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Potential strategies for bioremediation of microplastic contaminated soil

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DOI: 10.1016/j.enceco.2024.05.001

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 Environmental Chemistry and Ecotoxicology 6 (2024) 117–131

Contents lists available at ScienceDirect

Environmental Chemistry and Ecotoxicology

journal homepage: www.keaipublishing.com/en/journals/environmental-
chemistry-and-ecotoxicology/

Potential strategies for bioremediation of microplastic contaminated soil

Charu Thapliyal a, Anshu Priya b, Salam Bhopen Singh a, Vivekanand Bahuguna a,
Achlesh Daverey a, c, *
a
School of Biological Sciences, Doon University, Dehradun 248012, Uttarakhand, India
b
Amity Institute of Biotechnology, Amity University, Uttar Pradesh, Sector –125, Noida 201303, India
c
School of Environment and Natural Resources, Doon University, Dehradun 248012, Uttarakhand, India

A R T I C L E I N F O A B S T R A C T

Keywords: The escalating production and ubiquitous presence of plastics and their degradation products, such as micro­
Microplastic plastics and nanoplastics, pose a significant environmental threat. Microplastics enter the soil through various
Soil pollution pathways, including agricultural practices, plastic degradation, and wastewater disposal. Herein, we discussed
Bioremediation
the harmful effects of microplastics on the physicochemical properties of soil, plant growth, terrestrial fauna, and
Biodegradation
microbial activity, potentially affecting the stability and nutrient cycle of the soil ecosystem. This review delves
Phytoremediation
Synthetically engineered microbes into recent advances in potential microplastic bioremediation approaches, such as phytoremediation strategies
utilized by plants and their associated microbes to accumulate, immobilize, and even degrade microplastics.
Rhizosphere microorganisms play a crucial role in the degradation of microplastics, potentially utilizing them as
a carbon source. Soil animals like earthworms, snails, and mealworms can also contribute significantly to
bioremediation by ingesting and degrading microplastics through their gut microbiota. Various soil microor­
ganisms, including bacteria and fungi, can degrade different microplastics with the help of enzymes such as
laccase, esterase, peroxidase, oxidoreductase, and hydrolases and depolymerise the larger polymer chains into
smaller units that ultimately mineralize them into CO2, H2O, and CH4. Genetic engineering and synthetic biology
are also used to create strains with enhanced microplastic degrading and mineralization capabilities. It holds
promise for efficient bioremediation but requires further research for real-world application and scalable
implementation. Overall, this review comprehensively highlights the potential of bioremediation approaches and
future recommendations for tackling microplastic pollution. Further research and development are crucial for
enhancing biodegradation efficiency and scaling up this strategy for environmental protection.

1. Introduction culminating in an estimated 12 billion tons of plastic waste anticipated

to accumulate in terrestrial ecosystems by 2050 [6]. This ominous
The escalating global production of plastic materials, driven by their projection underscores the confluence of inadequate disposal practices,
versatile utility, cost-effectiveness, and durable properties, has engen­ protracted degradation rates, and suboptimal recycling initiatives.
dered significant environmental concerns, particularly exacerbated by The gravity of plastic pollution is underscored by the explicit
the compounding factors of population expansion and rapid industrial­ declaration from the United Nations Environment Programme (UNEP),
ization [1,2]. In 2018, plastic production reached 360 million tons, acknowledging it as an eminent environmental crisis [6]. The intricate
rising to 370 million tons in 2019 and 390 million tons in 2021 [3,4]. interplay between plastics and the environment unfolds through the
Projections indicate a trajectory towards 450 million tons by 2025 and a actions of both biotic and abiotic agents, including UV radiation, tem­
staggering 1480 million tons by 2050 [5]. This substantial surge in perature variations, moisture, collision, hydrolysis, abrasion, and ther­
plastic production underscores a pivotal environmental issue, mal oxidation [7,8]. This intricate choreography results in the

Abbreviations: PLA, Polylactic acid; FTIR, Fourier-transform infrared; HDPE, High -density polyethylene; HHCB, 1,3,4,6,7,8-hexahydro-4,6,6,7,8,8,-hexamethyl-
cyclopenta[g]benzopyran; LDPE, Low -density polyethylene; LDPP, Low -density polypropylene; MP, Microplastics; NP, Nanoplastics; PE, Polyethylene; PET,
Polyethylene terephthalate; PS, Polystyrene; PU, Polyurethane; PVC, Polyvinyl chloride; SEM, Scanning electron microscopy; TCA, Tricarboxylic acid cycle; UNEP,
United Nations Environment Programme.
* Corresponding author at: School of Environment and Natural Resources, Doon University, Dehradun, Uttarakhand 248012, India.
E-mail address: [email protected] (A. Daverey).

https://doi.org/10.1016/j.enceco.2024.05.001
Received 23 January 2024; Received in revised form 2 April 2024; Accepted 4 May 2024
Available online 10 May 2024
2590-1826/© 2023 The Authors. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co. Ltd. CC BY-NC-ND 4.0 This is an open access article
under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
C. Thapliyal et al. Environmental Chemistry and Ecotoxicology 6 (2024) 117–131

