Environmental Chemistry Letters (2023) 21:1787–1810

https://doi.org/10.1007/s10311-022-01552-4

REVIEW PAPER

Biotechnological methods to remove microplastics: a review

Uttpal Anand1 · Satarupa Dey2 · Elza Bontempi3 · Serena Ducoli3 · A. Dick Vethaak4,5 · Abhijit Dey6 ·
Stefania Federici3

Received: 6 October 2022 / Accepted: 25 November 2022 / Published online: 8 February 2023
© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2023

Abstract
Microplastics pollution is major threat to ecosystems and is impacting abiotic and biotic components. Microplastics are
diverse and highly complex contaminants that transport other contaminants and microbes. Current methods to remove
microplastics include biodegradation, incineration, landfilling, and recycling. Here we review microplastics with focus on
sources, toxicity, and biodegradation. We discuss the role of algae, fungi, bacteria in the biodegradation, and we present
biotechnological methods to enhance degradation, e.g., gene editing tools and bioinformatics.

Keywords Microplastics · Incineration · Microplastic degrading microorganisms · Biodegradation · Synthetic biology ·

Biotechnological interventions · CRISPR

Introduction 2022; Su et al. 2022; Federici et al. 2022). In 2018, global

plastic production increased to more than 360 million tons,
Plastic pollution is considered to be one of the most signifi- and it is expected to triple by the year 2050 (Gumel et al.
cant threats to global ecosystems and is known to impact 2013; Plastics Europe 2020). According to a survey made
both the abiotic and biotic components (Ogonowski et al. by Plastics Europe (the Association of Plastics Manufac-
2018; Everaert et al. 2020; Lusher et al. 2021; Liu et al. turers in Europe) and the European Association of Plastics
Recycling and Recovery Organisations, Asia is the largest
producer and consumer of plastic goods, with China contrib-
* Satarupa Dey
[email protected] uting the lion’s share (32%) to this “white pollution”, while
the rest of Asia produces nearly 19%.
* Abhijit Dey
[email protected] Whereas, Europe, Canada, Mexico, and the USA produce
fewer plastics than Asia (Tiwari et al. 2020; https://p​ lasti​ cseu​
* Stefania Federici
[email protected] rope.​org/​knowl​edge-​hub/​plast​ics-​the-​facts-​2021/).
Moreover, the coronavirus disease 2019 pandemic has
1
Zuckerberg Institute for Water Research, Jacob Blaustein increased the use of one-time useable plastic wares, gloves,
Institutes for Desert Research, Ben-Gurion University masks, tissues, and other personal protective equipment
of the Negev, 8499000 Midreshet Ben Gurion, Israel
which along with the generation of municipal waste has
2
Department of Botany, Shyampur Siddheswari exacerbated the plastic pollution crisis (De-la-Torre and Ara-
Mahavidyalaya, University of Calcutta, Ajodhya,
Shyampur, Howrah 711312, India gaw 2021; Morgana et al. 2021; Patrício Silva et al. 2021;
3
Yang et al. 2022). Most single-use masks and personal pro-
Department of Mechanical and Industrial Engineering,
INSTM Unit of Brescia, University of Brescia, Via Branze
tective equipment are made up of various polymeric sub-
38, 25123 Brescia, Italy stances such as polypropylene, polyurethane, polyacryloni-
4
Department of Environment and Health, Vrije Universiteit
trile, polyethylene, and polystyrene. Also, the imposition of
Amsterdam, Amsterdam, Netherlands lockdown has led to the surge in the use of different types of
5
Institute for Risk Assessment Sciences, Department
plastic largely composed of high-density polyethylene, low-
of Population Health Sciences, Faculty of Veterinary density polyethylene, polypropylene, and polyethylene tere-
Medicine, Utrecht University, Utrecht, Netherlands phthalate. The incorrect disposal and poor waste manage-
6
Department of Life Sciences, Presidency University, 86/1 ment of plastic items have led to their ubiquitous presence in
College Street, Kolkata, West Bengal 700073, India

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1788 Environmental Chemistry Letters (2023) 21:1787–1810

all environments globally, and due to their persistent nature only few articles have addressed plastic degradation focus-
and low biodegradability, they will remain in these environ- ing on the use of modern biotechnological methods in the
ments for prolonged periods of time (Andrady 2017; Aragaw enhancement of microplastic degradation and there remains
2020; Patrício Silva 2021). a lack of knowledge with respect to biotechnological inter-
It has been reported that around 4.8–12.8 million metric ventions for microplastic removal (Danso et al. 2019; Patrí-
tons of plastic debris are disposed of in the ocean without cio Silva et al. 2021). Therefore, it is crucial to summarize
a proper management strategy (Jambeck et al. 2015). This and analyze the current state of knowledge to determine
disposal of plastic debris into the ocean has led to a series microplastic degradation by microorganism, as well as to
of impacts on marine life (Galloway et al. 2017; Mendoza promote a better understanding of how modern biotech-
et al. 2018; Peng et al. 2020) and human health (Keswani nological methods can be enabled to manage and degrade
et al. 2016). They result in problems like ingestion, entangle- microplastics.
ment, and suffocation to different marine species leading to This review provides some background information on
reduced life quality, and impairment of feeding and repro- the impact of microplastic sources, their effects on marine
ductive ability (Staffieri et al. 2019; Wilcox et al. 2018). In life including microalgae, and their potential impact on
addition, most plastics are positively buoyant and can be humans. Then it categorizes the different types of micro-
transported over a long distance, acting as carriers of non- organisms and enzymes associated with the degradation of
native and invasive species (De-la-Torre et al. 2021). microplastics. Finally, different biotechnological methods
Wasted plastics, exposed to ultraviolet (UV) irradiation to increase the efficiency of bacterial cell degradation of
and other environmental degradation processes, can degrade microplastics and their possible application in field studies
and fragment into smaller pieces: large microplastics are discussed at length.
(1 mm–5 mm), microplastics (1 µm–1 mm) and nanoplastics
(1 nm–1 µm) (Waller et al. 2017; De-la-Torre 2020; Atugoda
et al. 2022). However, there is no scientific consensus on the Sources of microplastics
definition of microplastics. Here we define microplastic as
plastics that are ≤ 5000 µm (or ≤ 5 mm) in diameter (ISO/ Microplastics derive from two different origins. The primary
TR 21960:2020), with "large microplastic" referring to the microplastics are generated from cosmetics, household prod-
larger portion (> 1000 µm or > 1 mm) (Frias and Nash 2019; ucts, drug delivery systems (Patel et al. 2009), and polymeric
GESAMP 2019; Hartmann et al. 2019; Hale et al. 2020; raw materials (pellets, flakes, powders) composed of poly-
Koelmans et al. 2020). Microplastics are the major contribu- ethylene, polystyrene, polyvinyl chloride, polyamide nylon
tors to plastic pollution in the marine ecosystem, freshwater 6 and polypropylene, among others. Personal care products
ecosystem, soil ecosystem, and agroecosystem (Nizzetto such as toothpaste, scrubs, cleaning materials and cosmet-
et al. 2016; Mendoza et al. 2019; Wang et al. 2019b; Wong ics are known to contain irregularly shaped microplastics of
et al. 2020; Chia et al. 2021; Razeghi et al. 2021) and are 0.5 to < 0.1 mm in diameter, which are mainly marketed as
known to be globally ubiquitous (Jambeck et al. 2015). They “micro-beads” or “micro-exfoliates” and contribute to pri-
also contaminate soil through sewage sludge and wastewater mary microplastics (Fendall and Sewell 2009). However, the
which are widely used as fertilizers. Moreover, plastics are existing wastewater treatment plants have shown that tertiary
also used in agriculture mainly in a greenhouse, low tunnels treatment of water is not a source of microplastic pollution,
and mulching and also as a coating for fertilizers, hormones, as these pollutants are effectively removed by the skimming
pesticides, and packaging material (Nizzetto et al. 2016). and settling treatment processes (Carr et al. 2016).
Recently, several studies have investigated the distribu- The secondary microplastics are generated due to exten-
tion, uptake, fate, behavior, effects, and removal strategies sive fragmentation of large plastic items or particles in pres-
of microplastics (Bahtt et al. 2021; Wong et al. 2020; Anik ence of environmental factors such as high temperature and
et al. 2021). Nevertheless, the effectiveness of the methods exposure to UV radiation, stress, reactive ozone, oxidation,
developed for microplastic remediation still remains unclear. and atmospheric pressure (Tiwari et al. 2020; John et al.
Research in microplastic degradation has progressed focus- 2021). Polymeric materials can withstand oxidative-ther-
ing on biological and non-biological approaches. Microplas- mal degradation only when antioxidants and stabilizers are
tic treatments enabled by the action of microorganisms such added. Physical abrasion also generates secondary micro-
as algae, fungi, and bacteria are considered attracting tools plastics. Moreover, biological agents like bacteria, fungi and
for cost-effective and eco-friendly degradation approaches. algae are known to produce a plethora of enzymes which
While research papers and reviews have recently been pub- play a crucial role in microplastic degradation (Chia et al.
lished on the microorganism-mediated degradation and 2020; John et al. 2021; Othman et al. 2021; Chen et al. 2022;
remediation strategies (Chen et al. 2022; Bahtt et al. 2021; Manzi et al. 2022; Miri et al. 2022; Zhu et al. 2022).
Qin et al. 2021; Kotova et al. 2021; Cholewinski et al. 2022),

