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Abbreviation (ISO4): Prog Chem      Editor in chief: Jincai ZHAO

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Progress in Chemistry >

2025 , Vol. 37 >Issue 1: 112 - 123

DOI: https://doi.org/10.7536/PC240706

Microplastics Special Issue

Microbial Degradation of Environmental Microplastics

  • Hongqin Guo 1, 2 ,
  • Kai Yang 1 ,
  • Li Cui , 1, *
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  • 1 Key Laboratory of Urban Environment and Health, Institute of Urban Environment, Chinese Academy of Sciences, Xiamen 361021, China
  • 2 University of Chinese Academy of Sciences, Beijing 100049, China
*e-mail:

†These authors contributed equally to this work.

Received date: 2024-07-10

  Revised date: 2024-09-29

  Online published: 2024-12-25

Supported by

National Natural Science Foundation of China(22241603)

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Abstract

Due to the highly stable chemical properties of plastics, plastic wastes disposed into environments are difficult to degrade and can only be broken down into microplastics with smaller particle size and larger surface area through the weathering process. Microplastic pollution has become one of the most pressing environmental issues. There is an urgent need to reduce microplastic pollution in order to protect the ecological and human health. Biodegradation of microplastics can ultimately convert microplastics into environmentally friendly substances such as biomass, CO2, CH4 and H2O or other valuable intermediates. It is thus an environmentally friendly technology to potentially make microplastics harmless and resourceful. This paper reviews the present understanding of microplastics biodegradation processes, the influencing factors, the microbial and enzymatic resources for microplastics degradation, and the up-to-date approaches for mining plastics-degrading microbial resources. It finally provides perspectives on the challenges of current research and the direction of future research on microplastic biodegradation.

Contents

1 Introduction

2 Microplastic biodegradation process

2.1 Degradation pathway

2.2 Influence factors

3 Microplastic biodegradation resources

3.1 Degrading bacteria

3.2 Catabolic enzymes

3.3 Synthetic community

4 Mining strategies for microplastics-degrading microorganisms

4.1 Culture-dependent methods

4.2 Culture-independent methods

5 Conclusion and outlook

Key words： microplastics; plastic biodegradation; biodegradation process; biodegradation resources; mining strategies

Cite this article

Hongqin Guo , Kai Yang , Li Cui . Microbial Degradation of Environmental Microplastics[J]. Progress in Chemistry, 2025 , 37(1) : 112 -123 . DOI: 10.7536/PC240706

1 introduction

Plastic is a polymer compound synthesized by polymerization. It is widely used in many aspects of modern society because of its durability. With the extensive use of plastic products, plastic pollution has become one of the most urgent environmental problems facing mankind, which has attracted great attention of the international community. According to statistics, the total amount of plastics produced in the world by 2015 was 8.3 billion tons, of which 4.9 billion tons were buried or released into the environment[1]Due to the stable physical and chemical structure, plastics are difficult to degrade naturally in the environment and can persist for a long time, with a half-life of 0.19-5000 years（Table 1）[2-3]Waste plastics are constantly broken through weathering in the environment to form micro plastics with smaller particle size and larger specific surface area (diameter less than 5 mm), or even nano plastics (diameter less than 1000 nm)[4⇓-6]It is difficult to recover from the natural environment and poses a potential serious threat to the ecosystem and human health. Microplastics can enter human body from respiratory tract and digestive tract, and can be detected in many tissues and organs of human body, including intestine, lung, liver, kidney, blood, placenta, etc., causing potential harm to human nervous system, immune system, reproductive system, etc[7]The United Nations Environment Programme has listed plastics as a global emerging pollutant[8]In May 2022, China promulgated the action plan for the treatment of new pollutants to further strengthen the treatment of plastic pollution at the national level. In order to protect the ecological environment and people's health, it is urgent to reduce and control micro plastic pollution.
Table 1 Types, structures and properties of common plastics[2-3]

Table 1 Plastic types，structures，and properties[2-3]