fragmentation of plastics into macroplastics (>5 mm), microplastics MP are often hydrophobic, that affect their interactions with soil water
(MP) (1 μm to 5 mm), and nanoplastics (NP) (<1 μm), with the latter content and organic matter, potentially influencing their transport and
category, MP, assuming particular significance due to heightened retention in soil. MP have the potential to adsorb various chemicals from
toxicity and enhanced environmental mobility [6]. MP pollution rep­ the surrounding environment. The presence of adsorbed substances on
resents a notable environmental challenge on a global scale, particularly MP surfaces may alter their interactions with soil particles and influence
across Asia, China, India, Indonesia, Philippines, Vietnam, Thailand, and their distribution in soil. The combined effects of these physicochemical
Bangladesh contend with significant plastic contamination within their attributes determine the behavior of MP in soil environments, including
soil ecosystems. This issue stems from various factors such as the their transport, retention, and potential impacts on soil ecosystems. Of
widespread adoption of plastic mulching in agricultural practices, notable concern is the discernible prevalence of MP in soil, and eclipsing
inadequate waste management strategies, and the rapid pace of indus­ quantities observed in marine environments. An overwhelming 80% of
trialization. The presence of MP particles in soils can alter soil charac­ marine MP trace their origin from terrestrial sources [18,19]. This ac­
teristics and diminish soil fertility, affect microbial populations and centuates the role of soil as a substantial repository, functioning as a
diversity, and impede crop development. Agricultural areas situated critical sink for MP in the global ecosystem. The fate of MP within soil
proximate to urban centres, bodies of water, and coastal zones are matrices is contingent upon several factors, including the type of plastic
especially susceptible to MP pollution. The ramifications of MP monomer (e.g., polyethylene (PE), polypropylene (PP), polystyrene
contamination extend to diminish soil quality, poor ecosystem services, (PS)) and their intricate interplay with soil characteristics, encompass­
and low agricultural output, underscoring the urgent need for mitigation ing shape, size, charge, density, texture, composition, hydrophobicity,
efforts [9–11]. Over temporal scales, the properties of MP encompassing and adsorptivity [20,21]. Polyethylene terephthalates (PET) are poly­
morphology, mechanical strength, and functional groups undergo dy­ ester molecules that typically undergo degradation primarily through
namic modifications in response to key physiochemical parameters such hydrolytic cleavage and photo-oxidation under ambient environmental
as polymer type, size distribution, environmental conditions, soil conditions [22]. While, Polyethylene (PE) is characterized as an inert
composition, aging and weathering, biological interactions, anthropo­ polyolefin that exhibits gradual degradation in natural environments.
genic inputs etc. [12,13]. Notably, an elevated rate of photo-oxidation has been observed in LDPE
MP ingress into the environment through direct and indirect compared to HDPE, attributed to the greater occurrence of reactive
anthropogenic activities, spanning agriculture, packaging, cosmetic branch points within the low-density polymer [23]. Facilitated by larger
applications, industrial processes, and waste disposal practices [14,15]. pore sizes, MP exhibit heightened vertical infiltration in sandy soils as
These MP are categorized based on their polymer composition, which compared to plastic fibres and films, accentuating their ubiquity and
predominantly includes polyethylene (PE), polypropylene (PP), poly­ pervasive distribution [24,25]. The ramifications of this widespread MP
ethylene terephthalate (PET), polyvinyl chloride (PVC), polystyrene distribution pose a global concern, manifesting in deleterious impacts on
(PS), polyamide (PA) and polyethylene-terephthalate glycol (PETG). the physicochemical properties of soil and perturb the functionality of
Each of these polymers exhibits distinct chemical structures and bonding soil microorganisms, plants, and animals in various ways, including-
configurations. PE, PP and PVC are composed primarily of carbon and disruption of microbial community, toxicity to microorganisms, bio­
hydrogen atoms linked by single covalent bonds, forming linear or accumulation in plants, alteration of soil physical properties, impair­
branched chains. PET consists of repeating units of ethylene glycol and ment of nutrients, affect plant growth, physical damage to soil fauna,
terephthalic acid, connected by ester bonds. PVC consists of vinyl and disruption of soil food webs [26]. Prior studies delve into various
chloride monomers linked by covalent bonds, forming a linear or mechanisms within the domain of MP bioremediation. Certain micro­
branched structure. PS comprises of styrene monomers linked by cova­ organisms exhibit enzymatic capabilities that facilitate the breakdown
lent bonds, forming a linear polymer chain. PETG incorporates tere­ of specific polymer structures, thereby contributing to MP degradation
phthalate and ethylene glycol units joined by ester linkages, [27,28]. These enzymes serve as biological catalysts, breaking down the
contributing to its distinct polymer structure. The types of bonds present molecular structure of MP into smaller components including oligomers,
within these polymers dictate their chemical stability, degradation rates, monomers, CO, CO2, H2O, CH4, carboxylic acids, anhydrides, and esters.
environmental interactions, and consequently their persistence and These products are more readily assimilated by the microbial commu­
environmental impact as MP [16]. Various environmental factors play a nity as a source of carbon and energy. The microbes utilize these
pivotal role in the fragmentation of MP, fostering their enduring pres­ degraded MP as substrates for metabolic processes, incorporating them
ence in ecosystems and posing significant risks to both ecological sys­ into their cellular machinery for growth and proliferation [29]. This
tems and organisms therein. These factors encompass mechanical stress, phenomenon underscores the remarkable adaptability of microorgan­
UV radiation, temperature, pH, chemical weathering, biological activ­ isms to utilize novel carbon sources, including synthetic polymers like
ity, and the duration of exposure. The dispersion of MP in soil is subject MP, and highlights their potential role in mitigating plastic pollution in
to the influence of various physicochemical attributes, such as their the environment through biodegradation processes. Besides, certain
shape, size, electrical charge, density, texture, composition, hydropho­ plant species display the ability to absorb and accumulate MP within
bicity, and adsorption capacity, among others [12,13]. Smaller MP their tissues, offering a potential avenue for soil remediation strategies
generally have greater mobility and can penetrate deeper into soil [30,31].
layers, while larger ones may be more confined to surface soils [17]. Size There are significant international efforts aimed at mitigating plastic
also impacts the surface area available for interactions with soil particles and MP pollution that includes Basel Convention, which addresses the
and organisms. Irregularly shaped MP may have varied interactions with control of transboundary movements of hazardous waste, including
soil particles compared to spherical ones, potentially influencing their plastic waste [32]. Within the European Union, directives like the
distribution. The density of MP relative to soil particles influences their Single-Use Plastics Directive and the Circular Economy Action Plan are
settling behavior and vertical distribution within soil profiles. MP with pivotal in reducing plastic waste, thus indirectly tackling MP pollution.
densities similar to soil particles may exhibit different transport be­ Furthermore, regional agreements, like the Barcelona Convention and
haviors compared to those with contrasting densities. Different soil the OSPAR Convention, highlight regional efforts to protect marine
textures have varying pore sizes and structures, which can affect the environments from plastic pollution, including MP. Moreover, MP
movement and retention of MP. Fine-textured soils with smaller pores degradation significantly contributes to several Sustainable Develop­
may retain MP more effectively than coarse-textured soils. The chemical ment Goals (SDGs), including mitigating MP threats to marine life (SDG
composition of MP can also influence their interactions with soil com­ 14), promoting responsible consumption and production patterns (SDG
ponents. For instance, MP containing additives or surface coatings may 12), safeguarding water quality (SDG 6), protecting terrestrial ecosys­
exhibit different affinities for soil particles compared to pure polymers. tems (SDG 15), addressing health risks (SDG 3), and fostering innovation

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in materials science and waste management (SDG 9). India has imple­ development to enhance biodegradation efficiency and scale up these
mented a series of policies and regulations to address plastic and MP strategies for effective environmental protection. Moreover, the review
issues, notably through the Plastic Waste Management Rules (PWM) underscores the value of the assessment in synthesizing existing litera­
introduced in 2016. These rules regulate the production, usage, and ture on MP in soil, providing a comprehensive understanding for re­
disposal of plastic materials with a focus on environmentally sound searchers, policymakers, and stakeholders. The synthesis of these
management and safe disposal. Additionally, several states and union insights seeks to propel a more nuanced understanding of MP dynamics
territories have imposed bans on single-use plastics to curb pollution and in soil environments, fostering informed strategies for mitigation and
encourage alternatives. Various initiatives by governmental, non- environmental stewardship.
governmental, and private entities aim to promote plastic waste man­
agement, including awareness campaigns and recycling programs. The 2. Effects of microplastics on soil ecosystem
National Clean Air Programme (NCAP) underscores the importance of
improved waste management practices to mitigate air pollution, often Soil, being a complex mixture of minerals, moisture, air, and organic
exacerbated by plastic burning. Furthermore, there is a concerted effort and inorganic matters, plays a pivotal role in environmental conserva­
towards research and innovation for biodegradable plastics and alter­ tion and security. MP in soil induce significant alterations in crucial
native materials to lessen reliance on conventional plastics, involving physicochemical properties such as changes in soil porosity, bulk den­
government agencies, research institutions, and private companies [33]. sity, water-holding capacity, organic matter content, pH, electrical
Despite a heightened collective awareness regarding the behavior of conductivity, and adsorption capacity [34–36]. The impact of MP on the
MP, knowledge gaps persist concerning their transformations in soil and soil ecosystem is contingent upon various factors, including their type,
their intricate effects on soil ecosystems. In summation, this compre­ concentration, shape, size, and duration of exposure, emphasizing the
hensive review aims to elucidate the current state of knowledge on the need for a nuanced understanding of these parameters [37]. The lower
multifaceted impacts of MP on soil, ranging from physicochemical al­ density of MP compared to soil particles leads to a reduction in soil
terations to ecological consequences. The review highlights the pressing carbon storage, consequently affecting bulk density [36,38]. The hy­
environmental threat posed by the widespread presence of plastics and drophobic nature of MP diminishes soil aggregate stability, resulting in a
their degradation products, particularly MP, in soil ecosystems. Various decrease in water-stable aggregates and an increase in soil water evap­
pathways through which MP enter soil and their detrimental effects on oration, though effects can vary [39]. For instance, polyester fibres
soil physicochemical properties, plant growth, and microbial activity increased the water-holding capacity of soil, contrasting with poly­
have been discussed. The review emphasizes recent advances in biore­ ethylenes, polyamides, and polyacrylics fragments [1,39,40]. Studies
mediation strategies, including phytoremediation and genetically engi­ reveal diverse pH responses to different types of MP. For example, low-
neered microbial degradation, as promising approaches to mitigate MP density polypropylene (LDPP) elevates soil pH, while fibres, high-
pollution. It underscores the importance of further research and density polyethylene (HDPE), and biodegradable polylactic acid (PLA)