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Environmental Chemistry Letters (2023) 21:1787–1810 1789

Fig. 1  Microplastics impact on living organisms. A Microplastics cause metabolic dysfunctions, and cause impairment of photosynthe-
can enter aquatic fauna food chain, causing intestinal blockage and sis due to shading effects. C The three main routes of human micro-
alteration in nutrients absorption, endocrine disruption, immunologi- plastics exposures are identified as ingestion, inhalation, and dermal
cal and neurological effects, and loss of reproductive functionalities. contact, triggering inflammatory and immune reactions. Created with
B Microplastics and toxic leachate can damage microalgal cell walls, BioRender.com

Secondary microplastics are mostly generated from a addition, microplastics provide a novel habitat for the growth
large plastic object made from the same polymers improp- of microbial biofilms containing algae, bacteria and fungi
erly disposed of in land and water systems, which undergoes and can potentially spread microbial pathogens and antimi-
physical abrasion leading to the weakening of the chemical crobial resistance (Zettler et al. 2013; Wu et al. 2019; Guo
bonds and subsequent oxidative-thermal degradation (Ger- et al. 2020; Yuan et al. 2020).
ritse et al. 2020). Other secondary microplastic sources As most microplastics have a size range similar to the
include the disintegration of synthetic fibers during the foods that are normally consumed by the zooplankton,
washing of clothes and commercial activities like thermal they can accidentally enter the food chain (Gregory 2009).
cutting of polystyrene. The increased use of single-use plas- This selection of microplastics instead of food particles
tics has contributed to the overproduction of polyethylene, can cause a loss of energy resources and also sublethal
polypropylene, and polyethylene terephthalate products. effects on the species reproductive pattern (Enders et al.
Moreover, a wide range of electronic, automobile, textile, 2015). Moreover, the ingested plastic can cause intestinal
and paint industries also discharge microplastic products blockage, which subsequently reduces the absorption of
into the water bodies and river catchment areas which can nutrients and also causes a change in hormonal balance
lead to microplastic pollution (Kay et al. 2018; Wang et al. (Derraik 2002). The improper absorption of the nutrients
2019b; Chia et al. 2021). may result in a decrease in the energy reserves and defi-
ciency in food assimilation which in turn impacts growth
and reproduction (Besseling et al. 2013; Wright et al.
Impact of microplastics on marine 2013a) and also decreases the ability of the organisms to
organisms and humans survive in adverse environmental conditions (Bugoni et al.
2001).
Microplastics could have a huge impact on the aquatic flora Microplastic exposure can also cause other sublethal
and fauna (Sathicq et al. 2021) (Fig. 1a) as they act as a effects in marine organisms, including oxidative stress,
vector for the transport of absorbed heavy metals, bacte- altered gene expression, inflammation, and effects on the
rial fish pathogens, multidrug resistant E. coli, persistent immune system and central nervous system (for review see
organic pollutants etc. (Enders et al. 2015; Viršek et al. Wright et al. 2013b; Vethaak and Martinez 2020). These
2017; Caruso, 2019; Song et al. 2020), and the possible adverse effects may be caused by residual monomers and
leaching of chemical components added during their manu- additives release from plastic particles rather than by the
facturing process (Groh et al. 2019; Bacha et al. 2023). In particles themselves. Smaller microplastics below 10 µm

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1790 Environmental Chemistry Letters (2023) 21:1787–1810

were found to be more toxic to aquatic organisms than 39,000–52,000 microplastics each year. These microplas-
larger plastic particles. tics cause intestinal blockage and result in the inflamma-
The fishing and aquaculture industry is strongly impacted tory response and changes in gut microbe composition and
by microplastic pollution, and its viability and productivity metabolism.
are affected by the presence of plastic waste in water bodies As stated earlier, microplastics can also be inhaled and
(Rochman et al. 2015) and microplastic contaminated sea- the outdoor microplastic concentration is between 0.3 and
food products (Bråte et al. 2016; Smith et al. 2018; Curren 1.5 particles per ­m3, whereas the indoor concentration is
et al. 2020; Li et al. 2021; Pan et al. 2022). The ubiquitous, 0.4–56.5 particles per ­m3 (Dris et al. 2017). The deposition
persistent, and anticipated increase in microplastics pollu- of microplastics is largely dependent on the size and den-
tion could in the long run also have a significant impact sity of the particles. The less-dense smaller particles tend to
on marine biodiversity and ecological processes, such as deposit deepest in the lungs, causing the release of chemot-
primary producers at the basis of the food chain (Guzzetti actic factors and resulting in chronic inflammation (Oliveira
et al. 2018; Vethaak and Martinez 2020). et al. 2020a; b). The presence of microplastics in human lung
The impact of microplastics on the growth and diversity tissue and human blood was very recently confirmed (Jenner
of the microalgal population usually varied (Liu et al. 2019) et al. 2022; Leslie et al. 2022).
(Fig. 1b). According to Sjollema et al. (2016), uncharged It was also speculated that nanoparticles can transverse
polystyrene particles negatively affected microalgae growth the dermal barrier (Revel et al. 2018) causing low inflam-
of Dunaliella tertiolecta at high concentrations (250 mg/l) matory reactions and fibrous encapsulation (Oliveira et al.
and with decreasing particle size. The study made by Khoi- 2020a; b). Once in contact with mucous membranes or
roni et al. (2019) reported that the growth of Spirulina sp. absorbed by the body, microplastics generate oxidative stress
was severely impacted by the presence of high concentra- and cytotoxicity, mainly due to their persistent nature in the
tions of microplastics, especially due to shading effects and body and the leaching of toxic additives, which may result
reduced light intensity, with subsequent impairment of pho- in inflammation, immune reactions, neurological damage,
tosynthesis, and the damage of microalgal cell walls (Khoi- metabolic disruptions, deoxyribonucleic acid (DNA) dam-
roni et al. 2019). Li et al. (2018) reported that both poly- age, and even cancer (Wright and Kelly 2017; Revel et al.
ethylene and polypropylene gradually degrade and generate 2018; Rahman et al. 2021; Vethaak and Legler 2021; Gruber
microsized plastics and release potentially toxic additives et al. 2022).
including plasticizers, polychlorinated biphenyls, dichlorodi- Overall, microplastics and nanoplastics derived from
phenyltrichloroethane, and heavy metals such as cadmium, several sources, in particular from the environmental deg-
chromium, bromium, copper, and titanium which cause cell radation of waste plastics, can exert toxic effects on organ-
membrane damage and growth inhibition. Capolupo et al. isms in all trophic levels. They can enter aquatic fauna food
(2020) reported that in the case of Raphidocelis subcapitata chain, causing intestinal blockage and alteration in nutri-
and Skeletonema costatum the cell growth is inhibited due ents adsorption, endocrine disruption, immunological and
to the leaching of additives. neurological effects, and loss of reproductive functionali-
Raphidocelis subcapitata, on the other hand, showed a ties. Micro- and nanoplastics and toxic leachate can dam-
higher growth rate in presence of plastic microbeads (Can- age microalgal cell walls and metabolic dysfunctions, and
niff and Hoang 2018), and similar enhancement of cell impairment of photosynthesis due to shading effects. Micro-
growth and photosynthetic activity was evident in Dunaliella and nanoplastics can enter the human body through inges-
salina in presence of larger microplastics (Chae et al. 2019). tion, inhalation, and dermal contact, triggering inflamma-
This enhancement of growth seemed related to the trace con- tory, and immune reactions.
centration of additive chemicals such as stabilizers, phtha-
lates, and endocrine disruptors, which are leached out of
microplastics (Chae et al. 2019). Processes of microplastic degradation
The potential effect of microplastics on humans is far
from understood and requires further research (Vethaak and Most of the conventional methods discussed for the reuse of
Legler 2021). Microplastics have been reported in differ- microplastic degradation include a primary method where
ent foods such as mussels, commercial fish, and table salt the plastic scrap is re-introduced in the heating cycle of the
(Li et al. 2018) and three different pathways of exposure to processing unit, followed by the conversion of waste to new
microplastics are identified, including ingestion of food con- plastic products by blending it with a virgin polymer which
taining microplastics, inhalation of microplastics in the air, can considerably reduce the cost of production. Sometimes,
and dermal contact with these particles (Revel et al. 2018) plastic wastes are chemically or thermochemically altered to
(Fig. 1c). According to a study made by Cox et al. (2019), be recycled in the industrial loop. However, in most cases
it was estimated that each person usually intakes around due to poor management strategies, these microplastic

