Biodegradation: The Best Solution To The World Problem of Discarded Polymers
Biodegradation: The Best Solution To The World Problem of Discarded Polymers
Abstract
The widespread use of polymers has made our lives increasingly convenient by offering a more convenient and
dependable material. However, the challenge of efficiently decomposing these materials has resulted in a surge
of polymer waste, posing environment and health risk. Currently, landfill and incineration treatment approaches
have notable shortcomings, prompting a shift towards more eco-friendly and sustainable biodegradation
approaches. Biodegradation primarily relies on microorganisms, with research focusing on both solitary bacterial
strain and multi-strain communities for polymer biodegradation. Furthermore, directed evolution and rational
design of enzyme have significantly contributed to the polymer biodegradation process. However, previous
reviews often undervaluing the role of multi-strain communities. In this review, we assess the current state of
these three significant fields of research, provide practical solutions to issues with polymer biodegradation, and
outline potential future directions for the subject. Ultimately, biodegradation, whether facilitated by single bacteria,
multi-strain communities, or engineered enzymes, now represents the most effective method for managing waste
polymers.
Keywords Biodegradation, Polymers, Single-strain degradation, Multi-strain degradation, Enzyme engineering
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Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 2 of 16
Graphical Abstract
incineration, a process that releases hazardous chemi- to recycling efforts and diminish the reliance on fossil
cals into the atmosphere, including carbon dioxide, sul- fuels. These monomers have the capability to polymerize
fur dioxide, and dioxins (Zhao et al. 2020). In addition, into new macromolecular polymers, boosting recovery
incineration produces a large amount of hazardous and rates by 40% (Rajendran and Han 2022). The establish-
poisonous dust and slag compounds, which pose a sig- ment of a “recycle-generate-reuse” green cycle is made
nificant threat to the environment (Duval 2014; Wang et possible by employing biodegradable treatments in poly-
al. 2019a; Yan et al. 2023). The main destinations of the mer recycling, as depicted in Fig. 2. In conclusion, the
polymers now are shown in Fig. 1. most effective and environmentally sound approach to
Previous researches have shown that biodegradation handle waste polymers is through biodegradation.
stands out as a sustainable and ecologically friendly pro- Various microorganisms have undergone trails for
cess for handling waste polymers. Utilizing biological polymer degradation over the past few decades. Despite
enzymes and microbial strains, biodegradation converts efforts to identify more effective strains through screen-
waste polymers into smaller molecules or even mono- ing, most individual wild strains exhibit limited deg-
mers. Biodegradation, when contrasted with traditional radation capabilities. Aiming to achieve efficient
landfill methods, has the ability to preserve valuable biodegradation of diverse polymers, multi-strain com-
land resources by reducing solid volume by up to 80% munities have been proposed as an alternative to indi-
(Mostafa et al. 2018; Rai et al. 2021). Moreover, biodeg- vidual strains, as communities offer greater resilience to
radation exhibits a superior capacity for degradation. deterioration and higher efficiency. Furthermore, there
The rapid decomposition of the polyester polymer poly- is growing interest in enhancing the ability of strains to
ethylene terephthalate (PET) exemplifies the high effi- degrade polymers, which can be achieved by utilizing
ciency of biodegradation. Contrary to the lengthy natural enzyme engineering to modify their catalytic enzymes
deterioration process that takes place in landfills over and increase their catalytic efficiency. Compared with the
several decades, biodegradation can rapidly disintegrate previous review (Kotova et al. 2021; Pathak and Navneet
PET within a few days (DelRe et al. 2021). Furthermore, 2017), this review offers a comprehensive introduction
biodegradation plays a crucial role in reducing environ- to the three key areas of biodegradation, and provides a
mental pollutants. It can eliminate harmful gases, such rounded analysis of all categories of polymers. The analy-
as dioxins, and reduce greenhouse gas emissions by 60% sis traces the progression and transformation of poly-
when compared to incineration (Rajendran and Han mer degradation, starting from the examination of wild
2022). Beyond waste reduction, biodegradable polymer individual strains to communities and then to genetically
monomers generated through this process can contribute modified strains. The factors driving the advancement
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 3 of 16
trials (Altenhoff et al. 2021; Basik et al. 2022; Sarkar et it was believed that only Gram-positive bacteria were
al. 2021; Schmitt et al. 2019). By employing this method, capable of degrading natural rubber since all mentioned
they successfully isolated 50 species of bacteria capable rubber-degrading strains were identified as such. These
of degrading rubber from natural environments. Addi- strains that degrade rubber contain the lcp gene or its
tionally, another 46 species of rubber-degrading bacteria homologous sequence, which codes for the latex cleavage
were identified through screening multiple strain collec- protein (LCP). LCP is recognized as a crucial enzyme for
tion centers (Schmitt et al. 2019). After incubating these degrading natural rubber, as the insertion of the lcp gene
selected strains in medium with NR as a carbon source enables Escherichia coli to degrade natural rubber (Basik
for 30 days, the weight of NR was decreased by 10–30% et al. 2022). Nevertheless, Sharma’s discovery of Steroido-
and the average molecular weight of polymer decreased bacter cummioxidans strain 35Y challenged the perspec-
from 640 kDa to 25 kDa (Schmitt et al. 2019). All of the tive that only Gram-positive bacteria have the ability to
isolated strains were identified as Actinomyces spp. They degrade rubber. This strain was demonstrated to be the
were easy to distinguish during the initial screening pro- most effective among all known strains. Within seven
cedure because they could produce hyphae and create days, this particular strain led to a 60% decrease in NR
transparent areas when decomposing rubber on agar (Cui et al. 2023; Sharma et al. 2018). Imai et al. identified
plates with NR. Screening bacteria for rubber degrada- and separated three additional gram-negative bacteria
tion by this screening process can be challenging when capable of rubber degradation: Streptomyces sp. LCIC4,
the identifying bacteria exhibit no obvious indications Actinomyces sp. OR16, and Methylating bacteria sp.
