholipid biosurfactants cytidine diphosphate, cytidine triphosphate proves to be
a better solution in the cosmetic industry comparatively to the synthetic surfactants [164].

7.6. Environment and Bioremediation

The multi-biotech was used to market biosurfactants for the improved recovery process
(a subsidiary of Geodyne Technology). The biosurfactant produced by Bacillus licheniformis
JF-2 that seems highly helpful in the reduction of the viscosity of heavy crude oil has
reported characteristics like anaerobic, halotolerant, and thermal tolerant. Biosurfactants
were also attributed to the disappearance of crude oil from an oil tank [165]. Biosurfactants
can be used best in every way possible in the pulp and paper, carbon, textiles, ceramics
and uranium ore processing industries. Heteropolysaccharide was effectively used in the
ceramics processing industry, which was isolated from different sources. Biosurfactants
reduce surface tension between aquatic solutions and hydrocarbon mixtures. An analysis
of the soil was carried out using 18 strains of actinomycetes and 13 bacterial strains, with
high emulsification activity and an oil propagation property compared to the synthetic
surfactants. Oil spreading properties were isolated [166]. The selective means to test the
emulsifying activity of the isolates from mangrove hydrocarbon-contaminated sediment
were sucrose and Arabian light oil. The results finally increased the production of bio-
logical agents for the time of incubation. This method is thus ideal in the country with
low financial resources for the isolation and production of biosurfactants [167]. Fuel and
other oil material leaks lead to severe environmental pollution. The study currently in-
volves diesel degradation on Gram-negative microorganisms. A gravimetric analysis in
DJLB isolate is used to detect 65.4–83.12 percent diesel degradation. With an increase in
glucose consumption, the rate of diesel degradation and growth of the microbial cell have
increased. Thus, both diesel degradation and emulsification activity were increased by
glucose added. The increase in the production capacity for biofactants is announced in the
diesel degradation property. In diesel degradation, efficiency of Gram-positive bacteria
was identified from Stenotrophomonas sp. isolates DJLB, respectively [168]. An analysis
of the bacteria strain B160 showed that the surface tension was extraordinarily reduced
compared to other biostructure products, even under extreme pH conditions, tempera-
ture, etc. [169]. In general, the surface-active compounds during their growth phase were
Agronomy 2022, 12, 662 21 of 35