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Environmental Chemistry Letters (2023) 21:1787–1810 1791

Fig. 2  Processes of microplastic

degradation through abiotic
and biotic combined factors.
Microplastics derive from the
fragmentation of plastic debris
and can further degrade into
smaller plastic particles at the
nanoscale (nanoplastics). Due
to abiotic factors and extracel-
lular enzymes, nanoplastics are
degraded into oligomers and
monomers, then internalized
by microorganisms and used
as carbon source, resulting in
the complete mineralization of
plastic. Created with BioRen-
der.com

particles are not disposed of properly or segregated properly. by microbes (see Fig. 3 for a representative scheme of poly-
Most of them get mixed up with the organic components in a ethylene mineralization).
landfill, which is used for composting or anaerobic digestion, On complete mineralization, carbon dioxide is evolved
leading to excessive pollution and to the production of toxic along with the formation of several intermediate compounds
compounds such as dioxins, phthalates, tetrabromobisphenol which are used as a source of energy to promote the growth
A, polybrominated diphenyl ethers, and toxic metals such as of microbes. The different extracellular enzymes that play a
cadmium and lead (Verma et al. 2016). pivotal role in microplastics degradation include esterases,
Currently, several physical, as well as chemical methods lipases, lignins peroxides, laccases, and manganese perox-
are popularly used for disposing of microplastic particles ides, which increase the hydrophilicity of microplastics and
including incineration, landfilling, and recycling. Chemical convert them to carbonyl or alcohol residues (Taniguchi
recycling processes such as pyrolysis are extremely popu- et al. 2019). Hydrolases enzymes, such as lipases, esterase,
lar at the commercial level (Thiounn et al. 2020). In the and cutinase, act on plastic surfaces and degrade micro-
slow pyrolysis methods, the plastic waste is converted to plastics by enhancing the chain cleavage reactions. These
a mixture of char and tarry products which are treated at enzymes fail to diffuse into the polymer, but they act on the
three different temperatures including 300, 425 and 550 °C surface resulting in the formation of cracks. The monomers
(Dussud et al. 2018). There exist several pieces of research generated are assimilated into the cytoplasm of microbes and
which focus on the pyrolysis of polypropylene, polystyrene, finally enter into different metabolic pathways.
and polypropylene from where heat energy can be recov- Although extensive research has already been done using
ered. The contaminated, mixed, or degraded residues which extracellular enzymes in microplastics biodegradation, very
are not suitable for recycling can be used as feedstocks for little information is still now available on the role of intracel-
waste-to-energy strategies such as pyrolysis (an endothermic lular enzymes in the degradation of microplastics; moreover,
cracking process without oxidation), and incineration (oxi- the pathways involved in the uptake of monomers are still
dation of plastics) (Prata et al. 2020). not clear. Normally, after the fragmentation of the micro-
Degradation of microplastics can occur by physical, plastics, the metabolic intermediates with carbonyl and
chemical, and biological methods and the biological deg- hydroxyl groups are metabolized within the cell using the
radation process is associated with a plethora of enzymes tricarboxylic acid cycle and β -oxidation pathway (Taniguchi
(Padervand et al. 2020; Bacha et al. 2021; Fig. 2). The basic et al. 2019). This process is followed by the complete miner-
process includes steps like the degradation of polymers to alization of plastic debris into ­H2O, ­CO2, ­N2, and ­CH4 (Zet-
smaller particles, followed by the degradation of the smaller tler et al. 2013). Researchers have made a thorough research
polymers to oligomers, dimers, and monomers. This degra- on the process of surface colonization of the microplastics
dation is followed by mineralization steps which are aided by degrading consortium forming a biofilm on the particles.

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1792 Environmental Chemistry Letters (2023) 21:1787–1810

Fig. 3  Mineralization process

for polyethylene. Due to a
combined effect of abiotic fac-
tors and extracellular enzymes,
plastic undergoes bio-deteriora-
tion and bio-fragmentation pro-
cesses, resulting in the release
of oligomers and monomers.
Thanks to specific cell transport
mechanisms, monomers are
internalized by microorgan-
isms and enter the catabolic
pathways as carbon source. The
final products of cells’ aerobic
metabolism, which results in
the mineralization of plastic, are
carbon dioxide and water. Cre-
ated with BioRender.com