Fig. 1. Sources of MP in the environment and their effects on the physicochemical characteristics of soil, plants, soil animals, and microorganisms.

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reduce soil pH after one month of exposure [36,41,42]. Similarly, soil matter decomposition. Similarly, chlorinated paraffins, which are
electrical conductivity is also intricately related to the chemical nature widely used as flame retardants and plasticizers, can exert adverse ef­
of MP. Fig. 1 shows the sources of MP in the environment and their ef­ fects on microbial communities. These compounds are known for their
fects on the soil ecosystem. persistence in the environment and their toxic effects on microorgan­
MP, particularly low-density polyethylene (LDPE) and biodegrad­ isms. Chlorinated paraffins can interfere with microbial cell membrane
able plastics, can modify the soil carbon‑nitrogen ratio (C:N) over time, integrity, inhibit enzyme activities, and disrupt metabolic pathways. As
potentially due to carbon release during slow degradation [43]. Clothing a result, exposure to chlorinated paraffins often leads to a decrease in
fibres increase soil organic matter, while HDPE and PLA decrease it, microbial abundance and diversity, particularly among indigenous soil
with variations attributed to differences in biodegradation and carbon and aquatic microorganisms [59]. On the other hand, Bisphenol A
adsorption behaviors of the MP [34,44]. Moreover, MP affect soil quality (BPA), a common component of plastics and epoxy resins, can have
by suppressing soil enzymes activities including alkaline phosphatase, complex effects on microbial communities. While, some studies have
β-glucosidase, leucine-aminopeptidase, and dehydrogenase, and posing shown that BPA exposure can reduce microbial diversity by inhibiting
potential threats to the soil ecosystem [45]. the growth of Lysobacter, Steroidobacter, Variovorax, and Mycoplan [60].
The physicochemical characteristics of MP, including shape, size, Others have demonstrated that BPA can stimulate the growth of specific
polymer density, and chemical constituents, play a significant role in microbial populations such as Pseudomonas sp. and Arthrobacter sp.
shaping soil microbial activities and microbial diversity. Additionally, capable of metabolizing this compound as a carbon and energy source
MP undergo weathering processes through photooxidation and thermal [61]. Thus, the impact of BPA on microbial abundance and diversity
deterioration, leading to alterations in surface properties that influence may vary depending on environmental conditions, microbial commu­
microbial attachment and colonization [42,46,47]. MP undergo nity composition, and the availability of alternative carbon sources.
weathering processes, such as photooxidation and thermal deteriora­ The impact of MP on plants is intricately linked to its physico­
tion, leading to changes in surface properties that influence microbial chemical characteristics, influencing the vital biological processes such
attachment and colonization [48]. A complex network of microorgan­ as seed germination, growth, reproduction, photosynthesis, and defence
isms including bacteria, fungi, protozoa, and algae, forming a micro­ mechanisms [38,44,62,63]. Bosker et al. reported delayed seed germi­
plastisphere plays a crucial role in agroecosystem management [49]. nation and inhibited root growth in Lapidium sativum due to obstruction
Different types of MP selectively stimulate specific soil microbial taxa, of seed pores by MP [63]. Similarly, high concentrations of HDPE MP
impacting microbial richness and diversity on MP surfaces [50,51]. impeded seed germination, reduced shoot development, and biomass
Specific bacterial colonies form on MP surfaces, alter the microbial production in Lolium perenne [44]. The MP residues enter leguminous
composition in the surrounding environment. Certain genera of micro­ plants through root pores and interfered with stomatal functioning and
organisms known for degrading plastic polymeric substances such as nutrient transport. Jiang et al. highlighted oxidative damage, inhibition
Phenylobacterium, Arthrobacter and Streptomyces are found to colonize on of catalase enzyme activity, and genotoxic effects of PS MP on Vicia faba,
MP materials [52]. Phenylobacterium have potential to degrade poly­ resulting in reduced biomass production [37]. Thus, MP in the soil
cyclic aromatic hydrocarbons and use it to meet their growth and reduce growth, quality, and yield of the crops, affecting fertilizer effi­
respiration requirements [52] Arthrobacter were found to degrade pol­ cacy [64,65].
ycarbonate plastics forming polar by-products [53]. In a study on PE MP ubiquitously distributed in soil ecosystems enter animals,
film degradation, Streptomyces sp. demonstrated the ability to form including humans, through inhalation and the food chain. The physi­
biofilm on PE surface and degradative enzymes leading to alternation in cochemical properties of MP play a crucial role in determining their
the chemical structure of PE [54]. Owing to the ability to produce plastic harmful effects. In soil, MP serve as vectors, enhancing the bioavail­
degrading biochemicals and enzymes, these bacteria may be considered ability of several toxic pollutants, including heavy metals, by accumu­
as effective MP remediating and biodegrading tool. Studies report lating them on their surfaces [21,66]. Soil animals inadvertently ingest
varying effects of MP on soil microbial communities, with changes MP, leading to various toxic effects such as false satiety, energy deple­
observed in community structure, abundance, and diversity. The altered tion, hindered growth, and development [67]. MP ingestion also affects
physicochemical characteristics of soil due to the presence of MP, has the gut microbiota, metabolism, immunity, and fertility in soil animals
also been reported to affect arbuscular mycorrhizal fungi (AMF) growth [68–70]. Various soil animals, such as mites, moles, earthworms, go­
[55]. AMF are symbiotic organisms that form associations with the roots phers, and collembolan, contribute to the dispersion of MP in soil
of a wide range of plant species, playing essential roles in nutrient through surface attachment and movement, facilitating their transfer
cycling, soil structure maintenance, and overall plant health. Their across trophic levels in the food chain [71,72]. Earthworms, for
growth and colonization are influenced by environmental factors. AMF example, are affected by MP, leading to hindered growth, compromised
enhance plant resilience to cold stress by aiding in moisture retention, immune systems, and potential alterations in the carbon‑nitrogen ratio
increasing secondary metabolites, strengthening the immune system, (C:N) due to their contribution of mucus and excreta as nitrogen sources
and boosting protein content for cold stress adaptation. Moreover, [43,73]. Toxic mechanisms of MP on earthworms primarily involve
symbiotic AMF relationships improve water and plant interactions, gas histopathological damage and oxidative stress. When the concentration
exchange, and osmotic adjustment, while concurrently boosting chlo­ of MP in soil surpasses 0.1%, it affects earthworm growth and induces
rophyll synthesis, leading to elevated concentrations of metabolites in oxidative stress, leading to neural and DNA damage [74,75]. Similarly,
cold-stressed plants [56]. polyethylene terephthalate (PET) in soil induces oxidative stress and
The plastic additives, such as phthalate residues, chlorinated disrupts the gastrointestinal system in snails, reducing antioxidant
paraffin, Bisphenol A etc. released during weathering leach into the soil enzyme activity [69]. PE and polyvinyl chloride (PVC) in soil alter gut
and alter soil microbial community abundance, diversity, and activities microbiota in certain soil animals, impacting reproduction, growth, and
[57]. Phthalate residues, commonly found in plastics and various con­ survival [76,77]. PS MP affects the growth, reproduction, and survival of
sumer products, have been shown to exert inhibitory effects on micro­ nematodes and induces lipid metabolism disorders in the livers of mice
bial communities. These compounds can disrupt microbial cell [78]. Besides, certain plastic additives, such as vinyl chloride and
membranes, interfere with cellular processes, and serve as substrates for bisphenol A, induce mutagenic and carcinogenic effects, with bisphenol
certain microbial degradation pathways. Consequently, phthalate A disrupting endocrine functions in humans [21,79]. Thus, MP present
exposure often leads to a decrease in microbial abundance and diversity, in soil has far-reaching implications for soil animals, influencing their
particularly among rhizosphere bacteria [58]. This reduction in micro­ physiology, ecology, and potentially posing health risks.
bial diversity can disrupt ecosystem functioning and compromise
microbial-mediated processes such as nutrient cycling and organic