Polymers Abbreviation Molecular formular Backbone structure Estimated half-lives ( year)
polystyrene PS (C8H8)n >2500
polyethylene PE (C2H4)n 58~5000
polypropylene PP (C3H6)n 53~780
polyethylene terephthalate PET (C10H8O4)n 2.3~2500
polyvinyl chloride PVC (C2H3Cl)n >2500
polyurethane PU (NHCOO)n 10~999
polycaprolactone PCL (C6H10O2)n 0.19~3.1
polylactic acid PLA (C3H4O2)n 0.19~3.1
polyhydroxyalkanoates PHA (C5H8O2)n 0.19~3.1
polyvinyl alcohol PVA (C2H4O2)n 0.19~3.1
polyester PES (C6H10O5)n 0.19~3.1
polycarbonate PC (C16H18O5)n 0.19~3.1
polyhydroxybutyrate PHB (C4H6O2)nH2O 0.19~3.1
polyamide PA (C6H11NO)n 0.19~3.1
The traditional treatment of waste plastics is mainly landfill and incineration. However, these disposal methods are not environmental friendly. The landfill process not only occupies a lot of land resources, but also easily causes secondary pollution, such as soil and groundwater pollution caused by the release of harmful chemical additives (such as plasticizer, flame retardant, antioxidant, etc.) in plastics. Although incineration can reduce the amount of plastics, it will emit greenhouse gases and volatile toxic organic compounds, which is contrary to the concept of "carbon neutrality". Environmental microorganism is an important driving force of material cycle in ecosystem. In the long-term evolution process, microorganisms constantly adapt to various pollutants produced by human activities, and can even use specific biochemical reactions to degrade and transform them to obtain materials and energy. Compared with traditional plastic reduction measures, microbial degradation of micro plastics is the use of microorganisms widely existing in the environment to ultimately convert plastic waste into environmentally friendly substances such as carbon dioxide and water or other high-value intermediates, which has great application potential for reducing environmental micro plastics pollution. The research trends of microbial degradation of microplastics mainly include: ① in scientific understanding, from the separation and verification of degrading microorganisms to the analysis of degradation path and functional mechanism; ② In terms of research methods, from traditional isolation and culture to multi group technology, single cell technology, high-throughput screening technology and other methods; ③ In engineering applications, from the in-situ colonization of high-efficiency degrading bacteria to the design of synthetic functional flora and the transformation of key degrading bacteria and enzymes by genetic engineering. Its key milestones include: ① discovery of plastic biodegradation: in 2010, Albertsson[9]The phenomenon of plastic biodegradation by environmental microorganisms was revealed by radioisotope labeling, and the possibility of plastic biodegradation was proposed; ② Screening of degrading bacteria and revealing degradation mechanism: 2016, Yoshida et al[10]Screening and identification of pet degrading bacteria for the first time（Ideonella sakaiensis201-f6) and enzyme (petase) were used to analyze the degradation mechanism. Since 2015, Yang et al. And Luo et al. Have screened plastic degrading microorganisms from the intestines of plastic degrading insects (such as Tenebrio molitor, barley worm, earthworm, cockroach, etc.)[11⇓-13]③ genetic engineering and synthetic biology: Zhu et al. Since 2020[14]And Tournier et al[15]Through genetic engineering and synthetic biology technology, the structure and function of specific enzymes involved in the degradation of microplastics were redesigned, and the high-efficiency expression of key degradation enzymes and in-situ environmental remediation were carried out by engineering bacteria. For example, petase gene was highly expressed in E. coli and achieved good pet degradation effect[14]④ commercial application: in the past five years, many international research teams and enterprises have devoted themselves to developing microbial degradation technology that can be applied on a large scale, which has promoted the commercialization of plastic degradation technology[15]These research trends and milestones mark the rapid development of the field of micro plastic biodegradation and reveal the potential of microorganisms in plastic degradation. In this paper, the degradation mechanism, influencing factors, degradation resources and mining methods of microplastics are reviewed, and the current challenges and future research directions of microplastics degradation are prospected.

2 Microbial degradation of microplastics

Microbial degradation of environmental microplastics is the use of microorganisms and enzymes in the environment to ultimately convert microplastics waste into biomass CO2、CH4And H2The process of O and other environmentally sound substances or other high-value intermediate products is an environmentally friendly resource and harmless control technology.

2.1 Degradation process

The biodegradation process of microplastics can be divided into four stages, and the degradation of microplastics depends on the cooperative implementation of four degradation steps by microorganisms[16]。
(1) Biological degradation: change the chemical and physical properties of polymers. As a non-polar polymer, plastics have limitations in biodegradation due to their surface hydrophobic properties. The process of biological degradation starts with the adhesion and colonization of microorganisms on the polymer surface. The modification of the physical and chemical properties of the polymer surface caused by the formation of biofilms by microorganisms reduces the hydrophobicity and durability of plastic materials, thus promoting the degradation of micro plastics by microorganisms.
(2) Depolymerization: microplastics are cleaved into oligomers, dimers or monomers by enzymes released by microorganisms. Depolymerization mainly includes two main reactions, namely, the reduction of polymer molecular weight and the oxidation of low-mass molecules[17]Microorganisms colonizing on the surface of micro plastics further secrete extracellular enzymes and intracellular enzymes, which attack the molecular chain of plastics, breaking the end of the hydrolyzable chemical bond or molecular chain, producing monomers, dimers and other oligomers, resulting in the reduction of the molecular weight of micro plastics. At the same time, it is further oxidized under the action of oxygen and environmental ROS to form hydroxyl, carbonyl, carboxyl and other oxygen-containing functional groups[18-19]To promote the further degradation of microplastics. Hydrolases (such as lipase, esterase, protease, etc.) and oxidoreductases (such as laccase, monooxygenase, oxidase, peroxidase, alkane hydrogenase, etc.) are two main types of micro plastic degradation enzymes. Hydrolase is a kind of enzyme with the ability to catalyze the hydrolysis of substrates. It can degrade plastics with chemical bonds (such as c=o bond, ester bond) that are easy to hydrolyze in the structures of pet, PLA, PU, PVA, etc. Oxidoreductase is a kind of enzyme with lignin degradation ability, which can degrade micro plastics containing C-C single bond that is not easy to hydrolyze in PE, PS, PP, PVC and other structures（Figure 1）。
Figure 1 Biodegradation path of microplastics

Fig. 1 Biodegradation pathways of microplastics

(3) Assimilation: oligomers formed after chain breaking are transported to the cytoplasm of microorganisms in the assimilation stage. Oligomers formed after chain breaking will be released into the surrounding environment, and some oligomers with low molecular weight will enter the microbial cells for further metabolism through different membrane transport systems[17]。
(4) Mineralization: the oligomer entering the microbial body decomposes and metabolizes in the body to form Co2、CH4And H2O. Micro plastic derivatives enter microbial cells, gradually remove carbon atoms on the molecular chain of micro plastic polymer through β - oxidation mechanism, generate acetyl coenzyme A, enter the TCA cycle, oxidize organic substances and produce ATP to provide energy for cells[20]Finally, after a series of metabolism in vivo, the oligomer is mineralized and transformed into biomass in cells, and CO is produced at the same time2、CH4And H2O [21]。