shredded into the surrounding medium [170]. Polycyclic aromatic hydrocarbon (PAH)
contamination addresses the serious threat to human beings and the environment and has
thus been implemented to meet these requirements with efficient and cost-competitive
remediation through the use of PAH. Thus, the bioavailability of PAHs is enhanced by
the addition of bioreactors locally or ex situally [171]. The report was published. The
production of biosurfactants increases the aqueous naphthalene concentration, thereby
increasing the solubilities of the substrate through the production of biofactants [172]. In
several industries, the process of the micelle formation of Pseudomonas aeruginosa synthe-
sized rhamnolipid has been of greater relevance. Comparatively, less CMC was shown in
the neutral dirhamnolipids than the negative. Centrifugation and dynamic light dispersion
techniques demonstrate greater aggregate formation at concentrations above CMC [173].
When Pseudomonas aeruginosa biosurfactants were investigated, promising algicidal actions
were shown against species such as Heterosigma akashiwo that causes severe harmful algal
blooming (HAB). Other species causing HAB such as Gymnodinium sp. were also reported
to inhibit Prorocentrum dentatum by rhamnolipids. During the prolonged application period
of the rhamnolipids, serious algae ultrastructural damage was reported. The cell mem-
brane is mainly integrated so that other cell organelles can be damaged [105]. A highly
potential bioremediation product in several sectors was revealed by the biodegradation
of diesel-contaminated water and the soil by rhamnolipids from Pseudomonas aeruginosa
J4 and surfactin from Bacillus subtilis, ATCC21332. The highest amounts were achieved
under glucose when bio synthesizing activity of bacteria was analyzed under different
carbon sources, allowing a good insulation and analytical method for microorganisms
in various industries [174,175]. When the biological activity of Pseudomonas aeruginosa
LBI rhamnolipids is analyzed, exponential interface tension reduction properties, which
were still far higher and more environmentally so than their synthetic equivalents, have
thus become predominant in different industry sectors [143]. The results show that max-
imum biosurfactant efficiency was obtained from palm oil use as a carbon source when
the biosurfactant synthesis of Pseudomonas aeruginosa A41 was investigated under various
circumstances. The C18:2 were also reported to be a highly unsaturated fatty acid, the
highest activity of oil displacement [176]. An analysis of the treatment of a biosurfactant
for treatment of artificially contaminated pyrene soil has shown high surface activity [177].
The use of molecular biological tools has optimized hyper-manufacturing microbial strains
for biosurfactant production [178]. In a few instances, there was no correlation between
cell characteristics and the carbon sources used. [179]. The report was submitted. In some
cases, however, the produced biosurfactant cannot be used during the bioremediation
process due to different factors, like Staphylococcus sp. [180]. In some cases, however,
microbes biosurfactants, isolated from a contaminated site, may use this to treat them
themselves [181]. After 15 days of Lactobacillus pentosus treatment with biosurfactant, 63%
of the biodegreasal efficiency of octane by autochthonous microflora in soil is achieved.
The application by various biosurfactants (e.g., rhamnolipids, sophorolipids) has shown an
increase of 50% in halogenation of halogenated compounds, etc. [142,182]. Phenanthrene
degradation of specific hexadecans and their production source has been reported. The
Microbially Enhanced Oil Research (MEOR) [183] was a highly potential area of research.
Seed germination stimulation, soil removal of motor oil, the biosurfactant produced by
S. marcescens UCP 1549, and a non-toxic, environmentally friendly compound were also
attributable [184].

7.7. Industries
Upon the analysis of the manipueira medium as a substrate for the production of
biosurfactants of bacterial isolates, the results showed a significant reduction in the surface
tension of the substrate, thus finding a place in various industrial applications as the most
suitable alternative media [185]. The advancement and the qualitative analysis of the
biosurfactant rhamnolipid over the synthetic surfactant sodium dodecyl sulfate have been
reported using a new methodology known as the drop-collapse method. This methodology
Agronomy 2022, 12, 662 22 of 35

proves to be more accurate, advantageous, efficient and cost effective, when compared to
other methods for the same processes like du Nouy ring method [186]. Upon the analysis
of the rhamnolipid biosurfactant synthesized by Pseudomonas aeruginosa LBM10, various
significant surface properties have been reported, thus making a promising class of com-
pounds in various industrial applications [187]. The production of biosurfactants by the
PAH utilizing bacteria isolated from soil contaminated with petroleum wastes were ana-
lyzed. The results revealed the significant production of the biosurfactants under various
substrates such as phenanthrene and naphthalene [188]. An anionic monorhamnolipid
generated by Pseudomonas aeruginosa was tested for its capacity to eliminate residual
hexadecane from sand columns. This research identified exploring the influence of low-
concentration rhamnolipid on cell transport in a variety of natural soils and resolving the
potential mechanisms [189]. Upon the experiments with Pseudomonas aeruginosa grown
on nitrate and protease peptone media, a direct relation between the enhanced biosur-
factant production and glutamine synthetase activity has been reported [14]. A recent
analysis revealed the methods for the enhancement of the di-rhamnolipid production over
the mono-rhamnolipid, synthesized by the strain Pseudomonas aeruginosa J16, thus for the
implementation in various industries [190]. A 100% efficiency for the removal of heavy
metals from soil and sludge by the biosurfactants has been reported [191]. The hazardous
spent hydrodesulfurization (HDS) from the petroleum refineries is treated with various
biosurfactants like rhamnolipids, etc., for the analysis of bioleaching, and significant re-
sults were obtained [192]. The heavy molecular biosurfactants were attributed with high
emulsifying properties, whereas the low molecular biosurfactants were attributed with
surface tension reducing properties. The combined effects of the biosurfactants synthesized
by Acidithiobacillus sp. and Meyerozyma quilliermondii have been attributed with significant
bioleaching properties against heavy metals such as copper, cadmium, etc. [192].