of reaction. Actinomycetes, such as Nocardia and Myco- NS21. LCIC4 exhibited the highest rubber degradation
bacterium, as well as non-actinomycetes like Corynebac- capability, degrading 70% of NR in just 50 days. This deg-
terium, do not form a transparent areas when degrading radation resulted in an average reduction in molecular
rubber. This makes it more difficult to screen for these weight of rubber from 400 to 23 kDa (Imai et al. 2011).
strains (Basik et al. 2021; Prakash et al. 2024). Previously, The identification of rubber oxygenase (RoxA), a key
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 5 of 16
enzyme required for the degradation of NR in gram-neg- et al. discovered that Bacillus subtilis, Pseudomonas aeru-
ative bacteria, was first reported in Xanthomonas 35Y. ginosa, and Streptomyces possess the capacity to degrade
The enzymes RoxA and LCP play a crucial role in the vulcanized rubber. The utilization of these three strains
degradation of NR by breaking the double bond between on vulcanized rubber resulted in a decrease in cross-
rubber monomers (cis-1,4-polyisoprene) through oxida- linking within the vulcanized structure by 17.2%, 10.7%,
tive cleavage (Suzuki et al. 2022). and 43.4%, and a reduction in the cleavage of C-C bonds
Vulcanization is a commonly used process to enhance by 16.1%, 16.8%, and 18.1%, respectively, as compared
the stability and optimize the properties of natural rub- to the control. The data suggest that the desulfuriza-
ber. Nevertheless, microbial strains capable of degrading tion reaction and rubber degradation occurred simul-
NR are unable to directly degrade vulcanized rubber due taneously, which means that the vulcanized rubber was
to their inability to break the C-S or S-S bonds present directly degraded (Aboelkheir et al. 2019). Ceriporiopsis
in vulcanized rubber. In other words, these strains lack subvermispora, a type of White-rot basidiomycetes, dem-
the capacity for desulfurization. These strains can only onstrates superior efficiency in both desulfurizing and
participate in the degradation of vulcanized rubber after degradation of the main structure of vulcanized rubber
it has been desulfurized. Consequently, the initial pro- compared to other organisms. Exposure of vulcanized
cess of degrading vulcanized rubber can be split into two rubber to Ceriporiopsis subvermispora for 200 days led
distinct steps: desulphurization and degradation. The to a 29% decrease in sulfur content and a 69% decrease
elimination of sulfur from vulcanized rubber is mostly in the frequency of S-C bonds. In contrast, the control
accomplished by the use of various species of Thiobacil- group showed no degradation of S-C bonds or removal of
lus bacteria. For instance, the content of sulfur in vulca- sulfur (Chen et al. 2020). The main microorganisms capa-
nized rubber decreases by 30% after undergoing a 30 days ble of degrading rubber and their degradation capabilities
treatment with Thiobacteria (Calabrese et al. 2021). Sub- are summarized in Table 1.
sequently, it is necessary to introduce microorganisms
that can degrade rubber in order to dismantle the pri- Degradation of the plastic family
mary chain structure. This complicates the degradation Plastic is a broad term that encompasses various poly-
process of vulcanized rubber. To simplify and optimize mers, such as polyethylene (PE), polypropylene (PP),
the biodegradation of vulcanized rubber, it would be polyvinyl chloride (PVC), and polystyrene (PS). These
advantageous to have a single strain capable of achieving materials have taken the place of wood, glass, and metal
both desulfurization and degradation. Further research in several applications. Plastics, renowned for their dura-
revealed that certain microorganisms have the ability bility, were once believed to be non-biodegradable unless
to carry out desulfurization and degradation processes subjected to certain treatments. However, the pretreat-
simultaneously. This means that a singular species of ment of plastics is a complicated and energy-intensive
microbe can decompose vulcanized rubber. Aboelkheir
procedure, so it is necessary to investigate strains that There have been few attempts to biodegrade PVC due
can directly degrade polymers. to its high stability. Only a small number of microorgan-
PE is a highly prevalent thermoplastic material. Early isms have demonstrated the capability to degrade PVC
studies have indicated that certain types of bacteria, (Giacomucci et al. 2019; Shilpa et al. 2022). White-rot
such as Bacillus, Rhodococcus, and Pseudomonas, as well basidiomycetes degrade low-molecular-weight PVC in
as fungi like Aspergillus and Fusarium, are capable of circumstances under limited nutrients conditions (Chow
degrading polyethylene when exposed to ultraviolet (UV) et al. 2023). Both Pseudomonas citronellolis and Bacillus
radiation or heat treatment (Gómez-Méndez et al. 2021; flexus demonstrate depolymerization activity on PVC
Soong et al. 2023; Sun et al. 2022a). Recent studies have films (Giacomucci et al. 2019). The combination of Asco-
identified several microorganisms capable of degrading mycetes and Chaetomium globosum (ATCC 16021) has
PE without the need for any special treatment, includ- demonstrated the capacity to degrade PVC (Vivi et al.