2.2 influence factor

2.2.1 Physical and chemical properties of microplastics

The type, structure, crystallinity, molecular weight, hydrophilicity, hydrophobicity and additives of microplastics will affect the biodegradation efficiency of microplastics[20]The degradation of microplastics mainly depends on its chemical composition. Compared with other synthetic polymers, starch based plastics, such as PLA, PVA, PHA, PBAT, which are made of starch and its derivatives as the main raw materials, are easy to be invaded by microorganisms and secrete enzymes to decompose starch into smaller carbohydrates, with strong biodegradability[22]However, biodegradable plastics usually need controlled conditions to decompose effectively. For example, the degradation rate of PLA and PBAT is 90% within 180 days under anaerobic composting at 70 ℃[22]However, in natural environments such as seawater, soil and landfill sites or through household composting, the degradation rate is much slower and usually takes decades[22]Polyester micro plastics, such as pet, PU, PCL, etc., contain hydrophilic groups such as hydroxyl, carboxyl, carbonyl and hydrolyzable ester bonds, which are easy for microbial colonization and enzymatic hydrolysis[23]However, polyolefin micro plastics contain a stable carbon chain skeleton composed of high bond energy C-C bonds, lack of groups that are easy to be oxidized and hydrolyzed, and are less likely to be biodegraded[22]In addition, the structure of microplastics also affects biodegradability. For example, the water absorption of PLA with different stereostructures is different, and the degradation efficiency of racemic PLA with high water absorption is higher than that of racemic PLA under the action of protease K[24]The crystallinity of microplastics also affects the degradation of microplastics. The chemical structure and properties of polymers with higher crystallinity are more stable, while the molecular chains in semi crystalline or amorphous polymers are loosely arranged, and there are more voids and active sites between the molecules of micro plastics, which are more vulnerable to attack by microorganisms and enzymes. Therefore, choosing plastics with lower crystallinity or reducing the crystallinity of micro plastics can promote the biodegradation of micro plastics[25]In addition, the molecular weight of microplastics is also an important factor affecting the degradation of microplastics, and the lower molecular weight is easier to be used and degraded by microorganisms and enzymes. For example, intestinal microorganisms of Tenebrio molitor and Hordeum vulgare can efficiently degrade low molecular weight（Mw<6.2 kDa) of PP, PE, PS and other polyolefin micro plastics, which can only be degraded to a limited extent for high molecular weight micro plastics[26]Microplastics with high surface hydrophobicity limit the adhesion of microorganisms on the polymer surface, limit the formation of biofilm, and reduce the degradation of microplastics by microorganisms and enzymes in biofilm. At the same time, various additives contained in some micro plastic products, such as preservatives, antioxidants, plasticizers, flame retardants, curing agents, fungicides, heat stabilizers, etc., will also affect the biodegradability of micro plastics. For example, the nucleophilic bromide ion decomposed by flame retardant decabromodiphenyl ether at high temperature is easy to react with the positively charged group of polycarbonate (PC) chain, accelerating the depolymerization of PC in subcritical water; The non-polar part of the plasticizer di-n-octyl phthalate is embedded in the PC molecular chain, which inhibits the further diffusion of solvent molecules to the polymer and the hydrolysis of PC[27]。

2.2.2 environmental factor

Environmental factors play an important role in improving the effectiveness of microbial growth on polymer surface. Environmental factors such as light, temperature, humidity, oxygen, pH value, etc. will change the structure and properties of microplastics and affect the degradation of microplastics. At the same time, they will also affect the diversity, relative abundance, life activities and metabolic potential of microorganisms, thus affecting the degradation of microplastics by organisms[16,20]For example, low pH is conducive to the formation of hydrogen peroxide during photooxidation, thus promoting the degradation of microplastics[28]The factors such as solar radiation, high temperature, hydraulic impact, sand abrasion, weathering and fragmentation in the environment will accelerate the aging of microplastics, making the surface of microplastics more rough and reducing the hydrophobicity. At the same time, the carbonyl, carboxyl, hydroxyl and other oxygen-containing functional groups formed by aging increase the probability of microbial contact and colonization, so as to improve the degradation efficiency of microplastics[20]。

2.2.3 Biological factors

Biological factors include biological species, microplastic degrading enzymes and gene differences. The difference in the ability of bacteria and fungi to biodegrade micro plastics is due to the difference in the types and active sites of enzymes produced by different microorganisms, which affects the contact efficiency between enzymes and substrates, thus affecting the degradation of micro plastics by enzymes. On the other hand, extracellular polymers secreted by microorganisms can improve the adhesion ability of microorganisms on the surface of microplastics, thereby improving the degradation ability of microorganisms on microplastics. The long-term domestication of animals' intestines, plant roots and phyllospheric microorganisms in the hot spot of microplastics pollution formed the hot spot of microplastics degradation through long-term selection and evolution. Animals such as cockroaches, earthworms, barley worms, Tenebrio molitor, etc. can break the micro plastics by chewing, stomach grinding and other ways to change the physical and chemical properties of the micro plastics. At the same time, the intestinal microorganisms and enzymes secreted by them can oxidize the micro plastics, produce small molecular polymers, and promote the degradation of the micro plastics[13,29 -30]Some plant rhizosphere microorganisms, aquatic algae and the enzymes secreted by them can change the physical properties of microplastics and promote the degradation of microplastics. For example, in the respiratory roots of mangrove plantsAlternaria alternataSuch fungi can colonize and secrete laccase and manganese peroxidase on the surface of LDPE, destroy the structure of microplastics, produce low molecular weight polymer fragments, and promote the degradation of microplastics[31]In addition, the degradation effect of microflora composed of a variety of bacteria is often more obvious than that of a single strain. For example, a single degrading bacterium can reduce the weight of microplastics by 20%, while when four strains are used, the weight loss can reach nearly 50% within 140 days[32]。

3 Microplastic degrading bacteria and enzymes

The environmental micro plastic pollution is serious, which needs to be reduced and controlled. It is helpful to realize the targeted reduction and control of microplastics in different environments, improve the scientificity and accuracy of microplastics pollution prevention and control, and protect the ecological environment and human health by excavating the efficient and high colonization microplastics degradation resources in environmental microorganisms, deeply analyzing their metabolic pathways, degradation mechanisms and interactions, designing and constructing synthetic microflora containing a variety of specific functional microorganisms, optimizing the structure-activity relationship and analyzing their principles.