7.8. Pharmaceuticals
Biosurfactants were used as antibacterial, antifungal, antiviral, adhesive agents, im-
munomodulatory molecules, vaccines and also in-gene therapy. The adaptive nature by the
formation of different surface forms of Bacillus cereus ATCC14579 has been analyzed, and
its hemolytic activity and Gram-positive bacteria growth inhibition property have been
reported [193]. Many characteristic features like anti-tumor properties; self-assembling
potential into nanoparticles creates a great interest upon the application of this cyclic
lipopeptide biosurfactant [194]. Due to their non-toxic and safe characteristics, biosurfac-
tants were comparatively preferred than synthetic surfactants in the microemulsions drug
delivery systems (MDDS). Out of all the biosurfactants, lipopeptide and glycolipids were
the most preferable ones [195]. In the process of gene transfection, biosurfactant-based
liposomes were highly efficient compared to synthetic liposomes. [32]. The problems in the
drug delivery system are well addressed by the microspheres, nanoparticles, micelles, and
liposomes [196–199]. In the future, the use of biosurfactants, either alone or in combination
with other antibiotic or chemotherapeutic treatments, could provide a viable strategy for
combating diseases, biofilm development, and microbial growth. Additionally, new discov-
eries about biosurfactant’s ability to operate as an anticancer, immunomodulating, wound
healing, and drug delivery agent have piqued researchers’ interest in biosurfactant’s ability
to perform the aforementioned functions [159]. By utilizing a metagenomic approach,
a new gene related to biosurfactant production and hydrocarbon degradation has been
reported [200].

8. Critical Micelle Concentration of Various Biosurfactants

In recent years, microbial biosurfactants with low CMC, the potential to minimize sur-
face and interfacial stress and excellent emulsifying activity, coupled with good biodegrad-
ability and low toxicity, have drawn extensive attention [201]. Surfactant as an amphiphilic
compound may decrease the free interfacial energy to change the interface or surface area
at low levels. Surfactant molecules may shape micelles at high concentrations, above
Agronomy 2022, 12, 662 23 of 35

their CMC [202]. Using biosurfactants at concentrations greater than CMC to chemical
surfactants helped to decrease the absorptionratio, improve solutions and accelerate PAH
degradation [203]. At concentrations above the CMC, the biosurfactants increased the
partition rate of fluorine, phenanthrene and pyrene [204]. Phenol can assist in improved
solubilization of PAHs by biosurfactant. When phenol is applied to the solution, the biosur-
factant CMC reduces due to the development of mixed micelles. The cause for the enhanced
dissolution of PAHs could be the combined micelles [205]. A biodegradable, non-toxic
biosurfactant, Pseudomonas frederiksbergensis has been successfully obtained from piggery
waste water. This biosurfactant has been shown to improve the removal of ethylbenzene in
BTF. The removal rate of ethylbenzene at the 1000 mg/m3 inlet and of EBRT of 30 s peaked
at 87.2 percent with 0.1 CMC dosed biosurfactant. P. aeruginosa J4 generates rhamnolipids,
which lower water surface tension to 31 mN/m at a critical micelle concentration of about
50 mg/L. For diesel and kerosene, the biosurfactant attained emulsion indexes of 70 and
78 percent, respectively, at about 300 mg/L [206]. Similar findings have been reported in
swine waste water, suggesting that it might be an acceptable platform for the production of
biosurfactants. At 25 ◦ C, the CMC of the biosurfactant produced from Bacillus stratospheri-
cus sp. A15 strain was 46.8 mg/L. In the near future, BS15 might be created as an alternative
to antibiotics [207]. Biosurfactants have a molar mass of 500 to 1500 Da, and their effectivity
is assessed by CMC values, which typically vary from 1 to 2000 mg. Polymeric micelles
have a CMC in the range of 106–107 M, which is about a thousand times lower than the
CMC of low molecular weight surfactants, making them thermodynamically very stable
even at low concentrations [208]. The CMC level of biosurfactants is typically 1 to 200 mg
L21 [209]. B. Licheniformis induces lipopeptide lichenysin. It displays the strongest surface
and chelating action, indicating that antimicrobial activity is present in lichenysin. It has
an improvement in surface tension in water of approx. 29 mN m−1 and a lower CMC
15 mg L−1 [210]. Similar analysis was reported in B. Subtilis RSL 2 strain which developed
biosurfactant of 3.5 g/L at pH 4.0, 25◦ C and used in 7 days for 1 g/L crude petroleum as the
only C-source, defined as critical micelle concentration (CMC) lipopeptides of 0.5 g/L. In
comparison, the simultaneous feed of 0.5 CMC biosurfactant improved the biodegradation
of oil by 72 percent [211]. Tensile CMC provided by B. subtilis ATCC21, 332 was recorded as
50 mg L21 by [206] incorporating CMC reduced from activated carbon to 10 mg L21. [212]
recorded B. salmalaya 138SI biosurfactant CMC of 0.4%.