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3. Bioremediation strategies and NP by plants and rhizoremediation by soil microbes and their
enzymes.
3.1. Phytoremediation
3.1.2. Immobilization and degradation of microplastics by rhizosphere
Phytoremediation is a kind of in-situ restoration technology which microorganisms
involve plants and associated soil microorganisms to decrease the con­ Plant roots entrap MP, making them immobile, thus, decreasing their
centrations of pollutants such as MP and NP in the soil ecosystems [80]. bioavailability to the other parts of the plant and reducing the potential
As mentioned in the previous section, terrestrial plants are significantly harm to other soil organisms. Although there is no report available on
affected by the harmful effects of MP and NP, however, they have the MP immobilization in terrestrial plants, hydrophytes such as Thalassia
potential to manage MP pollution through various phytoremediation testudinum [88], Lemna minor [89,90], and Fucus vesiculosus [91]are
mechanisms discussed below. capable of adsorbing MP leading to reduced mobility and bioavailability
in the aquatic environment. Adhesion of MP/NP on large leaves surfaces
3.1.1. Microplastics accumulation and uptake by plants is also a possible way of phyto-stabilizing MP. The adsorption of MP on
Size, shape, surface charge, composition and mechanical properties leaf surfaces is a result of various interactions like electrostatic or Van­
of MP play a crucial role in its accumulation and uptake by plants. MP der Waals interactions, hydrogen bonds, micropore filling, and π-π in­
especially NP <100 nm in diameter can enter the plant cell nucleus by teractions between MP/NP and polysaccharides and other components
penetrating through nuclear membrane thereby disrupting the struc­ such as lipids, proteins, cellulose and lignin, phenolic compounds, sur­
tural and functional properties of chromatin [81,82]; for example, broad face active compounds, minerals etc. present on plant leaves [92]. Thus,
bean (Vicia faba) [37]and Cress (Lepidium sativum L.) plant [83]. Nega­ leaves provide a large sink to pool atmospheric deposition of MP.
tively charged NP like PS-SO3H and positively charged NP like PS-NH2 Mycorrhizal fungi associated with plant roots can potentially immobi­
were found to be accumulated in Arabidopsis thaliana [84]. The uptake of lize MP/NP, further enhancing their removal from the environment. For
smaller MP takes place through endocytosis as the size of the endocytic example, AMF which is present in 90% of the plant species and functions
vesicles is <200 nm in diameter [85]. Root tips are the most susceptible to promote phosphorus uptake by plants and protect host plants against
part of the plant for MP uptake [37,82]. Various reports suggest the environmental stresses, including plastics and heavy metal contamina­
accumulation of PS microbeads/MP of size 0.1–5 μm by the roots and its tions [93]; and Ectomycorrhizal fungi (e.g. Lactarius delicious) helps in
transportation to various parts of Wheat (Triticum aestivum L.) [86], reducing the MP associated stress by enhancing the potassium and
Maize (Zea mays L. var. Jubilee) [87] and carrot (Kurodagosun) [83]. phosphorus absorption from the environment, which is beneficial to its
However, the larger MP of 0.2- 200 μm, enter through crack-entry own metabolic activities [94].
pathway in the lateral roots of plants [81,84]; for example, Wheat To resist the MP uptake process, plant releases mucilaginous exu­
(Triticum aestivum) and lettuce (Lactuca sativa) [81]. Thus, MP, specif­ dates rich in organic acids and amino acids that enwrap MP and stops its
ically NP are easily translocated because of their high bio-accessibility entry inside the plant (Fig. 2) [84,95]. Several bacteria (e.g., Serratia
and bioavailability to the plant cell. The evidence of phytoaccumula­ plymuthica), fungi (e.g., Laccaria Laccata) and their enzymes present in
tion and translocation of MP and NP in various parts of the plants hints soil ecosystem are capable of degrading MP (e.g. PS, PLA, and PET) by
the removal of these contaminants from MP polluted environment. Fig. 2 depolymerising their carbon backbones [96]. Root exudates are a good
shows diagrammatic representation of uptake and accumulation of MP carbon source for MP degrading microorganisms, promoting their

Fig. 2. Schematic diagram showing phytoremediation strategies: accumulation and uptake of MP and NP by plants and rhizoremediation by soil microbes and
their enzymes.