3.1 Microplastic degrading microorganisms

More than 416 microorganisms have been reported to have the potential to degrade microplastics[33]It is mainly distributed in Pseudomonas, actinomycetes, Firmicutes and Bacteroides[16,22]（Figure 2）. Actinomycetes, α - Proteaceae, β - Proteaceae, γ - Proteaceae, bacillus, flavobacteriaceae and other bacteria are the main plastic degradation microorganisms. Of which,Pseudomonas aeruginosa、Rhodococcus sp.、Amycolatopsis sp.、Aspergillus sp.、BacillusSp. andCladosporiumSp. is the main bacterial species that has been proved to be related to the degradation of microplastics[34]For example, Pseudomonas aeruginosa can degrade micro plastics such as PS, pet, PE and PVC, while bacillus can degrade micro plastics such as PS, pet, PE and PP[17,33,35]In recent years, the research on insect feeding on degradable plastics has provided a new research direction and treatment strategy for the efficient biodegradation and resource utilization of waste plastics. In the past decade, the intestinal microorganisms and enzymes of Lepidoptera and Coleoptera larvae, such as Tenebrio molitor, Spodoptera litura, Hordeum vulgare, Tenebrio SMUs, and pyralid borer, have been found to be able to degrade a variety of micro plastics, including PS, PE, PP, PVC, PLA, pet, etc[20,36]Among them, the main potential micro plastic degradation functional bacteria in insect intestines include Pseudomonas, spirulina, Enterococcus, Acinetobacter, etc[21,37]。
Figure 2 Microplastic degrading bacteria

Fig. 2 Microplastics degrading bacteria

In addition to bacteria, fungi also have the ability to degrade micro plastics. The filamentous branching structure of fungi has a strong ability to extend and penetrate, and the mycelia degrade microplastics by exocytosis. Fungi have powerful enzyme system, adsorption capacity and the ability to produce natural biosurfactants. They can use micro plastic substrates as a source of carbon and electrons to provide materials and energy for cells[38]At present, more than 256 fungi have been reported to be able to degrade microplastics, among which Ascomycota, Basidiomycota and Mucor are the main microplastics degrading fungi[33]（Figure 3）. For example, Rhizopus, Mucor, Aspergillus and other fungi can degrade PLA, and Mucor, Aspergillus flavus and Streptomyces strains are also involved in the degradation of starch based PE[23]Penicillium bacteria can degrade a variety of micro plastics, including PLA, PU, PE, PCL, PHA, PP, etc[33]。
Figure 3 Microplastic degrading fungi

Fig. 3 Microplastics degrading fungi

It is worth mentioning that although a large number of microplasticizer degrading bacteria have been screened out from the environmental microbial community, the microplasticizer degradation ability of some microorganisms has not been fully verified due to the differences in characterization methods and experimental operations. Even if the degradation ability has been verified, the degradation effect of micro plastic degradation bacteria in the actual environmental application is not ideal, and it still faces the challenges of weak colonization ability, poor stability of key degradation enzymes and low degradation efficiency[39]In addition, the vast majority of environmental microorganisms (>90%) can not be isolated and cultured in the laboratory environment[40]The main limiting factors include unknown metabolic needs, unknown physiological status and low species abundance. However, many uncultured microorganisms may play an important role in the in-situ environmental degradation of microplastics. Moreover, in-situ degradation may involve the division and coordination of multiple microorganisms[41]It is difficult to effectively degrade and fully reduce the micro plastic pollution in the environment by using a single strain[42]Finally, microplastics are easy to absorb toxic pollutants from the surrounding environment, which may be toxic to microplastics degrading microorganisms[43]So as to inhibit its degradation activity. Therefore, it is urgent to use culture independent technology to find strains or flora resources with in-situ high-efficiency microplastics degradation ability and colonization in polluted environment.