The attachment process of the microbes occurs through sev- products of cells’ aerobic metabolism, which results in the
eral mechanisms including biofouling, and degradation of mineralization of plastic, are carbon dioxide and water.
plasticizers followed by the attack on the backbone of the
polymer which is subsequently associated with hydration
and penetration of microbes in the polymer structure. Techniques to monitor microplastic
Moreover, for efficient biodegradation, several factors are biodegradation
required which include the availability of potential microbial
degrading organisms which possess suitable enzymes and Different techniques have been applied to study microbial
metabolic pathways and other environmental factors such as degradation of microplastics, which includes weight loss
temperature, pH, salinity, and moisture content (Raddadi and measurement due to leaching, ­CO2 production due to deg-
Fava 2019; Syranidou et al. 2019; Matjašič et al. 2021; Miri radation of low molecular weight polymers and loss of addi-
et al. 2022; Lin et al. 2022). The biodegradation of micro- tives which affect the strength of microplastics (Baldera-
plastics is also influenced by the surface and the structure of Moreno et al. 2022). To get direct proof of the degradation
the polymer, amorphous and crystalline regions, crystal size, process, morphological, chemical, thermal, and structural
and lamellar thickness of polymers. Shabbir et al. reported properties are investigated using various techniques/methods
polyhydroxyalkanoates depolymerase enzymes to hydrolyze such as scanning electron microscopy, laser diffraction par-
the chains structures in the amorphous state on the surface ticle, differential scanning calorimetry, dynamic light scat-
of fragmentation films followed by erosion of chains in the tering, X-ray diffraction, etc. (Huang et al. 2022). Chemi-
crystalline state (Shabbir et al. 2020). cal changes are usually tracked by vibrational spectroscopy
In brief, microbial microplastic degradation involves techniques, such as Fourier transform infrared spectroscopy,
abiotic and biotic combined factors. Microplastics derive nuclear magnetic resonance, mass spectrometry, and gas
from the fragmentation of plastic debris and can further chromatography (Donelli et al. 2009; Chamas et al. 2020;
degrade into smaller plastic particles at the nanoscale (nan- Ivleva, 2021; La Nasa et al. 2021; Miao et al. 2020; Du et al.
oplastics). Due to a combined effect of abiotic factors and 2021).
extracellular enzymes, plastic undergoes bio-deterioration The gravimetric weight loss method is another widely
and bio-fragmentation processes, resulting in the release of used method for the determination of the biodegradation of
oligomers and monomers. Thanks to specific cell transport microplastics. However, this method should be used care-
mechanisms, monomers are internalized by microorganisms fully as biodegradation of microplastics is an extremely
and enter the catabolic pathways as carbon source. The final slow process (Raddadi and Fava 2019), and depends on
the incubation time and the assay conditions. Spectroscopy

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Environmental Chemistry Letters (2023) 21:1787–1810 1793

techniques can be used to determine the microplastics bacteria and algae including Chlorella fusca var. vacuolate,
biodegradation efficiency of microbes, by monitoring the Chlamydomonas mexicana, Stephanodiscus hantzschii, and
changes in the chemical functional groups of the polymer Chlorella vulgaris (Hirooka et al. 2005; Li et al. 2009; Ji
due to microbial activity (Singh and Sharma 2008). These et al. 2014).
changes may occur in hydrogen bonding, end-group modifi- In most cases, the degradation of microplastics is associ-
cation, cross-linking and copolymer compositions. Fourier- ated with the formation of biofilms on the surface of poly-
transform infrared spectroscopy is considered one of the mers. Several cyanobacterial strains, including the genus
most efficient methods for the detection of chemical changes Microcystis, Rivularia, Pleurocapsa, Synechococcus,
in the polymer and reference spectra of a wide variety of Prochlorothrix, Leptolyngbya Calothrix, and Scytonema,
polymers are available in libraries for comparison (e.g., were also able to form biofilms on the microplastic poly-
Celina et al. 1997). mers (Bryant et al. 2016; Debroas et al. 2017; Dussud et al.
Scanning electron microscopy allows the detection of 2018; Muthukrishnan et al. 2019). Besides cyanobacterial
microbial biofilm formation and surface degradation by species, diatoms are also present in the biofilms which help
monitoring the physical aspects of the polymer surface. in photosynthesis (AmaralZettler et al. 2020).
The evaluation of polymer biodegradation can be done by With the recent advances in different biotechnological
checking the formation of cracks and holes in the polymer processes, several genetically modified microalgal cell fac-
(Raddadi and Fava 2019). tories can be created which are capable of producing and
Modification of microplastics tensile strength and elon- secreting enzymes required for plastic degradation (Shen
gation at break are the signals of microbial biodegradation. et al. 2019). Green microalgae Chlamydomonas reinhardtii
Indeed, microbial degradation leads to significant changes was genetically modified to produce polyethylene tereph-
in mechanical properties and biochemical modification of thalate hydrolase, able to degrade polyethylene terephtha-
polymers due to the formation of cross-linking bonds or film late films and terephthalic acid (Kim et al. 2020). A similar
disintegration and shortening of the polymer chain (Nowak modification was also successfully done in P. tricornutum
et al. 2011; Othman et al. 2021). which produced polyethylene terephthalate hydrolase and
showed catalytic activity against polyethylene terephthalate
and the copolymer polyethylene terephthalate glycol (Moong
Role of algae in the degradation et al. 2019).
of microplastics In short, microalgae could serve as effective microplastic
degraders, thanks to their capability of using plastic mono-
Microalgae and their enzymes and toxins can be effectively mers as carbon source by producing degrading enzymes
used in the biological breakdown of polymeric material and the ease of culture. The possibility of genetically engi-
(Moog et al. 2019; Chia et al. 2020; Manzi et al. 2022). The neering algae strains to enhance degradation capability has
main advantage is that they do not require a rich carbon provided a promising environmentally friendly solution to
source for growth when compared to the bacterial system biologically degrade polyethylene terephthalate using micro-
and are adapted to a wide variety of habitats where most of algae via synthetic biology.
the microplastics occur (Yan et al. 2016). Microalgae are
known to colonize the plastic surfaces in wastewater streams
and this adhesion initiates plastic degradation by the produc- Fungal degradation of microplastics
tion of ligninolytic and exopolysaccharide enzymes. Mostly
these polymers serve as a carbon source and increase the The fungi largely consist of a diverse group of organisms
cellular proteins and carbohydrates and increase the growth which are largely saprotrophs, or opportunistic or obligate
rate. Very recently, surface degradation or breakdown of parasites. They have tremendous adaptivity and can grow
low-density polyethylene sheet through algal colonization in a wide range of habitats both aquatic and terrestrial eco-
has been identified using scanning electron microscopy systems under diverse environmental conditions. As well as
(Sanniyasi et al. 2021). being able to tolerate several toxic chemicals and metals,
Algal biodegradation occurs mainly in different processes they produce a diverse range of extracellular enzymes and
such as corrosion, hydrolysis, penetration, fouling, etc. (Chia natural biosurfactants such as hydrophobins that can degrade
et al. 2020). Both Oscillatoria subbrevis and Phormidium complex polymers into simple monomers, making them a
lucidum were also found to be able to colonize the surface source of electrons and carbons for microorganisms, thus
of low-density polyethylene and degrade it without any pro- facilitating the degradation and mineralization of complex
oxidative additives or pretreatment (Sarmah and Rout 2018). pollutants (Olicón-Hernández et al. 2017).
Bisphenol A, an additive with estrogenic activity commonly The main genus associated with the degradation of differ-
found in the polymers, was degraded by a combination of ent types of polymers such as polyethylene, polypropylene,

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1794 Environmental Chemistry Letters (2023) 21:1787–1810