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growth and activity. These bio-degraders can either transform MP into (Galleria mellonella) [110] and super worms (Zophobas atratus) [111] can
plant biomass and contribute to carbon sequestration or mineralised into degrade, and metabolize PS, PE, and PVC [112]. Other soil invertebrates
inorganic carbon in the form of CO2 and CH4 leading to greenhouse like snails (Achatina fulica) can ingest and degrade PS plastics [113] and
emissions and contributing to global warming [97,98]. Therefore, various bacterial strains present in the gut of earthworms (Lumbricus
further research is needed to better understand the impact of MP on terrestris) can decay LDPE MP [114]. The potential MP degrading soil
plant metabolic activities and carbon cycle, as both the potential ben­ animals are represented in Table 1.
efits and environmental risks are associated with phytoremediation. Kasahun et al. summarized the significant role of earthworms in
Rhizosphere microorganisms such as Cyanobacteria, Actinobacteria, maintaining soil ecosystems by accumulating heavy metals and
Bacteroidetes, Gemmatimonadetes, Proteobacteria, Ascomycota and Basi­ degrading MP. They highlighted the protective role of drilodefensins, a
domycota in conjunction with root exudates, offer promising potential metabolite in earthworms’ guts, against oxidative stress caused by plant
for MP rhizoremediation potentially utilizing them as a carbon source polyphenols [123]. Earthworms, such as Eisenia fetida, enhance their
[99]. It also plays a crucial role in plant detoxification promoting the antioxidant properties in response to MP exposure by secreting various
plant growth [100]. A recent study demonstrated the tolerant effect of enzymes like glutathione, glutathione-related enzymes, catalase,
an ornamental plant, Mirabilis jalapa L. upon exposure to galaxolide acetylcholine esterase and malondialdehyde, superoxide dismutase, and
(HHCB; 1,3,4,6,7,8-hexahydro-4,6,6,7,8,8,-hexamethyl-cyclopenta[g] lipid peroxidation enzymes [124,125]. They also form symbiotic re­
benzopyran) and PS MP and NP. HHCB when used solely, enhanced the lationships with bacteria and fungi in their gut, minimizing the harmful
activity of antioxidant enzymes like superoxide dismutase (>200%), effects of MP. Studies showed Lumbricus terrestris surviving well in 1%
indicating the plant’s capability to mitigate oxidative stress induced by MP-contaminated soil, but mortality increased significantly with MP
MP and NP. Co-exposure of HHCB and PS enhanced the activity of su­ exposure alone. Earthworms’ gizzards aid in MP fragmentation, sug­
peroxide dismutase by 93.8%. This co-exposure might have exerted gesting a potential bioremediation solution [115]. Earthworms can
synergistic effects on oxidative stress pathways. For example, HHCB- serve as test animals for monitoring MP toxicity. Researchers developed
induced production of reactive oxygen species combined with PS- a statistical model correlating LDPE-induced oxidative stress with the
induced cellular stress may resulted in higher levels of oxidative dam­ physiological response of earthworm microbial communities, revealing
age compared to individual exposures. In response, cells might have detoxification mechanisms. Earthworms remain unaffected at low LDPE
upregulated SOD activity more significantly to counteract the increased concentrations but suffer oxidative damage at higher levels. Microbial
oxidative stress caused by the combined exposure. Stomatal conduc­ communities inhibit lipid peroxidation to counteract LDPE-induced
tance and transpiration rate were also affected by the exposure of MP oxidative stress, becoming more active with increased LDPE concen­
and NP suggesting changes in water uptake efficiency of the plant. These tration. These findings offer a framework for monitoring ecotoxicity and
contaminants also affect various pathways like carbohydrate synthesis mitigating MP pollution, suggesting earthworm propagation in agri­
and energy metabolism in M. Jalapa [101]. Further research is needed to cultural lands for bioremediation and sustainable soil management
optimize the phytoremediation process and long-term effectiveness of [126].
M. jalapa plant. Similarly, various soil fauna significantly contributes to the biore­
There are many rate-limiting steps involved which can slow down mediation of MP in the soil. Recent studies have shown that mealworms
the overall process of rhizoremediation. One of the limitations is un­ (Tenebrio molitor) utilize ancient lignin digestion mechanisms to biode­
availability of contaminants due to low solubility or poor soil conditions. grade polystyrene with the assistance of their gut microbiota. This is
Many plants rely on rhizosphere microorganisms for degradation of MP/ because most of the insects, worms, and other soil animals rely on
NP. However, their activity can be influenced by many factors such as lignocellulose, a plant component as their primary dietary source. The
nutrient availability, soil type, pH, moisture content and plant species bonds within synthetic plastic polymers can resemble to those found in
etc. This may affect the biodegradability of rhizosphere microorganisms. natural polymers like lignin. The larvae of mealworms can metabolize
Indigenous microbial population in the soil may also compete with PS and CS MP by increasing the expression of various genes such as
rhizosphere microorganisms for resources and space, thereby reducing Lac640, superoxide dismutase, and CYP6/4 which leads to the oxidation
the efficiency of rhizoremediation. Even if contaminant reaches the of C–C bonds of both polystyrene and lignin. This resulted in the con­
roots, plant itself may not be able to uptake or metabolize the contam­ version of MP into their monomers and aromatic residues. The aromatic
inant. Environmental factors such as temperature, moisture, and oxygen
availability may also influence the rate of contaminant degradation and
microbial activity in the rhizosphere. Suboptimal environmental con­ Table 1
ditions can slow down the bioremediation process. Soil animals capable of degradation of plastics.
Despite these limitations, rhizoremediation still holds promising Soil animal Microplastic Biodegradation Duration Reference
potential. Various novel approaches like genetic engineering can also be degraded efficiency
used to enhance the ability of plants and microorganisms to withstand, Lumbricus LDPE, PLA, Survived with 0% 35 days [115]
bioaccumulate, and metabolize MP contaminants. Chemical agents and terrestris polybutylene mortality
biochar can also be included in the soil to enhance bio-stimulation and adipate
terephthalate
co-immobilization to improve the MP rhizoremediation process further Tenebrio PS 7.4 % PS Wt loss 60 days [116,117]
[102–104]. Therefore, rhizoremediation can be a promising approach molitor
towards MP and NP management. A critical evaluation of various MP PE 12% loss in the 36 days [118]
sources, their exposure pathways and vulnerable recipients is essential faeces of larvae
PE, PU, PS 69.7, 53.2, 46.5% 58 days [119]
to design effective phytoremediation strategies [105].
weight loss
Zophobas PS 36.7 % of PS 16 days [108]
3.2. Bioremediation by soil animals atratus converted to CO2
PP 20.4 % decrease 8 days [120]
in Mw of frass
The abundance of MP and NP in the soil ecosystem leads to greater
G. mellonella PE 43.3% wt loss by [121]
interactions of MP with terrestrial biota such as earthworms, snails, 100 insects
mealworms, waxworms, super worms, slugs, centipedes, grubs etc. Achatina Expanded 14 times wt. loss 28 days [113]
Various studies have highlighted the importance of numerous insects, fulica. Polystyrene EPS
soil animals, and their gut microbiota in degradation of MP [106–108]. Dark PS foam 26.03 % wt. loss 31 days [122]
mealworm
For example, yellow worms (Tenebrio molitor) [109], wax moths