3.2 Microplastic degrading enzyme

Enzyme is an efficient biocatalyst, which has many advantages compared with chemical processes, including: (1) high efficiency catalytic ability, making each basic reaction step speed up to 1020times[44](2) low environmental impact, reducing energy consumption and waste generation[44]Enzymes are considered to be renewable and biodegradable catalysts, which can be achieved through less expensive and environmentally friendly fermentation processes. At present, the commonly used databases of plastic degradation genes and enzymes include the Kyoto Encyclopedia of genes and genomes (KEGG)[45]Carbohydrate active enzyme database (cazy)[46], metabolic pathways and enzymes database (metacyc)[47], microbial metabolism and genome database (biocyc)[48]Plastic microbial degradation database (PMDB)[49], plastics degrading microorganisms and proteins database (plasticdb)[33]And bioremediation enzyme prediction database (remedb)[50]Etc. These databases can provide information on microbial metabolic pathways, microbial enzymes and functional genes for the degradation of microplastics, so as to evaluate the biodegradation process. At present, more than 20 enzymes have been reported to participate in the biodegradation of a variety of micro plastics, including laccase, esterase, hydrolase, hydroxylase, dehydrogenase, monooxygenase, oxidoreductase, peroxidase, chitinase, keratinase, lipase, etc[33]（Figure 4）. Enzyme mediated depolymerization is one of the initial key steps in the biodegradation of microplastics[51]Depolymerizing enzymes such as PLA depolymerizing enzyme, PCL depolymerizing enzyme, PHA depolymerizing enzyme, etc., can catalyze the hydrolysis of polymer skeleton, resulting in the cracking of polymer into oligomers and monomers, making the microplastics easier to be contacted by other enzymes[44]Oxidases such as laccase, oxygenase, monooxygenase, peroxidase, etc. promote the oxidation of complex microplastics molecules and decompose them into smaller molecules or produce soluble compounds through oxidation, so as to promote the further degradation of microplastics[21,52]For example, laccase can oxidize polymer chains of micro plastics such as PS, PE, PVA, etc., monooxygenase can cleave aromatic rings of PS by cleavage of main chain and side chain, and lignin peroxidase and manganese peroxidase play an important role in the degradation of PE. Hydroxylase can catalyze hydroxylation reaction, increase the hydrophilicity of microplastics, make the substrate more soluble in water, and be further degraded and metabolized by microorganisms[44,53-54]Hydrolases can further accelerate the decomposition of polymer chains into smaller and more easily metabolized components through hydrolysis reaction, and play an important role in the biodegradation of microplastics. It has been reported that hydrolases have the ability to degrade plastics such as PCL, pet, PLA, PA and PHA[55]E.g. e.osaka（Ideonella sakaiensis）The two hydrolases petase and mhetase can hydrolyze pet and the intermediate monohydroxyethyl terephthalate to produce terephthalic acid and ethylene glycol, respectively[10]Dehydrogenase is also essential for the microbial metabolism of decomposition products produced by the degradation of microplastics. When microplastics are decomposed into smaller organic molecules by other enzymes, dehydrogenase is responsible for catalyzing the oxidation of these molecules to produce reducing coenzymes (NADH or NADPH), which can then be used in the electron transfer chain to produce ATP. This process allows microorganisms to obtain energy from the decomposition products of microplastics to promote their growth and microplastics degradation process[56]。
Figure 4 Microplastic degrading enzyme

Fig. 4 Microplastics degrading enzymes

Microplastic degrading enzymes are closely related to natural polymer degrading enzymes. It has been reported that there are highly similar pathways for the degradation of PS and lignin[21]The microplastic degradation ability of enzymes may be derived from their long-term selective adaptation, evolution and evolution to the degradation of natural polymers, thus helping microorganisms and their hosts use specific nutrients (e.g., microplastic feeding insect larvae) and adapt to environmental changes (e.g., microplastic pollution)[52]Therefore, more natural polymer degrading enzymes need to further explore their potential for the degradation of micro plastics.

3.3 Synthetic flora

Synthetic flora is a specific functional flora co cultured by multiple synthetic species. Due to the metabolic interaction and functional complementarity between flora, compared with a single flora, synthetic flora has the characteristics of strong colonization ability, good functionality and strong environmental adaptability. In the field of microplastics degradation, synthetic bacteria can be designed to carry specific enzymes, or they can be designed to have synergy and cooperate with each other to effectively degrade specific types of microplastics, so as to help solve the problem of microplastics pollution. For example, thiamine degrading Bacillus thiamine isolated from activated sludge of sewage treatment plants and landfill sites（Aneurinilyticillus aneurinilyticus）Soil Bacillus brevis（Brevibacillus agri）Bacillus brevis（Brevibacillus Sp.) and Brevibacillus brevis（Brevibacillus brevis）The results showed that the mass loss of LDPE, HDPE and PP was 58.2%, 46.6% and 56.3% respectively within 140 days, while the mass loss of microplastics was only 22.2%~28.2% when the strain degraded alone[32]In addition, studies have shown that the synergistic degradation of polymer by marine microbial flora has better degradation effect[57]Similarly, the synthetic bacterial community constructed by using the micro plastic degradation strains isolated from the plastic waste treatment plant showed higher PE degradation efficiency than the single strain[58]In addition, the reasonable configuration of synthetic bacteria can use micro plastics to produce other high value-added products. For example, qiqingsheng and others designed pet degrading bacteria（Yarrowia lipolytica Polf）Bacteria producing polyhydroxybutyrate (PHB)（Pseudomonas stutzeri）The biodegradable plastic PHB with economic value can be synthesized at the same time of degradation of pet[59]。
Based on the functional quantification and targeted design of the synthetic microflora, the functional microflora for degradation and synthesis of microplastics was customized, and its construction mechanism and functional principle were analyzed, which could provide a new strategy for the enhancement and targeted reduction of microplastics, and realize the prevention and control of environmental microplastics pollution. At the same time, it is necessary to pay attention to biological safety and environmental risks in the process of using synthetic flora to ensure the safety and sustainability of its application.