ing Pseudomonas putida IRN22, Acinetobacter peddler 2019). Although these strains exhibited the capacity to
IRN19, Micrococcus luteus IRN20, Pseudomonas aerugi- degrade PVC, the extent of degradation is limited. Nev-
nosa PAO1, Pseudomonas aeruginosa ATCC, Pseudomo- ertheless, the co-cultivation of Pseudomonas citronello-
nas putida, Pseudomonas syringae, Pseudomonas sp. E4, lis and Bacillus flexus on a PVC sheet exhibited robust
Comamonas, Delftia, and Maltophilia (Meyer Cifuentes degrading activity. After a 45 days treatment period, the
et al. 2023; Montazer et al. 2019; Peixoto et al. 2017; Wei average molecular weight of the PVC was decreased by
et al. 2022). Nevertheless, their ability to degrade PE is 10%, accompanied by a weight loss of 19% approximately
limited, resulting in less than a 1% loss in PE weight. This (Giacomucci et al. 2019). This is the sole example demon-
presents a challenge for the efficient polymer degrada- strating a greater effect of biodegradation treatment on
tion and recycling. In the following studies, four strains, PVC.
were isolated from the marine environment: Cobetia PS is a thermoplastic material that has exceptional
sp. H-237, Halomonas sp. H-255, Exigobacterium sp. optical and chemical properties. It is worth noting that
H-256, and Alcanivorax sp. H-265. All of these bacte- this type of plastic is highly biodegradable and can
ria exhibited the ability to degrade substances. Among undergo degradation without any prior treatment. Geo-
these strains, H-255 had the most potent ability, resulting bacillus stearothermophilus FAFUA011, Bacillus cereus,
in a weight loss of 1.7% in PE after 90 days of treatment and Bacillus gottheilii cereus are capable of thriving on
(Khandare et al. 2021). A separate team of scientists has PS, which was the only available carbon source (Xing et
discovered three additional bacteria, including Kocuria al. 2021). Treatment of PS with actinomycetes (Rhodo-
palustris M16, Bacillus pumilus M27, and Bacillus sub- coccus ruber sp. C208) resulted in a weight reduction of
tilis H1584, which exhibit superior degrading capabili- 0.8% over 8 weeks (Sun et al. 2022b). After undergoing
ties in the pelagic waters of the Indian Ocean. The weight a 56 days treatment with Geobacillus stearothermophi-
loss of PE after 30 days of treatment with these three lus FAFUA011, PS exhibited a 4.2% decrease in mass
strains was 1.5%, 1.7%, and 1.8%, respectively, while the and a decrease in the average molecular weight ranging
crystallinity dropped by 10.3%, 8.6%, and 4.6%, respec- from 17.4 to 18.2% (Xing et al. 2021). The weight of PS
tively (Harshvardhan and Jha 2013). Up to now, Bacillus decreased 5.8% and 7.4% after being treated with Bacil-
subtilis H1584 has been proven to be the most effective lus cereus and Bacillus gottheilii for 40 days, respectively
strain for degrading PE directly. While these strains had (Auta et al. 2017). To a certain extent, PS is one of the
limited degradation capacity, with none of them causing most easily degradable plastics.
PE weight losses above 2%, this finding is significant since In short, the majority of plastics are not biodegradable
it challenges the conventional belief that PE is not inher- unless they are pre-treated. Additionally, the requirement
ently biodegradable. for pretreatment can increase the intricacies and costs of
Similar to PE, PP is also a widely used plastic, but these degradation process. Plastics are typically stable and not
has been little research on its direct degradation. Most easily biodegradable. Recent investigations have chal-
research has employed pre-treatment techniques, includ- lenged the conventional belief that plastics are not biode-
ing γ-irradiation, UV irradiation, and heat treatment, fol- gradable by nature. The main microorganisms capable of
lowed by deterioration. For example, Bacillus is capable degrading plastics and their degradation capabilities are
of degrading UV-treated PP (Devi et al. 2023). Only three summarized in Table 2. Nevertheless, the direct biodeg-
strains of Pseudomonas spp., Vibrio sp., and Alcaligenes radation process exhibits limited efficiency, as the major-
spp. have been shown to degrade PP without any prior ity of plastics undergo direct biodegradation, resulting in
treatment (Kelly et al. 2021). However, the degradation a mass loss of 10% or less. Consequently, further in-depth
efficiency is minimal, with only 4% degradation observed studies are necessary.
after 40 days treatment.