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rings of these residues were further cleaved by various enzymes like vulcanized rubber with 18.13% carbon loss and a 52.90% reduction in
alkylglycerol monooxygenase, aldehyde dehydrogenase, alcohol dehy­ cross-linking degrees after four weeks of incubation [129]. Besides,
drogenase, and peroxidase. Long-chain fatty acid CoA ligase, carbox­ other factors that affect the cross-linkage between rubber polymers
ylesterases, and lipases hydrolyse the resulting products to generate include temperature, pressure, humidity, exposure to chemicals, and
long-chain fatty acids which are either stored in the host cell or enter radiation. These factors can influence cross-linking reactions, which
tricarboxylic acid (TCA) cycle to generate metabolic energy for the determine the strength, durability, and flexibility of rubber polymers.
growth and development of the larvae. These enzymes are not directly Additionally, another member of the Tenebrio genus, Tenebrio obscurus
involved in plastic degradation, whereas indirectly help in the break­ (dark mealworms), also degrades PS and PE in a similar manner with the
down of MP made from lipid-based polymers, into degradation products help of their gut microbiome [122]. When these gut microorganisms
that can be further metabolized by microorganisms and their enzymes utilize PE and PS polymers as carbon source, CO2 is primarily produced
[127]. as a metabolic byproduct through oxidation under aerobic conditions,
Soil invertebrates like snail (Achatina fulica) are widely distributed whereas CO2 and CH4 are generated as end products under anaerobic
worldwide and often encounter with soil MP. Researchers have found conditions [58].
that after ingesting MP through various food sources like plastic- Another soil animal, super worm (Zophobas atratus) can feed on
contaminated leaves and litter present in the soil, these soil in­ styrofoam (a form of PS) as a sole food source with an average con­
vertebrates can egest MP in their faeces. PS-fed snail faeces form sumption rate of 0.58 mg/day. In the gut of its larvae, plastic degrada­
biodegradable intermediates with new oxidized functional groups, as tion occurs by depolymerizing long-chain PS monomers to low
evidenced by FTIR and 1H NMR spectra. Tenebrio genus also produced molecular weight products. A research group has compared the PS and
such intermediates during PS biodegradation [109,122]. Gut microor­ polyurethane (PU) foam degrading capacity of Z. atratus and T. molitor
ganisms play a role in the degradation and depolymerization of PS, for a duration of 35 days. The average rate of consumption by Z. atratus
resulting in lower molecular weight polymers. is 49.24 mg PS/larvae and 26.23 mg PU/larvae, which is 18 and 11
Interestingly, following the ingestion of PS, the snails exhibited a times higher than T. molitor, respectively [130]. Fig. 3 shows general
shift in their gut microbiota associated with polystyrene biodegradation, mechanism of biodegradation of MP and NP by soil animals.
as revealed by high-throughput sequencing analysis. The study Soil animals and insects-based remediation of MP and NP is a highly
demonstrated an increase in Enterobacteriaceae from 11.9% to 64.4% efficient and cost-effective method that does not generate secondary
and Sphingobacteriaceae from 6.7% to 25.5% after MP ingestion. pollutants. It is a promising tool with the potential to combat MP
Notably, there was no discernible negative impact of PS ingestion on the pollution, offering a sustainable solution for future generations.
growth and development of the snails [113].
Tenebrio moliter larvae, with the help of their gut microbiome (e.g.
Streptococcaceae and Spiroplasmateceae), can biodegrade PS and PVC 3.3. Degradation of microplastics by microbes or their product
products into CO2 [107] [112]. T. molitor can partially mineralize
polyvinyl chloride (PVC), releasing 2.9% of chloride from the total PVC MP being synthetic polymers made up of carbon and hydrogen, are
ingested. It can also biodegrade vulcanized styrene-butadiene rubber by found to be attacked by microorganisms. Various studies have reported
decreasing the cross-linkage between the rubber polymers as monitored the colonization of microbial communities such as bacteria, archaea,
by thermal analysis, FTIR, X-ray diffraction, and SEM analysis [128]. fungi, algae, diatoms, viruses, and protozoans in the form of biofilms
Actinobacteria, specifically Streptomyces species, also exhibited the over MP surface, which is known as “plastisphere” [131]. Interpreting
capability to degrade vulcanized rubber by breaking cross-link sul­ the role of MP in harbouring rich microbial consortium and the action of
fur‑sulfur or carbon‑sulfur bonds while keeping the main polymer chain microorganisms in the degradation of MP is of great ecological benefit
intact. For instance, Streptomyces sp. demonstrated devulcanization of [132]. The spread of microbial consortium in form of biofilm over the
MP occurs at several different stages and follows a temporal succession

Fig. 3. General mechanism of MP bioremediation by gut microbes and their enzymes presents in soil animals.