4 Mining methods of micro plastic degradation microbial resources

Microplastics degrading microorganisms can convert environmental microplastics into harmless substances such as carbon dioxide and water or other high-value intermediates, so as to realize the harmless and resource utilization of environmental microplastics. The development of efficient and sensitive micro plastic degradation microbial resource mining methods and Characterization Technologies will help to further explore micro plastic degradation microbial resources and screen in-situ high-efficiency degradation microorganisms, which is of great significance for environmental micro plastic pollution control. The main methods are divided into dependent and independent cultivation（Figure 5）。
Figure 5 Mining methods of micro plastic degradation resources

Fig. 5 Mining strategies of microplastics degradation resources

4.1 Culture based approach

The isolation and cultivation of microorganisms is a classic method of microbial resource mining, and it is also the most critical step to understand and use microorganisms. Microorganisms have evolved in different niches in the natural environment for a long time, forming rich and diverse genetic resources, which have important application potential in the remediation of polluted environment. Microplastics degrading microorganisms are generally isolated and cultured from the hot areas of microplastics pollution in the environment, including agricultural plastic film soil, marine sediments, garbage collection stations, insect intestines, etc. The traditional isolation and culture process includes inoculating environmental samples enriched with microplastics degrading microorganisms into a selective medium with microplastics as the sole carbon source, enriching and subculturing them under appropriate culture conditions, and then separating and purifying microorganisms with potential plastic degradation ability by gradient dilution method or plate scribe method, and then verifying the plastic degradation ability of the purified degrading microorganisms, and analyzing their phylogenetic species based on 16S rRNA gene sequence[10,60-61]At present, many researchers use this method to isolate and obtain degradation strains that can use micro plastics as the sole carbon source and energy, and study its degradation pathway and molecular mechanism. Yoshida et al[10]Pet degrading bacteria which can use pet as the main energy and carbon source were isolated and cultured from the waste recycling stationIdeonella sakaiensis201-f6, and pet hydrolase, which plays a key role in degradation, was found to reveal its degradation mechanism. Ju Feng et al[52]A polyvinyl chloride degrading bacterium isolated from the gut of insect larvaeKlebsiellasp. EMBL-1， It can depolymerize and use PVC as the only energy source.

4.2 Culture independent approach

4.2.1 Omics technology and big data algorithm

In recent years, with the progress of sequencing and detection technology and the reduction of cost, the wide application of omics technology provides a technical means independent of pure culture for studying and understanding the composition and function of microorganisms in complex environment. Metagenomic technology provides a large amount of information for in-situ understanding of the composition of environmental microbial communities and the functional genes of micro plastic degradation through high-throughput sequencing of environmental samples' DNA, and then splicing, annotation and analysis of gene sequences[62⇓-64]For example, Zhou et al[63]Metagenomic technology was used to study the microplastic degradation genes in the subtropical estuarine environment of China. It was found that human activities significantly affected the abundance of plastic degradation genes, and metagenomic box technology was used to reveal that a variety of microbial groups have microplastic degradation potential. By sequencing the transcripts of environmental samples, macrotranscriptome technology can study the degradation activity of active microorganisms and micro plastics in the environment, and compare the differentially expressed genes and degradation pathways under different conditions. Mamtimin et al[21]Macrotranscriptome sequencing of Tenebrio molitor intestinal microorganisms showed that the pathway of PS and lignin degradation was highly similar, indicating that the plastic degradation ability of insects may be due to their long-term selective adaptation to the degradation of natural lignin.
The integration of genomics, transcriptomics, proteomics, metabonomics and other omics data can provide a more comprehensive understanding of the degradation process and mechanism of microplastics[65]. Wright et al[61]The succession process of pet degradation by marine microplastics microbial community was studied by using multiomics technology. Two pet degrading bacteria were isolated from the microbial community and the degradation mechanism was further revealed. Meyer Cifuentes et al[41]The degradation process of polyadipic acid (PBAT) by marine microbial enrichment was analyzed by multi omics integration. It was found that the degradation was completed by the division and cooperation of multiple microorganisms. It was revealed that the depolymerization process was mainly completed by the attached biofilm community, and all microbial communities were involved in the degradation process of monomer substances. Although multiomics technology reflects the degradation potential of microplastics, it also provides new insights for the cultivation of microplastics degrading microorganisms in an in-situ environment. Most microorganisms in the natural environment are still difficult to isolate and cultivate. The metagenomic box tool can assemble the genome sketch of environmental micro plastic degrading microorganisms based on the sequencing results, so as to analyze and predict their growth conditions, nutritional needs, metabolic physiology, etc[66⇓-68]It provides the basis for cultivating the key strains of micro plastic degradation in complex communities, and brings hope for using the rich microbial genetic resources of uncultured microorganisms.
The difficulty of omics technology lies in bioinformatics analysis, and it relies heavily on the existing information in the database. The diversity of microorganisms in the natural environment is very high, while the known genome information in the database is relatively limited, and the speed of genome analysis algorithm and software development lags far behind the development of sequencing technology and a large number of sequencing data[42]In addition, the information in the database is helpful to understand the metabolic pathways of microorganisms, so that they can be isolated and cultured by changing the culture conditions[67-68]Big data, combined with artificial intelligence and machine learning algorithms, opens up a new prospect for tapping the functional potential and mechanism of micro plastic degrading microorganisms. According to the existing information of genes or enzymes related to the degradation of micro plastics in the database, the algorithm can be used to predict the potential degradation gene resources in the microbial sequencing data. For example, Sankara Subramanian et al[50]Based on the hidden Markov model (HMM) search algorithm, the remedb tool was developed to simplify the identification of micro plastic degradation microorganisms. Danso et al[69]Based on HMM algorithm, 504 genes of potential PET hydrolases were searched from the database, and 4 of them were verified. Using similar ideas, Tan et al[70]A dioxygenase his1, which can degrade polypropylene, was identified and heterologously expressed, and its degradation function was verified. In addition, using the global genome database to obtain the information of degradation genes and potential degradation microorganisms can promote the understanding of the core degradation flora and its degradation mode. Qianhaifeng et al[71]Using machine learning model to predict and draw the map of global marine microplastics degradation potential, it is found that there is a high potential for microbial degradation in areas with serious microplastics pollution, and Proteus is the main group of microplastics degradation bacteria.