9. Recent Advances
9.1. Influence of Biosurfactant-Producing Bacteria on Compost
Composting is delineated as high microbial activity that results in the degradation of
most environmentally friendly resources, mainly organic mixes, and the stability of organic
residues. Microbes are extensively used in the degradation of organic matter, because of
the extensive use of microbes in degradation in recent decades, biosurfactants were widely
used in bioremediation and biodegradation; it is conceivable to use their benefits to increase
the compost quality too. Therefore, it is understood that biosurfactants, which are amphi-
pathic microbial compounds, may lower liquid–solid surface tension and increase organic
matter bioremediation [213]. Furthermore, boosting bacterial growth in the presence of
biosurfactant enhanced organic matter decomposition. The readily metabolized chemicals
in the substrates, as well as the sudden rise in enzyme concentration and microbial biomass,
contributed to the abrupt decomposition of organic matter. The impact of rhamnolipid and
tween 80 on bacterial diversity were analyzed, where the rhamnolipid promotes microbial
growth in composting. In the composting process, synergisms of Bacillus sp. and Strepto-
myces sp. resulted in a greater rate of breakdown of organic materials. The combination of
biosurfactant generating bacteria consortium and biosurfactant containing cell suspension
results in a rise in bacterial communities in composting; indicating that biosurfactants did
not impede the development of bacteria in composting and even had a minor stimulatory
impact on their growth. Biosurfactant producing bacteria can be cultivated in low cost
Agronomy 2022, 12, 662 24 of 35

medium like whey, and the obtained wild bacterial strains will help in speeding up of
decomposition of organic matter [214].

9.2. Effects of Biosurfactant on Green Waste Vermicomposting

Vermicomposting has recently gained popularity in green waste disposal. Along
with biosurfactant, vermicomposting is proven as a promising technology, as well as an
environmentally friendly method used in treating green waste disposal [215]. Microbial
inoculation and rhamnolipid inclusion will have hastened the decomposition of organic
matter into humic-like compounds, resulting in higher humic acid concentration in the
additive treatments [216]. Rhamnolipid will enhance nitrogen concentration by boosting
the activities of nitrogen-fixing microbes, as well as quicken the degradation of organic
carbon, leading to an increase in combined total nitrogen contents [217]. The substantial and
positive relationships between enzymatic interaction and bacteria populations show that
increased enzyme activity in vermicomposts was related to increased microbial numbers.
The impacts of utilizing the biosurfactant rhamnolipid, the lignolytic and cellulolytic
fungus Phanerochete chrysosporium, and the free-living nitrogen-fixing bacteria Azotobacter
chrococcum to vermicomposting green waste using Eisenia fetida revealed that the addition
of rhamnolipid of a single microorganism or in consortium substantially increased the
growth rate. The most efficient vermicomposting approach for green waste management
was recommended to combine vermiwash with microbial consortia of P. chrysosporium and
A. chrococcum [218].