and polyethylene terephthalate includes Zalerion mar- the preservation of hyphal wall integrity and formation of
itimum, Aspergillus niger, Cladosporium, and Penicillium spore wall and utilized cofactors like heme, NADPH + ­H+,
simplicissimum (Paço et al. 2017; de Oliveira et al., 2020a, and FAD.
b; Devi et al. 2015), which use microplastics as sole carbon On the other hand, extracellular enzymes include hydro-
source after degradation by extracellular enzymes. They lases which are involved in the breakdown of complex pol-
promote the formation of different types of chemical bonds ymers (Sánchez, 2009) and increase the solubility of the
(having carbonyl, carboxyl, and ester functional groups) and pollutants subsequently reducing bioaccumulation (Olicón-
decrease their hydrophobicity. Similar degradation of polyu- Hernández et al. 2017). Enzymes belonging to the class II
rethane was evident in fungal strains such as Aspergillus peroxidases such as manganese peroxidase and lignin per-
fumigatus, Aspergillus tubingensis, Cladosporium pseudo- oxidase, laccases, and dye-decolorizing peroxidases, which
cladosporioides, Fusarium solani, and Penicillium chrys- oxidize a wide range of substrates, can be used as efficient
ogenum and strains of Pestalotiopsis microspora (Khan et al. tools for environmental cleaning. Lignin degrading fungi
2017; Álvarez-Barragán et al. 2016; Magnin et al. 2020; produce laccase which catalyzes the oxidation of aromatic
2015; Russell et al. 2011). and non-aromatic substrates such as chlorophenolic or non-
In most cases, serine hydrolase plays a pivotal role in phenolic compounds (polymethylmethacrylate and polyhy-
polyurethane degradation. Degradation of high-density poly- droxybutyrate (Straub et al. 2017). The thermostability of
ethylene from marine coastal areas by two fungal strains these enzymes may promote their uses in large-scale reactors
Aspergillus tubingensis VRKPT1 and Aspergillus flavus where the degradation of polypropylene can be carried out
VRKPT2 was reported to be 6.02 ± 0.2 and 8.51 ± 0.1%, at a high temperature, facilitating high kinetics reactions. A
respectively (Devi et al. 2015). Recently, Kunlere et al. detailed list of different fungi associated with microplastic
reported the promising degradation of low-density polyeth- degradation is reported in Table 1.
ylene by Mucor circinelloides and Aspergillus flavus isolated Overall, a wide variety of fungal strains are capable of
from a municipal landfill (Kunlere et al. 2019). degrading plastics into more environmentally acceptable
Pretreatment of the microplastics, for example, poly- compounds, thanks to the production of a plethora of intra-
ethylene, with chemicals such as nitric acid and sodium cellular and extracellular enzymes, including oxidases and
hydroxide is known to accelerate the rate of biodegrada- hydrolases, and natural biosurfactants such as hydrophobins.
tion of polyethylene by Aspergillus niger (Nwachukwu et al.
2010). Physical pretreatment processes including thermo-
oxidization at 80 °C for 15 days were required to cause the
degradation in low-density polyethylene mediated by Asper- Bacterial degradation of microplastics
gillus niger and Penicillium pinophilum, showing 0.57 and
0.37% after incubation over 30 months (Volke-Sepúlveda Diverse studies have been conducted using bacteria for the
et al. 2002). Similarly, Aspergillus spp. and Lysinibacillus degradation of microplastics. Bacteria capable of degrading
spp. showed 29.5% of biodegradation of UV-irradiated and microplastics have been isolated from a wide range of habi-
15.8% of biodegradation of non-UV-irradiated polymer films tats including contaminated sediments, wastewater, sludge,
(Esmaeili 2013). compost, municipal landfills (Mehmood et al. 2016; Awasthi
et al. 2020), and also from extreme climatic conditions like
the Antarctic soils, mangrove, and marine sediments. More-
Fungal enzymes associated over, microplastic degrading microbes have also been iso-
with the degradation of microplastics lated from the gut microflora of earthworms. It is generally
reported that microbes living in polluted sites often develop
Fungi produce a diverse range of intra and extracellular an ability to activate the enzymatic system responsible for
enzymes which can catalyze diverse reactions and have the microplastic degradation.
ability to degrade petroleum-based polymers. The intracel- Both pure cultures and bacterial consortiums can be used
lular enzymes perform a major role in fungal adaptation and for microplastic degradation. However, pure cultures pre-
detoxification processes (Schwartz et al. 2018). The enzyme sent several advantages in the degradation process, offering
systems associated with cytochrome P450 family epoxidases a convenient way to study metabolic pathways involved in
and transferases are associated with oxidation and conjuga- the process. Moreover, the impact of environmental fac-
tion reactions and help in the metabolism of aliphatic, alicy- tors such as temperature, pH, substrate characteristics, and
clic, and aromatic molecules. They perform a wide range of surfactants affecting the degradation process can be more
reactions such as epoxidation, sulfoxidation, desulfuration, easily monitored (Janssen et al. 2002). However, the main
dehalogenation, deamination, and epoxidation (Shin et al. disadvantage is an extremely slow rate of degradation. Thus,
2018). The cytochrome P450 families of enzymes help in more innovative methods are required to optimize conditions

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 Table 1  Microplastic degradation by fungi
Source of microbes Isolated fungal strains Type of microplastic Pretreatment Incubation period % of degradation Enzymes References
degraded

Not reported Aspergillus sp. Penicil- Polypropylene/butylene- 30 days De Oliveira et al. ( 2020a,
lium sp. adipate-co-terephthalate b)
Marine sediments Zalerion maritimum Polyethylene pellets 28 days Paço et al. (2017)
Not reported Bjerkandera adusta Polypropylene and bio- Gamma irradiated Ligninase Butnaru et al. (2016)
mass
Marine coastal area Aspergillus flavus High-density polyethylene 30 days Devi et al. (2015)
Environmental Chemistry Letters (2023) 21:1787–1810

VRKPT2
Endophytes of Humboldtia Aspergillus sp. Paecilomy- Polypropylene Laccase Sheik et al. (2015)
brunonis, Psychotria ces Lilacinus, Lasiodip-
flavida lodia theobromae
Waste dump Aspergillus niger, Asper- Low-density polyethylene UV-irradiated 28 °C and relative 24%, 60% and Nowak et al. (2012)
gillus terreus, Aureoba- humidity of > 90% for 58% of its initial
sidium pullulans, 84 days mass
Paecilomyces varioti,
Penicillium funiculosum,
Penicillium ochrochlo-
ron, Scopulariopsis
brevicaulis, Trichoderma
viride
Soil, wall paint coated Fusarium solani, Spicaria Polyester polyurethane 100% Ibrahim et al. (2011)
with polyurethane spp., Alternaria solani,
and Aspergillus flavus
Culture collection Penicillium pinophilum Low-density polyethylene 31 months Volke-Sepúlveda et al.
ATCC 11,797 powder (2002)
Not reported Aspergillus niger, Low-density polyethylene 28 days Chandra and Rustgi (1997)
Penicillium funicu-
losum, Chaetomium
globosum, Gliocladium
virens and Pullularia
pullulans

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1796 Environmental Chemistry Letters (2023) 21:1787–1810