4.2.2 Nanoscale secondary ion mass spectrometry

The study of environmental microorganisms at the single cell level can overcome the limitation of pure culture and achieve in situ functional research in complex environmental media. Due to the limitations of methodology, the current methods to determine the degradation of microplastics by microorganisms, such as measuring the weight loss of microplastics, determining the surface oxidation by infrared spectroscopy, and observing the surface morphology by electron microscopy, are indirect methods. It is difficult to determine whether the degradation of microplastics is caused by microbial processes or non biological processes. Moreover, these methods are not sensitive and direct enough, and need time-consuming experimental process. In addition, the existing methods are difficult to track the process from the carbon of microplastics to degradation products or microbial biomass. Nanosims is the analytical technology with the highest spatial resolution and sensitivity in the field of environmental microbiology. The principle is to use the primary ion beam to bombard the sample surface to produce a series of secondary ions for detection, so as to achieve high-resolution imaging of the elemental composition and isotopic distribution of the sample. At present, vaksmaa et al[72]utilize13C-labeled non degradable plastic PE and its application in marine environmentRhodotorula mucilaginosaCo incubation with yeast proved that UV aging13C-pe can be assimilated into its own biomass within 5 days, and its plastic degradation rate is estimated to be about 3.8% of its total mass per year. In addition, sander et al[73]utilize13C stable isotope labeled biodegradable plastic PBAT was co incubated with soil, and the incubated samples were imaged by nanosims. It was found that PBAT could be assimilated by filamentous fungi in soil to synthesize its own biomass, and the mineralization rate of PBAT by microorganisms during 42 days of incubation was determined13CO2Production. However, because nanosims is a destructive test, it is unable to carry out important downstream research on the detected micro plastic degradation microorganisms, such as sorting, sequencing and even subsequent culture. It is difficult to analyze the metabolic path of degradation microorganisms, which limits the in-depth understanding and utilization of degradation functional microorganisms.

4.2.3 Fluorescent labeling technology

In recent years, because of its high sensitivity and high-throughput detection, fluorescence analysis methods are increasingly used to screen microorganisms or enzymes that degrade micro plastics. The key problem of fluorescent screening is how to transform the degradation function of microplastics into fluorescent signal. At present, the fluorescent detection of the degradation of micro plastics is mainly focused on polyester plastics. Esterase can be used to screen esterase by the specific hydrolysis of carboxylate ester bond and the release of fluorescent probe. The principle is to modify fluorescent groups on small molecular esters of micro plastic monomers[74-75]The fluorescence is quenched during normal covalent bonding, and the fluorescent molecules will be released only after being hydrolyzed by enzyme, thus generating fluorescent signals. Zumstein et al[76]Fluorescein dilaurate was embedded into polyester matrix, and the hydrolysis process of polyesterase was characterized by detecting the production of fluorescein. In addition, using this principle and microfluidic platform, microorganisms or hydrolases with degradation function can be screened at the single cell level, thus providing a new technical means for mining the degradation resources of micro plastics. Duwenbin et al[75]A fluorescent activated droplet sorting platform was developed for high-throughput screening of pet degrading microorganisms or enzymes (petase). A single microorganism was dispersed in a single droplet and incubated with pet simulated substrate fluorescein dibenzoate. Petase could hydrolyze the substrate and send fluorescence signals to separate it in high flux. Using this method, nine pet degrading bacteria of different genera were obtained from the waste water samples of PET textile factory, and two potential PET degrading enzymes were obtained from the high activity strains, and the degradation function was verified by heterologous expression[75]. Xu et al[74]A polyurethane analog fluorescent probe was proposed. Naphthalimide derivative molecules were modified on both sides of polyester monomer. When degraded by polyesterase, the two probe molecules released fluorescence. In addition, the possibility of screening polyester degrading microorganisms or enzymes was verified by droplet microfluidic control. Although the above schemes provide new high-throughput technical means for exploring microorganisms and enzymes that degrade microplastics, the current strategy depends on the design and synthesis of fluorescent probes and plastic monomers.

4.2.4 Single cell Raman technique

Raman spectrum is a kind of molecular vibration spectrum, which can provide the phenotypic fingerprint information of bacteria at the single cell level, such as protein, nucleic acid, lipid, carbohydrate and pigment molecules. When combined with stable isotope labeling (e.g13C、2D、15N) After the microorganism assimilates the isotope labeled substrate, the heavy isotope of the newly synthesized biomolecule replaces the light isotope of the original biomolecule, resulting in the red shift of the corresponding Raman spectrum peak, which can effectively analyze the functional activity of a single microorganism in the in-situ environment[77⇓⇓⇓-81]Single cell Raman spectroscopy combined with stable isotope labeling has been used to study the in situ degradation of pollutants in complex environments. For example, Huang Wei et al. Used single cell Raman binding13C-labelled polycyclic aromatic hydrocarbons (PAHs) were analyzed by in-situ degradation bacteria in groundwater samples at the single cell level, and different species and in-situ degradation activities were linked by fluorescence in situ hybridization[82]It was found that uncultured bacteria played a key role in the in situ environmental degradation of polycyclic aromatic hydrocarbons[83]In addition, for systems without suitable stable isotope labeled substrates, single-cell Raman spectroscopy can be combined with heavy water labeling to study metabolically active bacteria. For example, in the soil with only fixed phosphorus, after incubation with heavy water, the activity of microorganisms with phosphate solubilizing function is significantly higher than that of non phosphate solubilizing microorganisms, producing characteristic C-D Raman peaks, while the low or inactive non phosphate solubilizing microorganisms have weak or no C-D peaks. Identification of protophosphate solubilizing bacteria in soil environment can be realized by C-D Raman marker peak[84]Based on similar principles, this method can also be applied to the study of cellulose degrading bacteria. When cellulose is the only carbon source, the heterogeneity of cellulose degrading bacteria 'degradation activity can be revealed at the single cell level[85]In addition, single cell Raman and heavy water labeling can be used to identify antibiotic resistant bacteria (requiring antibiotic pressure) and metabolically active bacteria in the microplastic biofilm in situ, revealing the metabolic activities and physiological status of microorganisms in the microplastic biofilm[86-87]。
Raman spectroscopy is a non-destructive detection. Combined with single cell sorting, it provides a new strategy of "separation before culture". It can carry out downstream directional sorting, sequencing and even culture of the target functional bacteria identified by Raman spectroscopy, and realize the joint analysis of phenotype and genotype[88]At present, based on the single-cell Raman sorting platform, the single-cell sorting and targeted metagenome research of antibiotic resistant bacteria in soil identified by Raman have been realized[89], and single cell Raman recognition of environmental phosphate solubilizing bacteria[84]And sorting culture[90]Although there is no report of using single cell Raman to identify micro plastic degrading bacteria, these cases show that this technology has great potential in tapping micro plastic degrading microbial resources.