9.3. Biosurfactants in Bionanotechnology

The combination of biosurfactants derived from microorganisms and nanoparticle is
now regarded as the next generation of green chemistry or bioengineering nanocatalyst
sources. The creation of nanoparticles using biosurfactant mediated synthesis has enormous
promise for environmental remediation [219]. However, the nanoparticles generated
using biosurfactant must be economically viable. Additionally, it must be energy efficient,
have a high rate of toxicant elimination, and be environmentally friendly. Microbes in
biosurfactants may stabilize and minimize nanoparticle production [205]. Microorganisms
have a role in the synthesis of nanoparticles such as gold, silver, and titanium. Among
many scientists and researchers, this innovative method of producing metallic nanoparticles
utilizing a reducing agent of biosurfactants presents new possibilities. Brevibacterium casei
MSA19 biosurfactants minimize nanoparticles and enable them to remain stable for around
two months by lowering aggregate formation via electrostatic force of attraction, enhancing
their use as an eco-friendly material for various products. Furthermore, research on
biosurfactant stabilized nanoparticles is still in its infancy [220]. Therefore, additional study
is urgently needed to stabilize nanoparticles utilizing biosurfactants before deploying them
in nanotechnologies [91].

10. Future of Biosurfactants

Biosurfactants are diverse entities with vast applications of biodegradation and biore-
mediation to clean up toxic contaminants. They are also used in the dairy, medicinal and
other sectors [220]. Such benefits and a wide variety of uses have culminated in continuous
interest in biosurfactants. In particular, an increasing debate is taking place regarding
environmental protection and the essential position of biosurfactants in the immediate
future, for example, utilizing sustainable by-products as substrates, minimizing waste
and eventually recycling treated waste. This has contributed to greater exposure being
given to these microbial products in manufacturing. The whole surfactants give increased
biodegradability and low toxicity to prevent the harmful effects of synthetic surfactants.
Amphiphilic, surface active materials are common in nature and are formed by plants, ani-
mals and microorganisms, including bacteria and fungi [221]. In complex biofilm systems,
biosurfactants are assumed to help the preservation of water and nutrient channels that
Agronomy 2022, 12, 662 25 of 35

are important to the survival and efficient attachment of the colony to the host, so a long
driven factor to handle the microbes for the production of synthetic surfactant.
The market performance of biosurfactants is currently constrained by high develop-
ment costs. Biosurfactant has many benefits over synthetic surfactant (Figure 10). Op-
timized conditions of growth/production with green substrates that are economically
viable and productive downstream will help produce a biosurfactant more sustainable and
economically viable. As such, for certain microorganisms, biosurfactants are important for
survival, promoting vital processes such as nutrient intake, associations with host surface
attachment, and detachment and degradation of competing microbes [222]. Two forms
of pollutants are produced when synthetic surfactants are used in industrial processes:
by-products of industrial activities and by-products of surfactants. Both are harmful for
humans and the atmosphere. In the environment, these toxins are persistent and they
are difficult to biodegrade [223]. A significant usage of biosurfactants will need a differ-
ent database, which would also allow researchers to operate very actively in the field of
biosurfactants with regard to their maximum contribution to industry and the ecosystem.

Figure 10. Comparison between biosurfactant and synthetic surfactant.