and improve the degrading bacterial isolates to shorten the polyethylene terephthalate, polypropylene, and polystyrene,
degradation process. The use of a consortium of bacteria is respectively.
usually preferred as it has been shown that biodegradation Few other bacteria associated with polypropylene deg-
by a single bacterium often results in the generation of toxic radation included Bacillus, Pseudomonas, Chelatococcus,
end products (Dobretsov et al. 2013), which can be success- and Lysinibacillus fusiformis, which were obtained from a
fully eliminated in a stable microbial community (Singh and wide variety of habitats including mangrove habitat, com-
Wahid 2015). post, cow dung, and land contaminated with plastic wastes.
The main process of degradation is represented by phys- Gut microflora of several arthropods like Tenebrio
icochemical degradation which reduces the polymer length molitor (mealworms) (Yang et al. 2015), Plodia interpunc-
and alters the functional groups of microplastics, making tella (Indian meal moth) (Yang et al. 2014) and Galleria
them more susceptible to microbial enzyme activity. Bio- mellonella (wax moths) (Kong et al. 2019) have also been
degradation using enzymes involves the action of lipases, reported to harbor microbes having microplastics biodeg-
esterases, laccases, amidases, cutinases, hydrolases, and radation properties. A study by Yang et al. (2014) isolated
carboxylesterases (Barth et al. 2016; Chen et al. 2020; two bacteria, Enterobacter asburiae YT1 and Bacillus sp.
Amobonye et al. 2021; Inderthal et al. 2021; Gómez-Mén- YP1, from the gut of waxworms capable of degrading poly-
dez et al. 2018). Thus, in-depth knowledge of the metabolic ethylene by decreasing the hydrophobicity and damaging
pathways and associated enzymes is necessary to perform the surface of polypropylene. A later study conducted by
an efficient biodegradation process. Yang et al. (2015) isolated a bacterial strain Exiguobacte-
Physiochemical pre-treatment, including chemical oxi- rium sp. from the guts of mealworms able to form biofilm
dizing agents, thermooxidation, and UV irradiation of and degrade polystyrene.
microplastics, is recommended to promote plastic biodeg- Efficient biodegradation of low-density polyethylene was
radation. These pre-treatments include UV irradiation, obtained using strains like Microbacterium paraoxydans and
nitric acid treatment and blending with polymers like starch Pseudomonas aeruginosa, which showed nearly 61.0% and
derivatives, cellulosic esters, polyhydroxybutyrate, poly(3- 50.5% degradation, respectively, within 2 months of incuba-
hydroxybutyrate-co-3-hydroxyvalerate), and polycaprolac- tion (Rajandas et al. 2012). Similarly, the biofilm of Pseu-
tone, which increase the biodegradability of polypropylene domonas sp. AKS2 has been reported to degrade low-density
(Gironi and Piemonte 2011). The addition of prooxidative polyethylene up to 5 ± 1% within an incubation period of
and biodegradable substances like starch to low-density 45 days (Tribedi and Sil 2013) without any pretreatment.
polyethylene, high-density polyethylene, polyvinyl alcohol, Likewise, degradation of polyethylene was also reported by
and polystyrene has been reported to enhance their biodegra- isolating Rhodococcus ruber C208 at the rate of 0.86% per
dability, promoting amylase activity (Zadjelovic et al. 2020). week (Sivan et al. 2006).
The earliest study of microplastics biodegrading micro- A consortium of bacteria consisting of Bacillus sp. and
organisms was conducted by Cacciari et al. (1993), using Paenibacillus sp. was able to reduce the dry weight of micro-
a consortium of Pseudomonas chlororaphis, Pseudomonas plastics by 14.7% in 60 days (Park and Kim 2019). Moreo-
stutzeri, and Vibrio sp. to degrade polypropylene. In the ver, Huerta Lwanga et al. (2018) investigated the earthworm
same study, the addition of starch was also reported to (Lumbricus terrestris)-mediated degradation of low-density
increase the biodegradation ability. polyethylene. The isolates from the gut included genera Act-
Later on, both Arkatkar et al. and Fontanella et al. inobacteria and Firmicutes which were also studied sepa-
reported biodegradation of polypropylene using a consor- rately and observed to be able to degrade low-density poly-
tium of Bacillus subtilis, B. flexus, Pseudomonas stutzeri, ethylene microplastics and release volatile compounds like
and Rhodococcus rhodochrous, respectively (Arkatkar et al. eicosane, docosane, and tricosane. A consortium of Entero-
2010; Fontanella et al. 2013). These microbial isolates were bacter and Pseudomonas from cow dung enhanced weight
found to form a biofilm, as reported in the study of Kowalc- loss up to 15% within 120 days (Skariyachan et al. 2021).
zyk et al. (2016), by isolating Achromobacter xylosoxidans. Several marine hydrocarbonoclastic bacteria such as
In a study conducted by Auta et al. (2018), two bacterial Alcanivorax borkumensis showed the ability to degrade
strains, belonging to Bacillus and Rhodococcus, isolated alkyl cycloalkanes, isoprenoid hydrocarbons, alkanes,
from mangrove sediments, showed polypropylene degrada- and branched aliphatic compounds (Davoodi et al. 2020).
tion efficiency of 4.0 and 6.4% after 40 days of incubation, The research was done on the same isolate that previously
respectively. They also reported that Bacillus cereus and showed biofilm formation on low-density polyethylene in the
Bacillus gottheilii were able to degrade microplastics (Auta presence of pyruvate, hexadecane and yeast extract and the
et al. 2017). B. gottheilii induced microplastics weight loss low-density polyethylene films (Delacuvellerie et al. 2019).
percentages of 6.2%, 3.0%, 3.6%, and 5.8% for polyethylene,

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It was also stated that the presence of alkanes modifies the monomers. Other bacterial strains can be genetically engi-
cell membrane hydrophilicity and produces biosurfactants neered by cloning the encoding genes of Ideonella sakaien-
to interact with the plastic surface and the formation of sis 201-F6, promoting the generation of modified strains
COOH/OH and C=O functional groups. Several actinomy- able to degrade polyethylene terephthalate in non-hazardous
cetes including Rhodococcus ruber and Streptomyces were monomers. Moog et al. (2019) introduced polyethylene tere-
also involved in polyethylene biodegradation (Sivan 2011). phthalate-hydrolyzing enzymes into photosynthetic micro-
Overall, among the different genera of bacteria asso- alga, Phaeodactylum tricornutum which showed efficient
ciated with microplastic degradation, 21% belonged to polyethylene terephthalate hydrolyzing activity.
Pseudomonas, about 15% to Bacillus and 17% derived Genetic modifications have also been made to promote
from mixtures of these two genera (Matjašič et al. 2021). the capture of polyvinyl chloride within the bacterial biofilm
Other bacteria associated with microplastic biodegrada- (Liu et al. 2021). Pseudomonas aeruginosa was genetically
tion included Enterobacter asburiae, Bacillus sp., Nocar- engineered by deleting the wspF gene to increase the for-
dia asteroids, Rhodococcus rhodochrous (Bonhomme et al. mation of sticky exopolymeric substances which enhance
2003), Streptomyces badius, Rhodococcus ruber, Coma- its capacity to accumulate microplastics in its biofilm.
monas acidovorans and Clostridium thermocellum (Paço Moreover, yhjH gene was designed under the control of
et al. 2019), Exiguobacterium sp., Ideonella sakaiensis an arabinose-induced promoter and was introduced into
(Tanasupawat et al. 2016), Pseudomonas chlororaphis, the bacterium. Since the function of yhjH was to decrease
Pseudomonas putida AJ, and Thermomonospora fusca cyclic dimeric guanosine monophosphate levels, induced
(Ghosh et al. 2013). A detailed list of different bacteria and expression of the gene reduced the biofilm formation suit-
actinomycetes associated with microplastic degradation is able enough to release captured microplastics. The synthetic
reported in Tables 2 and 3. ‘capture and release’ system would enable the creation of
To summarize, bacteria capable of degrading microplas- efficient microplastics scavengers for the bioremediation of
tics have been isolated from a wide range of habitats includ- aquatic ecosystems.
ing contaminated sediments, wastewater, sludge, compost, The advent of different genetic engineering methods has
municipal landfills, extreme environments, and microbiota. enabled us to manipulate the genetic materials of microbes
Bacteria have been tested for microplastics degradation, and enhance their biodegrading efficiency. Several proce-
both using pure cultures and microbial consortium. Bacte- dures involving recombinant DNA technology, gene clon-
rial consortium, in particular, show greater efficiency and ing, and genetic modification have been done to improve the
community stability. bioremediation ability of the microbes in presence of dif-
ferent hydrocarbons and heavy metals (Kumar et al. 2020).
However, till now very few works have been conducted on
Modern biotechnological methods the application of genetic engineering for creating a better
to enhance microplastic degradation strain for degradation of plastics.
These techniques are used for the construction of novel
Microplastics are gradually gaining attention due to their pathways and can alter enzyme specificity and their affinity
ubiquitous presence and negative impact on human health toward different microplastics. For successful gene editing,
and the ecosystem. Microplastic exposure has increased it is necessary to find suitable genes required for metaboliz-
many folds after the coronavirus disease 2019 pandemic due ing and degrading microplastics and suitable host organ-
to the excessive use of single-use personal protective equip- isms like E. coli in which these genes are expressed. The
ment products (Anand et al. 2022; De-la-Torre and Aragaw main processes involved are polymerase chain reaction, anti-
2021; Yang et al. 2022). Microplastic exposure may occur sense ribonucleic acid (RNA) technology, and site-directed
through inhalation, digestion, and dermal absorption and mutagenesis. Antisense RNA technology has emerged as a
might cause health issues like neurotoxicity, disrupt endo- new tool for genetic editing as artificially synthesized anti-
crine system, carcinogenicity, and metabolic disruptions sense RNA can effectively regulate the expression of genes
(Naqash et al. 2020; Rahman et al. 2021; see also Section in host cells. On the other hand, site-directed mutagenesis
“Processes of microplastic degradation”). Thus, there is a is also used to alter the activity of genes associated with
need for efficient microplastic bioremediation. microplastic degradation. In a recent study, Lameh et al.
Polyethylene terephthalate is a widely used thermoplastic (2022) reported mutation of carboxylesterase by in silico
polymer used for packaging which can degrade and generate site‐directed mutagenesis to produce BTA‐hydrolase in
microplastics. Isolates like, Ideonella sakaiensis 201-F6 are Archaeoglobus fulgidus to enhance its ability to degrade
reported to produce polyethylene terephthalate-hydrolyzing polyethylene terephthalate.
enzymes able to degrade polyethylene terephthalate to tere- The main enzymes associated with microplastic deg-
phthalic acid, and ethylene glycol which is non-hazardous radation, such as manganese-dependent peroxidase, were