4.2.5 Synthetic biology technology

Because some microplastics degrading bacteria have drug resistance genes or virulence factors, there may be potential ecological risks when they are directly applied to the environment. It is an important development direction to construct safer engineering bacteria for the scale production of key degrading enzymes and in-situ environmental remediation of microplastics through genetic engineering. With the development of molecular biology and genetic engineering technology, significant progress has been made in the development of new micro plastic efficient degradation microorganisms through gene manipulation[91]Through heterologous expression of the gene for the synthesis of microplasticizer degrading enzyme in engineering bacteria, the high-efficiency microplasticizer degrading engineering bacteria can be constructed for plastic degradation. For example, Zhu et al[14]The petase gene was introduced into E. coli by genetic engineering method for high-efficiency expression, and good degradation effect of PET was achieved in the simulated effluent environment of sewage treatment plant. Hu et al[92]Further, the coexpression system of adhesion protein and petase degrading enzyme was constructed in E. coli, and the pet degradation efficiency was improved by improving the adhesion between E. coli and pet. Tournier et al[15]By modifying the degrading enzyme itself to make it more resistant to high temperature, another solution to improve the degradation efficiency was proposed. At low temperature, pet is in crystalline state, and it is difficult for degradation enzymes to effectively combine with it and "digest". At high temperature of 65 ℃, pet can soften to form amorphous state, making it easier for degradation enzymes to interact with pet. However, the enzyme is easy to be inactivated at high temperature, and the genetically modified enzyme is redesigned to improve its thermal stability, so as to achieve efficient degradation of pet at high temperature. In addition, Alper et al[93]The machine learning system was used to predict the effect of the mutation of petase enzyme on its performance. Through the engineering transformation and testing of the mutant, an enzyme with significantly improved thermal stability and degradation activity was found, and the potential way of using degradation enzyme for micro plastic recycling on an industrial scale was proposed. In a word, the development of genetic engineering and artificial intelligence technology enables people to redesign the structure and function of specific enzymes involved in the degradation of microplastics and express the key degradation enzymes efficiently in combination with engineering bacteria. In the future, the combination of bioinformatics, artificial intelligence, synthetic biology and other interdisciplinary technologies will help to find green, efficient and sustainable biodegradation solutions for microplastics.

5 Conclusion and Prospect

Environmental microorganisms are an important driving force for the material cycle in the ecosystem, and have important potential to solve the increasingly severe environmental micro plastic pollution. Although the microbial degradation of microplastics has made some important progress, it still faces many challenges.
(1) Understand the microbial degradation law of microplastics: Although there are potential degradation bacteria in the environmental microplastics biofilm, the degradation efficiency is low and it is difficult to use directly. By analyzing the assembly mechanism and degradation law of degradation bacteria in microplastics, the core degradation microorganisms can be identified in time and space scale, which can promote the efficient reduction and control of environmental microplastics. In addition, in the future, attention should be paid to the effects of environmental factors such as pH, humidity and temperature on the structural characteristics and spatial distribution of micro plastic degradation bacteria, so as to provide a theoretical basis for optimizing the degradation efficiency.
(2) Development of in-situ analysis methods for the degradation of microplastics: due to the widespread presence of additives in microplastics, many existing indirect research methods for the degradation of plastics are not rigorous. Therefore, it is necessary to further standardize the characterization and analysis of the degradation process, and constantly improve the research field of microbial degradation of microplastics. The development of highly sensitive in-situ analysis methods and characterization techniques provides important technical support for in-depth understanding of the in-situ degradation process and molecular mechanism of microplastics, and mining efficient in-situ microbial degradation resources.
(3) Excavate in-situ efficient degradation microbial resources: Currently, the microbial resources for the degradation of micro plastics are still very insufficient in terms of the number of strains and degradation efficiency. At present, the rapid development of omics technology, single cell technology, artificial intelligence algorithm, synthetic biology and other technologies provides a series of powerful tools for the discovery, design and acquisition of micro plastic degradation genes and enzymes, which will greatly promote the mining of efficient degradation resources in various in-situ environments. In addition, through the artificial construction of synthetic bacteria, the microbial community with stable species composition, high degradation ability and strong environmental adaptability can be formed by using the metabolic interaction and functional complementarity between bacteria, and the synthesis of high value products can be used, which is expected to truly realize the green reduction of micro plastic pollution and turn waste into treasure.

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