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 7 of 16
Degradation of the polyester family the degradation of PBS is typically addressed by utilizing
Compared with plastics, polyesters can undergo bio- microorganisms capable of excreting lipase or by directly
degradation through hydrolysis reactions, as they are application of lipase. For instance, viscous lipase F, the
cross-linked through esterification reaction. Polyesters, enzyme was found in Rhizopus niveus, is capable of fully
including polybutylene succinate (PBS), polybutylene degrading PBS in a span of 17 days at a temperature of
succinate butadiene styrene (PBSA), polybutylene succi- 37 °C and a pH level of 7.0. The Asahi lipase from Chro-
nate butadiene terephthalate (PBAT), polycaprolactone mobacterium viscosum can completely degrade PBSA in
(PCL), and polylactic acid (PLA) are environmentally only 4 days (Arunrattanamook et al. 2023). Also, Rhizo-
friendly and biodegradable (Jiang et al. 2024; Satti and pus niveus, Alcaligenes sp., and Rhizopus oryzae have the
Shah 2020). ability to fully degrade PBSA within 6, 11, and 22 days,
PBS is a type of thermoplastic polyester with excep- respectively (Arunrattanamook et al. 2023).
tional processing and mechanical properties. PBS is PCL possesses exceptional characteristics, such as bio-
classified as a highly degradable substance because it is compatibility, favorable biodegradability, and compat-
formed as a polyester (Zhang et al. 2019). PBS, PBSA, ibility with various materials. It is synthesized through
and PBAT exhibit comparable degradation characteris- the ring-opening polymerization of the ε-caprolactone
tics due to their esterification through a common group monomer. Therefore, PCL is extensively utilized in the
called butylene glycol ester. According to reports, lipase manufacturing of drug carriers, plasticizers, and biode-
and cutinase enzymes released by bacteria can catalyze gradable polymers. Various microbial strains can effi-
the hydrolysis of ester bonds, leading to the degrada- ciently degrade PCL with high degradation efficiency. For
tion of polyesters (Lin et al. 2019; Shi et al. 2019). Nev- example, treatment of PCL with Chaetomium globosum
ertheless, degradation by cutinase results in degradation ATCC 16,021 for 28 days resulted in the formation of vis-
byproducts that have a diameter three times greater than ible micropores and cracks, which caused a significant
that of lipase. Furthermore, the PBS degraded by cutinase mass reduction of 75% (Vivi et al. 2019). Ahmed Nawaz
does not exhibit any alterations in crystallinity, whereas obtained Brevundimonas sp. MRL-AN1 from the soil,
the lipase-degraded PBS demonstrates a gradual reduc- which efficiently decomposed over 80% of the PCL within
tion in crystallinity over time (Shi et al. 2019). Therefore, 10 days. The enzyme excreted by this strain exhibited
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 8 of 16
stability throughout a broad spectrum of temperatures revealed and recognized conservative sites (Tyr139,
(20–45 °C) and pH levels (5–9), as well as in the pres- Tyr213, Arg259, and Thr46) of the RPA1511 hydrolase.
ence of diverse metal ions, surfactants, and organic sol- Modifying amino acids at specific sites such as His114,
vents. These characteristics render it appropriate for the Trp219, and Ala273 was found to enhance the reac-
decomposition of polyesters in intricate circumstances tion’s effectiveness, whereas modification of amino acids
(Nawaz et al. 2015). Furthermore, Brevundimonas sp. at Arg244 hindered the reaction (Wang et al. 2019b). In
MRL-AN1 has shown significant efficacy against various short, various modifications to the enzyme all had the
additional polyesters. Microorganisms capable of degrad- capacity to impact the degraded effectiveness of PLA by
ing polyesters can be found in various circumstances, the strain.
including extreme conditions such as Antarctica. These In conclusion, all polyesters exhibit exceptional biode-
microorganisms are not limited to typical soils. Aneta gradability and can be naturally degraded in environmen-
isolated 161 bacterial and 38 fungal strains from soil tal conditions. Nevertheless, the process of degradation
samples collected in Antarctica, all of which possess the can be expedited, and the efficiency of waste treatment
ability to biodegrade polyesters. Over 92% of the bacteria can be enhanced by conducting a screening to identify
and 77% of the fungus exhibited a high degrading activity, the dominant strains. Proteins extracted from bacteria
resulting in the etching and notching of PBSA, PBS, and with the ability to degrade polyesters are summarized in
PCL. Sclerotinia sp. B11IV and Fusarium sp. B3’M exhib- Table 3.