11. Prospective and Challenges of Biosurfactant Production and Application

• After many years of mediocre interest, biosurfactants have recently risen to the top of
many corporations’ agendas due to sustainability initiatives and green agendas.
• Biosurfactants may be customized for particular purposes and generated at a cost
competitive with chemical surfactants.
Agronomy 2022, 12, 662 26 of 35

• Several concerns must be addressed before large-scale exploitation may occur. The
two issues facing rhamnolipids are safety and yield.
• Despite their immune system impacts and virulence factors, rhamnolipids are likely
safe to use in many items, especially cleaning and laundry products.
• P. aeruginosa, being pathogenic is more challenging, although some companies have
clearly overcome it, and the identification of potential new nonpathogenic producer
organisms offers a potential solution, provided the products are suitable and yields
are acceptable.
• The quorum sensing system controls rhamnolipid synthesis in P. aeruginosa, prevent-
ing hyperproducing strains from being produced through mutagenesis and selection
or genetic manipulation. Inability to get large yields may prevent rhamnolipids from
being used in many applications.
• Thus, these do not appear to be any significant barriers to the widespread use of
biosurfactants in a variety of goods and applications over the next several years, and
we may anticipate a growing range of home items containing at least sophorolipids
and MELs on supermarket shelves.

12. Conclusions
The biosurfactant industry is a highly profitable and competitive industry that uses the
biodegradation and development benefits of the medicinal, cosmetic, petroleum and food
industry for renewable energy substrates. There has been a dramatic increase in the need for
surfactants in the world; however, most of the surfactants currently available are chemical
dependent. This study clarifies the potential benefits of biosurfactants for adapting their
actions in a number of applications. Moreover, this comparison with synthetic surfactants
helps to examine how physicochemical properties are affected by the composition of the
surfactants and to make appropriate formulation choices. Therefore, we focus on these
issues in order to provide a complete picture and perspectives for potential growth and
practical applications, through the sources, processes and physicochemical properties of
microbial biological factors. This is a description of the use of microbial biosurfactants
in accordance with our best experience, because it is largely due to basic simplicity, low
cost and widespread use. As the demand for biosurfactants is early on in growth, there
are a variety of niche applications where biofactants are used. The fetters found in this
sector are technical limitations, mainly their costs and disadvantages in terms of mixing
technology. Since understanding of biosurfactant producing strains needs to be extended to
include morphology, genetics and biochemistry, virulent strain screening and the advance
of process technology will help minimize production costs. Heavy metal remediation by
a biosurfactant happens either via complicated interaction with free metal residues or
by buildup at a solid–liquid interface, resulting in direct contact between the metal and
the biosurfactant. The biosurfactant metal complexes escape the soil surface and form
micelles as a result of the desorption pathway. Further precipitation and separation of
the biosurfactant from the metals is possible. Organic surfactants constitute a large part
of the surfactant industry with stronger controls over greener practices and responses to
enormous demand. Farmers are producing environmentally friendly surfactants from
numerous natural and renewable sources as they quickly become an attractive option on
the market. Using agro-industrial waste from both animal and plant origin to produce
biosurfactants might reduce production costs and make biosurfactants economically viable
and competitive with synthetic surfactants. Bio-based surfactants are meant to be used
to treat heavy metals, polluted soils and water, to treat skin conditions, to enhance oil
restoration, to preserve food and to eliminate plant disease. Recent research has shown that
using a biosurfactant in the aerobic composting of municipal waste, yard waste, and crop
residues enhances composting efficiency and product quality. Rhamnolipids are a kind of
biosurfactant that is widely utilized and accessible commercially on vermicomposting of
green manures. It is natural to use hereditary structure for the generation of mechanical
Agronomy 2022, 12, 662 27 of 35

biosurfactants using inexhaustible substrates as raw material in the future that super
dynamic microbial strains will be created.

Author Contributions: Conceptualization, methodology, software, validation and investigation—

B.R., N.K., M.K.A. and S.V.; writing—original draft preparation, E.G., P.P., N.K. and B.R.; writing—
review and editing, E.G., P.P., N.K., B.R., M.K.A. and S.V.; project administration and funding
acquisition, B.R. All authors have read and agreed to the published version of the manuscript.
Funding: This work was supported by the National Research Foundation of Korea by the Korean
Government (MSIT) (2020R1G1A1012069).
Conflicts of Interest: The authors declare no conflict of interest.

Nomenclature

EI24 Emulsification Index (%)

CMC Critical Micelle Concentration
Bs Biosurfactant
BSs Biosurfactants

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