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pattern [131]. The process starts with microbial colonization initiated Chaetomium, Curvularia, Aspergillus, and bacterial species within the
by pioneer microorganisms, which proliferate and spread throughout genera Corynebacterium, Acinetobacter, and Pseudomonas have also
the surface of MP. The progression of biofilm formation is accompanied exhibited the ability to degrade PU [145,146]. Oviedo-Anchundia et al.,
by a succession in the microbial population. The microbial diversity determined the potential of Penicillium, Geomyces, Mortierella species in
observed varies depending upon the variation in microhabitat around biodegradation of PU [147].
the MP, environmental conditions, composition, type and size of the MP Several studies have highlighted the role and mechanism of micro­
[133]. Studies have confirmed the variation in taxonomic specificity on organisms in the efficient biodegradation of MP [131,138,139]. The
plastics with different chemical composition. Gammaproteobacteria (39 microorganisms are reported to directly degrade the MP to derive car­
± 9%) dominated on PE surface compared to Alphaproteobacteria (34 ± bon and energy; or they produce several enzymes, metabolites and
6%), while on PLA was dominated by Alphaproteobacteria (54 ± 4%) as biochemicals that eventually act as catalysts in the degradation of
compared to Gammaproteobacteria (22 ± 5%). The presence of Plancto­ complex polymers into simpler units which are subsequently assimilated
mycetes was 20 ± 5% on PE and 10 ± 5% on PLA [134]. Further, the by the microorganisms [144]. Several enzymes such as polyurethanase,
microbial activity was also found to vary with the shape and size of the esterase, laccase, hydrolase, dioxygenases, and peroxidases have been
MP. In case of plastics with same mass, an irregular surface area reported to facilitate MP biodegradation processes [148,149]. These
demonstrated higher microbial abundance, diversity and activity as enzymes, however, offer limited solution and their activity, efficacy and
compared to a regular one, and with smaller size showed higher mi­ depend on several factors such as degree of polymerization, crystallinity,
crobial activity in contrast to large sized ones [135]. Polymers with high hydrophilicity of polymer, their breakdown products, temperature.
degree of crystallinity (~30-50%) such as PET based plastics have low Majority of enzymes such as esterases, hydrolases can efficiently
rate of biodegradability. The density of polymer also influences its hydrolyse amorphous, less crystalline polymers and perform better at
biodegradation. Study has revealed that the polymer arrangement in moderate temperature ranging between 55 and 65 ◦ C [150]. Further, in
low density polyethylene makes it more susceptible to oxidation and a study it was found that copper markedly affected the activity of bac­
biodegradation as compared to high-density polyethylene. Further, the terial laccases facilitating polyethylene degradation [151]. For enzymes
higher molar mass of high-density polyethylene also makes it difficult such as polyurethanase, dioxygenases, and peroxidases, the rate of
for microbes, biochemicals and enzymes to access the polymer chain for biodegradation was found to be inversely related to molecular weight,
biodegradation [27]. The prevalent pioneer microbial communities in degree of polymerization and crystallinity of polymer [150]. The
plastispheres in marine and estuarine generally are green algae, cya­ extracellular enzymes such as laccase, esterase, and peroxidase pro­
nobacteria, diatoms, Gammaproteobacteria, Marinobacter, Alcanivorax, duced by the microorganisms are found to have a significant role in
Vibrio spp. [136,137]. Upon maturation of plastisphere, microbes exposing the functional group of polymers like carbonyl, carboxylic,
including Bacteroidetes, Rhodospirillaceae, Rhodobacteraceae, Alphapro­ hydroxyl groups thereby increasing the overall hydrophilicity of the MP.
teobacteria, spirochetes emerge as dominant colonizers on MP substrates The increase in hydrophilicity facilitates efficient attachment of micro­
[138,139]. organisms to the MP surface, formation of short polymer intermediates,
The microbial community sequentially degrades MP by enzymatic their assimilation by microbial cells and ultimately biodegradation
hydrolysis, breaking down the polymer into oligomers, dimers, and [152]. Several hydrolases such as esterase, depolymerases and lipases
monomers, which are then assimilated as sole carbon and energy sour­ act to hydrolyse chemical bonds or side chains of MP and disrupt the
ces, ultimately undergoing complete mineralization to carbon dioxide structure and length of polymer chain [153]. Alkane hydroxylase is an
and water (Fig. 4). important enzyme produced in microorganisms such as Pseudomonas,
The degradation alters the chemical composition, functional groups, Micrococcus, Acinetobacter, Rhodococcus that is capable of aerobic
molecular weight, and tensile strength of the polymer. Biodegradation degradation of PE. These enzymes are involved in hydrolysis and
of MP is a prevalent phenomenon both in terrestrial and aquatic envi­ metabolism of linear n-alkanes molecules present in PE. The enzyme
ronments [22]. Microorganisms present in soil, landfills, sediments, initiates the reaction by hydroxylation of C–C bonds leading to pro­
oceans, estuaries have been found to be capable of degrading various duction of primary/secondary alcohols that get oxidized to aldehydes or
types of MP. Microorganisms such as Nesiotobacter, Achromobacter, ketones and subsequently to carboxylic acids. The hydrophilic carbox­
Micrococcus, Aspergillus, Acinetobacter, Staphylococcus, Arthrobacter, Ba­ ylic acids are later catabolized by microorganisms through the
cillus, Stenotrophomonas, Comamonas, and Pseudomonas are the major β-oxidation pathway and enter the tricarboxylic cycle [154]. Mon­
soil inhabitants which are efficient degraders of PE [140–142].) Species oxygenase, an important alkane hydroxylase enzyme has been demon­
of Ideonella, Aspergillus, Bacillus, Staphylococcus, and Saccharomonospora strated to facilitate extracellular oxidation and hydrolysis of PE
have also been identified as potential degraders of highly crystalline polymers. The enzyme produced by Pseudomonas putida has been shown
polymers such as PET [143,144]. Microorganisms such as Pseudomonas, to catalyse the hydroxylation of the terminal carbon in the alkane chains
Achromobacter are capable of degrading PVC [140]. Fungi such as of the polymer. Previous studies underscored the significance of bacteria

Fig. 4. Biodegradation of MP to generate carbon dioxide and water.

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strains P. knackmussii N1-2 and P. aeruginosa RD1-3 in the biodegrada­ biodegradation (Fig. 5). Further, the application of several gene editing
tion of polyethylene (PE), resulting in mass reductions of 5.95 ± 0.03% tools such as zinc finger proteins, clustered regularly interspaced
and 3.62 ± 0.32% in 8 weeks, by virtue of enzymatic activities such as palindromic repeats (CRISPR-Cas9) along with the knowledge of bio­
dioxygenases, monooxygenases and hydrolases [155,156]. Further, en­ informatics, and computational biology, have made the designing pro­
zymes such as cutinases produced by microorganisms such as Fusarium cess easy and precise by providing extensive details related to the
solani pisi, Thermobifidafusca, and T. alba have also been reported to biodegradation pathways, enzyme action, and associated genes [167].
hydrolyze polyester-polyurethane. Peptide engineering was shown to These tools offer a great advantage in predicting the biodegradation
further improve the catalytic activity of this enzyme 6.7-fold with a route and end products of MP biodegradation.
reduction in degradation time from 41.8 h to 6.2 h [157]. The extra­ Several microorganisms, including bacteria, fungi, algae, and di­
cellular enzymes are more efficient in corroding and degrading the atoms have been synthetically engineered to enhance their MP biodeg­
surface of the polymer as they usually fail to penetrate the polymer radation capabilities. Ideonella sakaiensis 201-F6 produces PET
structure; however, both intracellular and extracellular enzymes play hydrolases that hydrolyse PET to terephthalic acid, and ethylene glycol
significant role in MP biodegradation and microbial assimilation [158]. [165]. Attempts have been made to convert non-degrading strains to
PS serves as a carbon source for microorganisms, including Rhodococcus PET degraders through the transfer of genes encoding PET hydrolytic
ruber, through enzymatic oxidation involving laccase, hydroxylases, enzymes. Moog et al., have transformed Phaeodactylum tricornutum, a
oxidoreductases, and peroxidase; this process transforms PS into phe­ photosynthetic microalga into a PET degrader through genetic modifi­
nylacetate, subsequently entering the tricarboxylic acid (TCA) cycle for cation by the insertion of exogenous genes encoding PET hydrolytic
further metabolic processing [159]. Various studies have suggested the enzymes [168].
role of abiotic pretreatment to further facilitate MP biodegradation Genetically engineered E. coli, P. chrysosporium, and S. cerevisiae BY
[160,161]. Pretreatments such as UV irradiation, and thermal treatment 4741 were constructed with the ability to produce enzymes such as
have been shown to increase the hydrophilicity of the bioplastic or lead manganese-dependent peroxidase and laccase for efficient MP biodeg­
to the introduction of carbonyl and hydroxyl groups to the polymer radation [176,177]. Zurier and Goddard engineered and expressed the
making it more susceptible to microbial degradation [162]. Biodegra­ PETase enzyme in E. coli to enhance the degradation of MP fibres in
dation of PP was shown to be improved by UV irradiation followed by wastewater sludge by 17.4-fold [178]. Cloning and expression of the
the action of Bacillus flexus [163]. Another study on biodegradation of PETase gene in the green microalga Chlamydomonas reinhardtii also
UV pretreated PU samples revealed an improved degradation potential demonstrated efficient degradation of PET in 4 weeks under ambient
of Penicillium to 28.3%, and Geomyces and Mortierella achieving 24.9% condition [179]. Yeast, Yarrowia lipolytica, engineered by insertion of
and 26.3% respectively [147]. MP such as PET and PU composed of PETase gene also demonstrated its ability to degrade PET efficiently. The
carbon and heterogenous atoms can be efficiently biodegraded by degradation was further enhanced by 66% higher release of terephthalic
photooxidation followed by biodegradation [164]. Bacteria such as acid by supplementation of olive oil to the mutant culture. Oliveira and
Ideonella sakaiensis and Pseudomonas putida can efficiently depolymerize Almeida cloned pudA gene of Comamonas acidovorans encoding the
PET using enzymes such as PETase into terephthalate and ethylene esterase enzyme in E. coli to construct a genetically engineered MP
glycol which are then utilized as a source of carbon and energy [165]. degrader strain [180].
Pyrolysis of PET at 450 ◦ C facilitates the easy degradation of PET to Genetic engineering approaches have also been applied in biofilm
terephthalic acid which is then converted into polyhydroxyalkanoate by engineering to enhance the accumulation of MP molecules within it. In
strains like Pseudomonas putida [166]. The potential MP degrading mi­ this direction, Pseudomonas aeruginosa was modified by the deletion of
croorganisms and the associated enzymes are represented in Table 2. the wspF methylesterase gene to boost the synthesis of exopolymeric
substances for improved binding of PVC molecules [181]. In another
3.4. Synthetically engineered microorganisms study, PET biodegradation was enhanced by cloning the gene encoding
the polyester hydrolase enzyme responsible for PET degradation from
Biotechnological interventions in the form of synthetically engi­ Pseudomonas aestusnigri to E. coli [182]. There have been considerable
neered microorganisms have gained attention as sustainable bioreme­ attempts at genetic modification of microorganisms as well as enzymes
diation tools to manage MP pollution. Since all the indigenous to improve MP biodegradation. However, there are still gaps in the
microorganisms do not have the potential to degrade MP, genetic en­ transferability of the results from the laboratory to the real-world en­
gineering and synthetic biology approaches are being applied to the vironments. Moreover, metabolic pathways for biodegradation of
construction of synthetically engineered strains for improved diverse MP types also need to be extensively elucidated for constructing