ited significant biodegradation activity, degrading 49.7%
of PBSA and 33.7% of PCL, and 46.0% of PBSA and 49.7% The degradation process of polymer with additives
of PCL, respectively. These two strains had the lowest Additive polymers are a distinct category of polymers
optimal temperature requirement of 20 °C compared to whose properties are modified and optimized by the
all other strains that degrade polyester (Urbanek et al. addition of various additives during the manufacturing
2021). All of these strains exhibited considerable PCL process, such as emulsifiers, dispersants, and flame retar-
degradation efficiency, but none of them reached the dants. The term used to refer to these substances is “com-
highest level. The most effective strain for degrading PCL patibilizers”. For example, flame retardants are added to
is Penicillium oxalicum strain DSYD05-1, which was cre- certain polymers to reduce the flammability of the poly-
ated by Fan using UV irradiation of Penicillium oxalicum. mer (Pomata et al. 2024; Vahabi et al. 2021). This method
This strain resulted in up to 80% weight loss after treating can also enhance the biodegradability of polymers (Bher
PCL for 6 days (Khatua et al. 2024). et al. 2023), the processes for degrading polymers with
PLA is a completely biodegradable polyester produced additives are shown in Fig. 5. Research has demonstrated
by the polymerization of lactic acid from the conversion that employing starch as a compatibilizer enhances the
of a renewable resource such as starch. The monomeric rate of biodegradation and expedites the degradation of
lactic acid in PLA can be directly metabolized by micro- PS and PP (Zhang et al. 2022). The composite TPS/MA/
organisms. Research indicates that Rhodopseudomonas PLA is formed by combining polylactic acid with maleic
palustris exhibits remarkable degradation efficiency of anhydride (MA) and thermoplastic starch (TPS) as com-
PLA, capable of degrading approximately 40% of it within patibilizers. Ricardo observed that pure polylactic acid
36 h (Hajighasemi et al. 2016). The efficient degradation had a degradation rate of 40.4%, whereas TPS/MA/PLA
of PLA by the strain was principally facilitated by the showed a higher degradation rate of 82% after undergo-
release of its hydrolase RPA1511. Further investigations ing a 31-day degradation treatment (Camacho-Muñoz
Table 3 Summary of proteins derived from bacteria with the ability to degrade polyesters
Protein name Source Polyesters degradation capability Ref.
Lipase F R. niveus Degrade PBS within 17 days at 37 °C and pH 7.0. Arunrattanamook et al. 2023
Lipase Asahi C. viscosum Completely degrade PBSA within 4 days. Arunrattanamook et al. 2023
Lipase F-AP15 R. oryzae Degraded PBSA after cultivation for 22 days. Arunrattanamook et al. 2023
Lipase-QL Alcaligenes sp. Degraded PBSA after cultivation for 11 days. Arunrattanamook et al. 2023
PCL depolymerase Brevundimonas sp. strain More than 80% of PCL was degraded within 10 days. Nawaz et al. 2015
MRL-AN1
PCL-degrading enzyme P. oxalicum DSYD05 The weight loss can reach 80% after 6 days of cultivation. Khatua et al. 2024
PCL-degrading enzyme C. globosum ATCC 16,021 The mass loss was as high as 75%, after 28 days. Vivi et al. 2019
PCL-degrading enzyme Sclerotinia sp. B11IV The biodegradation activity was 49.7% for PBSA and 33.7% for Urbanek et al. 2021
PCL at 20 °C within 30 days.
PCL-degrading enzyme Fusarium sp. B3’M The biodegradation activity was 49.7% for PBSA 4.0% and PCL Wang et al. 2019b
49.7% at 20 °C within 30 days.
Hydrolase RPA1511 R. palustris Degraded almost 40% of PLA within 36 h of incubation. Wang et al. 2019b
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 9 of 16
notably superior to those achieved by individual strains a wider range of potential applications due to the pres-
(Park and Kim 2019). Additionally, a microbial commu- ence of several influencing elements in the treatment
nity consisting of multiple actinomycetes is more effec- process. Furthermore, microbial communities exhibit
tive than a single actinomycete at rubber degradation superior stability compared to individual strains. Hence,
(Nguyen et al. 2020). Recognizing the higher degradation the exploration of other microbial communities with
capacities possessed by natural microbial communities, the capability to degrade polymers is a highly intriguing
some researchers have attempted to construct artificial research endeavor.
microbial communities to facilitate polymer degrada-
tion. Pseudomonas otitidis strain SPT1, Bacillus aerius Polymer degradation using enzyme engineering
strain SPT2, Acanthopleuribacter pedis strain SPT3, and PET is a polymer formed by ester bonds between tere-
Bacillus cereus strain SPK1 were randomly combined phthalic acid and ethylene glycol. It is widely used, espe-
into microbial communities for comparison with single cially in packaging applications such as beverage bottles
strains on their ability to degrade PVC. It was determined (Sova et al. 2023; Srivastava et al. 2024). PET has received
that the combination of microbial communities was more extensive attention in biodegradation studies. Contem-
efficient in degrading PVC than single strains (Dhanraj et porary molecular biology techniques, including pro-
al. 2022). tein engineering and genetic engineering, are primarily
Following the discovery that the deterioration caused applied to enhance the biodegradation of PET.