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 Table 2  Microplastic degradation by bacterial isolates
1798

Source of microbes Isolated bacterial Type of microplastic Incubation % of degradation Biodegradation detection method/ References
strains degraded period techniques

13
Polluted soil samples Lysinibacillus sp. Polypropylene, polyeth- 26 days 4 and 9% Gas chromatography – mass Jeon et al.
ylene spectrometry, Scanning electron (2021)
microscopy
Cow dung sample Enterobacter sp nov. bt Low-density polyethylene, 160 days 64.25 ± 2% and 63.00 ± 2% Weight loss Skariyachan
DSCE01, Enterobac- polypropylene et al. (2021)
ter cloacae nov. bt
DSCE02, and Pseu-
domonas aeruginosa
nov. bt DSCE-CD03
Compost Bacillus cereus, Bacil- Polypropylene and poly- 6 months Fourier-transform infrared spectros- Jain et al.
lus thuringiensis, L-lactide copy; Thermogravimetric analysis (2022)
Bacillus licheni-
formis
Municipal landfill sediment Bacillus sp. and Paeni- Polyethylene 60 days 14.7 % Field-emission scanning electron Park and Kim
bacillus sp. microscope, Fourier transform (2019)
infrared spectrometer, Gas
chromatography-mass spectrome-
ter, Scanning electron microscopy,
Thermogravimetric analyzer
Biofilm composed Polyethylene, polypro- NA DNA extraction, amplification and Miao et al.
by Pirellulaceae, pylene sequencing (evaluation of the (2019)
Phycisphaerales, effects of substrate type on micro-
Cyclobacteriaceae, bial communities)
and Roseococcus
Earthworm gut Bacillus simplex and Low-density polyethylene 21 days Scanning electron microscopy Huerta Lwanga
Bacillus sp. et al. (2018)
Mangrove sediments Bacillus sp. strain 27 Polypropylene 40 days 4.0% Weight loss; Fourier-transform Auta et al.
infrared spectroscopy; Scanning (2018)
electron microscopy
Mangrove sediment Bacillus gottheilii Polyethylene, polyeth- 40 days 6.2%, 3.0%, 3.6%, 5.8% Weight loss; Fourier-transform Auta et al.
ylene terephthalate, infrared spectroscopy; Scanning (2017)
polypropylene, and electron microscopy
polystyrene
Bacillus cereus Polyethylene, polyeth- 40 days 1.6%, 6.6%, and 7.4%
ylene terephthalate,
polystyrene
Compost Bacillus thuringiensis Polypropylene and poly- 15 days 12% Fourier-transform infrared spectros- Jain et al.
L-lactide copy; Scanning electron micros- (2018)
copy; Thermogravimetric analysis
Compost Bacillus licheniformis Polypropylene and poly- 15 days 10% Fourier-transform infrared spectros- Jain et al.
L-lactide copy; Scanning electron micros- (2018)
copy; Thermogravimetric analysis
Environmental Chemistry Letters (2023) 21:1787–1810
 Table 2  (continued)
Source of microbes Isolated bacterial Type of microplastic Incubation % of degradation Biodegradation detection method/ References
strains degraded period techniques

Sewage treatment plants (STP) Microbial consortia Low density polyethylene, 140 days 47%, 58% and 56% Fourier-transform infrared spectros- Skariyachan
(including Aneu- high density polyethyl- copy; Scanning electron micros- et al. (2018)
rinibacillus sp. and ene and polypropylene copy; Atomic force microscopy;
Brevibacillus sp.) Energy dispersive spectroscopy;
Nuclear magnetic resonance; Gas
chromatography-mass spectrom-
etry
Mangrove sediments in Penin- Bacillus cereus Polypropylene 40 days 12% Weight loss Helen et al.
sular Malaysia (2017)
Mangrove sediments in Penin- Sporosarcina globis- Polypropylene 40 days 11% Weight loss Helen et al.
sular Malaysia pora (2017)
Sandy beaches in Northern Agios Onoufrios and Polystyrene films 6 months 0.19% Weight loss; Fourier-transform Syranidou et al.
Environmental Chemistry Letters (2023) 21:1787–1810

Crete, Chania, Greece Kalathas infrared spectroscopy; Scanning (2017)

electron microscopy
Pseudomonas aerugi- Polyurethanes 2.5% and 2.4 % Weight loss; Scanning electron Uscátegui et al.
nosa and Escherichia microscopy; Tensile strength and (2016)
coli elongation at break
Municipal solid waste Stenotrophomonas Polypropylene 90 days 20.3 ± 1.39% Weight loss; Fourier-transform infra- Jeon and Kim
panacihumi PA3-2 red spectroscopy (2016)
Nitrosomonas sp., High density polyethyl- 90 days 15%-20% (High density poly- Weight loss Muenmee et al.
Nitrobacter sp., ene, low density poly- ethylene), 5%-9% (polypro- (2016)
Burkholderia sp. and ethylene, polypropylene pylene),
Pseudomonas sp.
Plastic-eating mealworms Exiguobacterium sp. Polystyrene 60 days 7.4% ± 0.4% Cross polarization—magic angle Yang et al.
strain YT2 spinning nuclear magnetic reso- (2015)
nance; Thermogravimetric analysis
coupled with fourier-transform
infrared spectroscopy
National Environmental Engi- B. flexus + P. azotofor- UV treated polymers 12 months 22.7% Weight loss; Fourier-transform infra- Aravinthan
neering Research Institute, mans red spectroscopy et al. (2016)
Nagpur India
Plastic-eating waxworms gut Bacillus sp. ­YP1 Polyethylene films 28 days 10.7 ± 0.2% Weight loss; Fourier-transform infra- Yang et al.
Enterobacter asburiae 6.1 ± 0.3% red spectroscopy (2014)
­YT1
Compost Chelatococcus sp. E1 Low-molecular-weight 80 days 44.5% Gel permeation chromatogra- Jeon and Kim
polyethylene phy; Fourier-transform infrared (2013)
spectroscopy; Nuclear magnetic
resonance; Tensile strength

13
1799
 1800

13
Table 3  Microplastic degradation by actinomycetes
Source of microbes Isolated actinomycetes strains Type of microplastic degraded Incubation period % of degradation Biodegradation detection References
Method/techniques