Table 2
Microorganisms and enzymes capable of degradation of plastics.
Microorganism Enzymes Microplastic Biodegradation Duration Reference
degraded efficiency

Bacillus cereus LDPP 78.62 ± 2.16% 21 days [140]

Bacillus paramycoides PE 72.50 ± 20.53%
Altermonas PVC 1.76% 60 days [145]
Penicillium PU 18.3% 90 days [144]
Geomyces 16.4%
Mortierella 11.3%
Pseudomonas knackmussii N1–2 Hydroxylases, Monooxygenase, Polyphenol oxidase and PE 5.95 ± 0.03% 8 weeks [156]
Laccase
Penicillium simplicissimum Lipase PET 3.09% 28 days [169]
Pseudomonas Lipase PET 30% 28 days [162]
Alcaligenes faecalis LNDR-1 Lipase, CMCase, xylanase, and protease PE 21.72 ± 2.1% 14 days [170]
Pseudomonas and Enterobacter PE 10-15% 150 days [171]
Engineered strains of Clostridium PET film 60% 14 days [172]
thermocellum
Lysinibacillus sp. PP, PE 4 and 9% 26 days [173]
Pseudomonas aeruginosa DSM 50071 Serine hydrolase PS 2.6% 15 days [174]
Pseudomonas sp. AKS31 Hydroxylase LDPE 20% 45 days [175]

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Fig. 5. Approaches to construct synthetically engineered microbes.

engineered microbial species with the potential to enhance the scal­ with other native microbes, soil animals, and plants is still missing.
ability of biodegradation methods. Therefore, we must ensure the biosafety of genetically modified mi­
crobes in bioremediation applications for terrestrial ecosystems. The use
4. Future aspects of various cutting-edge technologies like genomics, transcriptomics,
proteomics, and metabolomics can also help in understanding the
The tremendous usage and increasing demands of plastics have interaction between MP and soil microbiota, their enzymes, and genes
created a new global issue- the disposal of plastic waste, which needs responsible for coding the proteins involved in the biodegradation
urgent attention. MP and NP, the degradation products of plastics, have process, which can provide a close insight into the molecular mechanism
become a major concern because of their potential ecological risks, and of MP biodegradation. The application of artificial intelligence can also
it is critical to find sustainable and safe solutions to this problem. This be utilized to accelerate the rate of MP biodegradation by genetically
review summarizes the harmful effects of MP/NP on soil quality, plant modified organisms. More in-depth, reliable, and sustainable bioreme­
growth and development, soil animals and their microbiota. The focus of diation of MP pollution in terrestrial ecosystems requires collaborative
the review is to describe various bioremediation strategies that can efforts from chemists, environmental engineers, biotechnologists, soil
minimize MP pollution in less explored terrestrial ecosystems. Despite physicists, and material scientists. These future directions are crucial to
global concern regarding soil MP contamination, existing reports have understanding the diversified effects of MP on terrestrial ecosystems and
many gaps. Most of the MP degradation studies are based on laboratory- developing proactive and sustainable solutions to this global concern.
scale research; however, it should also be conducted under realistic
conditions like terrestrial ecosystems, which include agricultural lands, 5. Conclusion
wetlands, forests, etc. One of the major challenges for performing field
studies is to identify and isolate the MP/NP contamination due to their MP pollution has become a pervasive problem in our environment,
very small size and presence in a heterogeneous soil system, which is a which has entered into the terrestrial ecosystem from diverse origins
complex system composed of matter, minerals, soil animals, microor­ including landfills, mulching films, sewage irrigation, industrial efflu­
ganisms etc. Therefore, effective monitoring tools are urgently needed to ents etc. Because of their smaller size, MP and NP can enter easily and
quantify the levels of MP contamination in the soil. There are many pose hazardous effects on terrestrial plants, animals, and soil microbes
reports confirming the role of rhizosphere microbes, soil animals, and and through the food chain to humans. Unfortunately, there is no per­
enzymes as MP bio-degraders. However, more information is needed manent solution to this global concern. Bioremediation could potentially
regarding MP degradation capabilities of other soil-based microorgan­ solve the problems associated with MP/NP. This review has provided
isms such as yeast and soil protists. Data on mixed biological soil pop­ comprehensive and insightful information about various bioremediation
ulation and their synergistic effect on MP degradation are also lacking. strategies to deal with MP pollution in terrestrial ecosystems. It
The in-depth molecular mechanism of MP biodegradation by terrestrial explained how plants naturally cope with the toxic effects of MP using
animals, microbes, enzymes, and other related proteins has also yet to be their rhizosphere microorganisms and enzymes. Soil animals also
explored. Further investigation of the potential health risks associated contribute significantly to degrading, depolymerizing, and mineralizing
with MP uptake and accumulation by plants and soil organisms and their MP and NP to inorganic carbon for use as their carbon source.
potential transfer to the food chain is crucial. Disposal of MP/NP Furthermore, biotechnological advancements in the field of MP biore­
contaminated plants and crops is still a challenge. Research should also mediation are also discussed. Overall, the present review provides
be conducted for the safe and sustainable disposal of MP contaminated several recommendations to overcome the knowledge gaps in the field
plants and soil animals. Many reports reveal the biotechnological and provides a valuable roadmap for future research on MP contami­
advancement in MP bioremediation by reconstructing genetically nation of terrestrial ecosystems.
modified microbes and optimizing their degradation pathways. How­ While it may not be possible to completely eradicate MP from the
ever, information about their efficiency in real-world conditions such as environment, efforts should be made to reduce plastic production and
variation in temperature, pH, or nutrient availability or their interaction consumption, improve waste management practices, develop

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