by microbial communities was more significant than PET exhibits exceptional durability and retains its
that caused by individual strains, it became imperative to structural integrity even after 15 years of natural degra-
investigate the specific contribution of each strain within dation in outdoor environments, despite belonging to
the microbial communities. Vagras isolated three micro- the polyester group (Ioakeimidis et al. 2016). Biodegra-
bial communities from El Bordo Poniente landfill, which dation treatments can expedite this process. Microbial
were named BP1h, BP3h, and BP7h, and deposited at the production of cutinases is the primary factor responsible
World Data Centre for Microbiology CFQ100 under the for PET biodegradation. These enzymes are quite similar
accession numbers CFQ-B-261, CFQ-B-269, and CFQ-B- to the other and usually reach their peak activity at the
264, respectively. The researchers examined the capacity PET glass transition temperature (Tg) of PET, which is
of these communities to degrade polyester and polyure- approximately 70 °C (Amanna and Rakshit 2023; Rich-
thane by measuring the various enzyme activities of the ter et al. 2023). Actinomyces thermophilus excretes the
microbiota and assessing the growth potential of par- most typical cutinase (Dąbrowska et al. 2021).However,
ticular microbial community strains in these polymers. enzymes from the wild-type strain, are highly susceptible
The majority of the microbial strains in the community to deactivation near the PET glass transition point and
were unable to thrive on these polymers independently. exhibit low thermal stability. Researchers have employed
Nevertheless, the strains exhibited the ability to thrive various methodologies to enhance the thermal resis-
on polymers when they were amalgamated to establish tance of enzymes. For instance, research has revealed
a microbial community. The researchers hypothesized that the addition of divalent metal ions, such as Ca2+ or
that these strains established a network of interactions Mg2+ ions, enhances the thermal stability of the hydro-
where each member had a distinct role, enabling them lase enzyme, allowing it to degrade PET at a tempera-
to sustain the growth of the microbial community (Var- ture of 65 °C (Qi et al. 2024; Serrano-Aguirre and Prieto
gas-Suarez et al. 2019). When the strains within a col- 2024). Instead, by substituting metal binding sites with
ony establish a stable network, there can be additional salt bridges or disulfide bonds, the hydrolase may operate
advantageous outcomes apart from the enhanced effi- at a temperature of 70 °C and degrade PET without the
ciency of the degradation process. Swiontek discovered need for divalent metal ions (Zhong-Johnson et al. 2024).
that a microbial consortium composed of Aeromonas Moreover the enzymes under investigation, namely
and Rhodococcus showed a significant ability to degrade Fusarium solani pisi cutinase (FsC), leaf-branch com-
polylactic acid (PLA), polyhydroxybutyrate (PHB), and post cutinase (LCC), and Thermobifida fusca hydrolases
polystyrene (PCL). The increased concentration of bac- 1 and 2 (BTA1 and BTA2), have been evaluated for their
teriostatic did not impact the degradation ability of the individual abilities to degrade substances. At a tempera-
microbial community as a whole. However, the presence ture of 65 °C, LCC demonstrated the highest efficiency
of bacteriostatic hindered the degradation ability of an in degrading PET, with an initial depolymerization rate
individual strain. These findings suggest that microbial of 93.2 mgTAeq.h− 1mgenzyme−1 and a 50% degradation
communities exhibit greater resilience to stress com- rate achieved within 12 h. In addition, the replacement of
pared to individual strains (Swiontek Brzezinska et al. divalent metal binding sites with disulfide bonds resulted
2020). This implies that microbial communities not only in an improvement in both catalytic activity and thermal
have enhanced degrading capabilities, but also provide stability of the LCC enzyme (Tournier et al. 2020a).The
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 11 of 16
enhancement of enzyme activity through the evolution (Fecker et al. 2018). Tournier utilized molecular docking
of enzymes is now possible due to recent advancements and enzyme contact-surface analysis strategies to iden-
in protein and enzyme engineering (Chen et al. 2017; tify mutagenesis sites to improve the catalytic activity
Wang et al. 2017). Similarly, modifications were imple- of LCC. A total of 209 mutants were obtained by satura-
mented to cutinase, a pivotal enzyme in the degradation tion mutagenesis of the screened sites. The most efficient
of PET, with the aim of enhancing the efficiency of PET LCC enzyme mutant converted at least 90% of PET to
degrade. Meng and Yang developed the mutation design monomer in 10 h, with a degradation efficiency of 16.7
tool Premuse, which they utilized to identify and modify ghydrolyzed PET L− 1 h− 1 (Tournier et al. 2020b). Despite the
two stable mutants (W159H and F229Y) from a pool of achievements, its large-scale applications are still ham-
1486 similar sequences with enhanced enzyme activ- pered by the remaining 10% of nonbiodegradable PET.