Antarctic soil Pseudomonas sp. ADL15 and Polypropylene 40 days 17.3% and 7.3% Weight loss; Fourier-transform Habib et al. (2020)
Rhodococcus sp. ADL36 infrared spectroscopy
Not reported Rhodococcus ruber strain Polyethylene 2 months 7.5% Weight loss; Scanning electron Sivan et al. (2006)
C208 microscopy
Soils from waste coal, a for- Bacterial consortia Low-density polyethylene 225 days 17.03% Weight loss; Fourier-transform Nowak et al. (2011)
est and an extinct volcano Arthrobacter viscosus, infrared spectroscopy; Scan-
crater Micrococcus lylae, ning electron microscopy;
Micrococcus luteus, elongation at brake
Bacillus mycoides, Bacillus
cereus, Bacillus pumilus;
Bacillus thuringiensis
Not reported Microbacterium paraoxydans Polyethylene (pre-treated with 2 months 61.0% Weight loss; Fourier-transform Rajandas et al. (2012)
nitric acid) infrared spectroscopy
Mangrove sediment Rhodococcus Polypropylene 40 days 6.4% Weight loss; Fourier-transform Auta et al. (2018)
infrared spectroscopy; Scan-
ning electron microscopy
Actinomadura sp. T16-1 Polylactic acid (production of 96 h Not available Enzyme activity Sukkhum et al. (2009)
(Enzyme production) polylactic acid-degrading
enzyme)
Rhodococcus ruber Polystyrene 2 months 0.8% Weight loss Mor and Sivan (2008)
Rhodococcus rhodochrous Two polypropylene films (Sta- 6 months Not available Fourier-transform infrared Fontanella et al. (2013)
ATCC 29,672 tistical copolymer and block spectroscopy; Proton nuclear
copolymer) magnetic resonance; ADP/
ATP ratio
Environmental Chemistry Letters (2023) 21:1787–1810
Environmental Chemistry Letters (2023) 21:1787–1810 1801

produced by a genetically engineered strain of E. coli and S. microplastic particles (Purohit et al. 2020). Various types of
cerevisiae BY 4741, similarly, laccase enzymes were pro- databases such as The University of Minnesota Biocatalysis/
duced by a modified genetic strain of E. coli BL21 and P. Biodegradation Database, The Environmental Contaminant
chrysosporium (Sharma et al. 2018; Paço et al. 2019). These Biotransformation Pathway Resource, MetaCyc database,
genetically modified enzymes are capable of better degrada- and BioCyc database related to biodegradation pathways
tion of polyethylene terephthalate. have been established to evaluate the process of biodegra-
Enzyme cutinase produced by microbes for the break- dation by providing information on the metabolic pathways,
down of polyester linkage can also be used in polyethylene the microbial enzymes and genes associated with the process
terephthalate degradation which acts at an optimal tempera- (Gao et al. 2010; Wicker et al. 2016; Karp et al. 2019; Caspi
ture of 75 °C. Genetic engineered yeast produces bacterial et al. 2020). These databases and computational methods
cutinase which prevents formation in strategic positions with help to recognize enzymes involved in a metabolic pathway
sugars which helps in the degradation of polyethylene tere- of interest and help in forecasting the biodegradation routes
phthalate (Shirke et al. 2018). Islam et al. (2019) reported of toxic chemicals, providing a platform in which a novel
that genetically engineered cutinase enzyme reduces the approach for the biodegradation of plastic can be designed
degradation time from 41.8 to 6.2 h when compared with (Ali et al. 2021).
wild strain. Despite all these advantages, the major disadvantage
A similar enhanced biodegradation ability was observed associated with bioinformatics is the lack of experimen-
in a consortium of marine microbes (Syranidou et al. 2019). tal data and its validation which is required for future
However, despite their better ability in laboratory conditions, research. Moreover, there is a wide knowledge gap
most of these genetically modified organisms have displayed between diverse groups of synthetic polymer degrader
unsatisfactory results in field studies. microorganisms and their responsible enzymes. Hence,
an extensive investigation is required to identify suitable
metabolic pathways for the degradation of polymers and
Gene editing tools their associated enzymes. In the near future, a combina-
tion of approaches, using bioinformatic tools, metabolic
Gene editing tools have been applied for genome engineer- engineering, genetics, molecular, and system biology may
ing of plants, animals, and microorganisms for the expres- help us to find a suitable and sustainable option for the
sion of specific genes (Paço et al. 2019; Tang et al. 2020; biodegradation of microplastics.
Nidhi et al. 2021; Ozyigit et al. 2021; Bhattacharyya et al.
2022; Biswas et al. 2022; Mandal et al. 2022; Jiang et al.
2022). With the advent of different types of gene editing Biosafety issues associated with genetically
tools such as zinc finger proteins, transcription activator-like modified organisms
effector nucleases, and more recently, the clustered regu-
larly interspaced palindromic repeats (CRISPR)/Cas9, the The advent of different genetic engineering techniques,
manipulation of organisms has become easier (Jiang et al. and synthetic biological and genetic tools has allowed
2013; Gaj et al. 2013). Genome editing also helps in the the development of genetically modified microbial scav-
manipulation of a gene of interest which can perform the engers for the mitigation of diverse types of pollutants
loss and gain of function experiments which alter the expres- (Mohamed et al. 2020; Wang et al. 2019a, b). However,
sion of different genes. several regulatory hurdles hamper the use of genetically
This strategy can be efficiently used to incorporate genes modified microorganisms in an onsite experiment. Also,
encoding enzymes like polyethylene terephthalate hydro- these genetically modified microbes have shown their effi-
lase, dehalogenase, esterase, depolymerase, and laccase ciency in laboratory conditions, but onsite experiments are
which are associated with microplastic degradation. Three required to validate their effectiveness.
different CRISPR sequences were identified in Streptomyces A diverse range of genetic tools has been developed
albogriseolus LBX-2 which makes it a suitable organism for only to prevent the negative impacts of genetically modi-
genetic engineering, where the main enzyme associated with fied organisms on the field, including antibiotic gene-
polyethylene degradation is oxygenase (Shao et al. 2019). free genetic engineering tools (Ji et al. 2019) and suicide
genetic systems (Honjo et al. 2019; Marguet et al. 2010;
Scott et al. 2017). By using synthetic biology and meta-
Bioinformatics bolic engineering, an attempt has been already made to
engineer microorganisms, which can be efficiently used as
Bioinformatics has also become an effective tool for self-eliminate microbial scavengers for the bioremediation
enhancing the biodegradation of plastic debris including

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1802 Environmental Chemistry Letters (2023) 21:1787–1810

of various toxic environmental pollutants (Moog et al. References

2019; French et al. 2020).
These techniques look promising and can be applied in Ali SS, Elsamahy T, Koutra E, Kornaros M, El-Sheekh M, Abdelkarim
the case of microplastics bioremediation. For biomedical EA, Sun J (2021) Degradation of conventional plastic wastes in
the environment: a review on current status of knowledge and
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Authors contributions All authors contributed to drafting and criti- lus sp. and Rhodococcus sp. isolated from mangrove sediment.
cal revision of the article and approved the article before submission. Mar Pollut Bull 127:15–21. https://​doi.​org/​10.​1016/j.​marpo​lbul.​
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Funding Not applicable. Danso D, Chow J, Streit WR (2019) Plastics: environmental and bio-
technological perspectives on microbial degradation. Appl Envi-
Declarations ron Microbiol. https://​doi.​org/​10.​1128/​AEM.​01095-​19
Awasthi AK, Tan Q, Li J (2020) Biotechnological potential for micro-
Conflicts of interest The authors declared no potential conflicts of in- plastic waste. Trends Biotechnol 38(11):1196–1199. https://​doi.​
terest. org/​10.​1016/j.​tibte​ch.​2020.​03.​002

Consent for publication All authors agreed on the publication.

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