ity. Compared to the wild type, the enzyme’s denaturing Cui et al. designed a mutant of PETase (TurboPETase),
temperature and catalytic efficiency (kcat/Km) increased with balanced thermostability and hydrolytic capacity, by
by 10.4 °C and 2 folds, respectively, while its degrad- incorporating a protein language model and force-field-
ing activity surged by nearly 40 folds at 40 °C (Meng et based algorithms. This mutant can nearly completely
al. 2021). Guo discovered that Ideonella sakaiensis 201- depolymerize 200 g of PET in 8 h, with a production rate
F6 has a distinctive cutinase known as IsPETase, which of 61.3 ghydrolyzed PET L− 1 h− 1 (Cui et al. 2024). Construct-
shares a high degree of similarity with LCC in terms of ing of a large mutant library by computational analysis
both its sequence and structure, and exhibits a high effi- can greatly accelerate the discovery of enzymes with high
cacy in decomposing PET. The hydrolytic activity of the heat resistance and high degradation activity. Limited
enzyme IsPETase was enhanced by substituting the resi- by the low flux of conventional evaluation methods, the
dues S214 with Ile, and I218 with Ser (Chen et al. 2021). amounts of mutant libraries are usually within 104. Cri-
Bifidobacterium thermophilus strain TfCut2 produces an bari proposed a high-throughput yeast surface display
IsPETase enzyme that exhibits excellent thermostability. platform that can rapidly evaluate mutants with more
Two enzyme mutants, G62A and G62A/I213s, were cre- than 107 enzymes. On this platform, each yeast cell can
ated by replacing four conserved amino acids (G62, T63, display different mutants. The enzyme activity is detected
I178, and I213) of TfCut2 with the corresponding amino by the change of fluorescence during the cleavage of the
acids of LCC. These mutants exhibited enhanced PET synthetic probe. Then, the highly active mutants are iso-
hydrolysis activity and improved thermal stability com- lated, which increases the screening flux by 1000 times
pared to the original LCC enzyme. The hydrolysis activity (Cribari et al. 2023). Previously strategies were based on
of G62A increased 4-fold compared to the wild type. In static protein conformation calculations, which could
addition, it showed a 5.5-fold decrease in the inhibitor’s hardly reflect the dynamic process of catalysis. For this
binding ability, hence reducing the inhibitor’s impact on reason, Zheng et al. devised a new computational strat-
the degradation process (Wei et al. 2016). egy (affinity analysis based on dynamic docking, ADD)
Determining the enzyme’s crystal structure is crucial to analyze the ligand affinity energy by molecular dock-
for understanding the mechanism of enzyme-catalyzed ing with the dynamic protein conformations. Compared
reactions and providing scientific guidance for enzyme to static protein conformations, dynamic conformations
modification. In order to analyze the mechanism of PET are more realistic and accurate, which facilitates the dis-
degradation catalyzed by IsPET, Austin identified the covery of more promising modification sites. The mutant
crystal structure of the IsPET produced by Ideonella LCC-A2, obtained by the ADD strategy, depolymerized
sakaiensi 201-F6 at a resolution of 0.92 Å, which is the over 90% of PET into terephthalic acid and glycol within
highest-resolution X-ray crystal structure of the apo- 3.3 h at 78 °C. This is currently the fastest PET depoly-
enzyme available in the database (Austin et al. 2018). merization rate on record (Zheng et al. 2024).
Meseguer used the crystal structure resolved by Aus- Increased enzymatic activity is, essential for PET deg-
tin to analyze the cause of the enhanced activity of the radation, as well as for the efficient release of the enzyme
PETase mutant (FAST-PETase). By employing classical into the extracellular environment. Limited studies have
and hybrid (QM/MM) molecular dynamics (MD) simu- been conducted on the synthesis of enzymes that degrade
lations, they determined that the mutation of N233K PET. Five distinct Bacillus signal peptides were evaluated
causes a series of changes that ultimately reduce the cata- for their effect on the secretion of PET hydrolase. The
lytic barrier and accelerate the PET degradation reaction results demonstrated that SP amy generated the high-
(García-Meseguer et al. 2023). Simultaneously with Aus- est volume of secretion, almost quadruple that of the
tin’s IsPET analysis, Fecker analyzed the crystal struc- native signal peptide SP. In addition, they observed that
ture of PETase at a resolution of 2.02 Å. This structure the upregulation in PET hydrolase expression was trig-
was used in molecular dynamics simulations, giving a gered by the low-strength P43 promoter. According to
firmer theoretical foundation for the evolution of IsPET the authors’ hypothesis, the weak promoter may allow
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 12 of 16
sufficient time for translation and folding processes. P43 enhancing the resilience of bacteria and enzymes in vari-
and SP Amy exhibited enhanced degradation activity on ous environmental conditions.
PET films (Wang et al. 2020). This suggests that the pro- Recent research has questioned the conventional
cess of degrading substances is improved by the effective method of separating distinct strains for the breakdown
release of enzymes. of polymers. The benefits of microbial communities have
PET is the most extensively studied polymer in terms been emphasized. Previous reviews have not specifically
of biodegradation. Currently, researchers are engaged in highlighted the significance of microbial communities in
the development and modification of potent enzymatic waste polymer degrading endeavors. This is the inaugural
agents to significantly enhance the efficiency of PET instance where microbial communities have been given
degradation, rather than identifying efficient microbial equal priority alongside the value of individual strains
strains. Table 4 summarizes the PET degradation capa- and synthetic enzymes. Microbial communities have
bilities of various PET-degrading enzymes after modifi- demonstrated superior efficacy in breaking down poly-
cation by protein engineering. Although contemporary mers compared to individual strains. They also exhibit
molecular biology techniques are mainly applied to PET enhanced stability and resistance, particularly when faced
degradation at present, they also open up possibilities for with the intricate combinations of polymers commonly
studying other forms of polymer degradation in future found in recyclable garbage. Engineered enzymes have
advancements. the ability to enhance the speed at which some polymers
break down, but they are not yet capable of effectively
Conclusions and future perspectives treating polymers that are combined together. Waste
Due to their exceptional characteristics, polymers are sorting methods are typically rudimentary, generally
widely employed. However, disposing of waste polymers grouping numerous polymers together in one category.
has always been a challenge because of the lack of appro- Dealing with intricate polymers using only one specific
priate treatment method. Existing methods have their strain or modified enzyme can provide difficulties. Prac-
limitations, more precisely, the process of biodegrading tical applications derive advantages from the flexibility
polymers is relatively sluggish and biodegradation has a of multi-strain communities in the treatment of various
lower resistance to unfavorable settings. Consequently, polymers. Techniques to augment the capacity of a soli-
scientists sought alternate methodologies, such as modi- tary strain to break down polymers can be extended to all
fying the enzyme to enhance its ability to degrade sub- strains within the community, while introducing supple-
stances. However, the majority of bacteria and enzymes mentary strains or engineered enzymes can expand the
lose their activity at elevated temperatures or when spectrum of polymers that can be decomposed, enhance
exposed to detrimental chemicals such as acids, alka- the efficiency of degradation, and greatly enhance the
lis, and antibiotics. Hence, additional comprehensive overall capacity of the community to degrade polymers.
research is necessary before the development of an engi- In the future, multi-strain degradation of polymers will
neered strain for waste polymer treatment. This involves be the most competitive biodegradation method. Ulti-
addressing challenges related to the slow natural degra- mately, biodegradation, whether facilitated by single bac-
dation process, exploring alternative methodologies, and teria, multi-strain communities, or engineered enzymes,
Table 4 Summary of proteins modificated by protein engineering with the ability to degrade PET.
Protein name Source Degradation capability Ref.
Hydrolases 1 and 2 T. fusca Reaching an initial PET-specific depolymerization rate of 3.2 mgTAeq.h− 1mgenzyme−1 at Tournier et
65 °C. al. 2020a
FsC F. solani Reaching an initial PET-specific depolymerization rate of 0.01 mgTAeq.h− 1mgenzyme−1 at Tournier et
65 °C. al. 2020a
LCC Leaf-branch 50% PET can be degraded within 12 h, reaching an initial PET-specific depolymerization Tournier et
compost rate of 93.2 mgTAeq.h− 1mgenzyme−1 at 65 °C. al. 2020a
TfCut2 G62A/I213s B. thermophilus Compared with the wild-type mutant, the degradation activity was increased by 2.7 folds. Wei et al.
strain KW3 2016
W159H and F229Y Screen out by Compared with the wild type, the degradation activity at 40 °C was nearly 40 folds higher. Meng et
Premuse al. 2021
ICCG LCC enzyme Degradating at least 90% of PET to monomer in 10 h, with a degradation efficiency of 16.7 Tournier et
mutant ghydrolyzed PET L− 1 h− 1. al. 2020b
TurboPETase a mutant of Depolymerizing 200 g of PET in 8 h, with a production rate of 61.3 ghydrolyzed PET L− 1 h− 1. Cui et al.
PETase 2024
LCC-A2 obtained by the Depolymerizing more than 90% of PET into terephthalic acid and glycol within 3.3 h at Zheng et
ADD strategy 78 °C al. 2024
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 13 of 16
Competing interests
now represents the most effective method for managing The authors declare that they have no competing interests.
waste polymers.
While using microbial communities mitigates some Author details
1
State Key Laboratory of Chemical Resource Engineering, Beijing
limitations associated with single strains, there are University of Chemical Technology, Beijing 100029, China
emerging issues that need further examination. It is 2
State Key Laboratory of Mycology, Institute of Microbiology, Chinese
essential to investigate the potential effects of introduc- Academy of Sciences, Beijing 100101, China
Chen Z, Shen X, Wang J, Wang J, Yuan Q, Yan Y (2017) Rational engineering of Howard SA, Carr CM, Sbahtu HI, Onwukwe U, López MJ, Dobson ADW, McCarthy
p-hydroxybenzoate hydroxylase to enable efficient gallic acid synthesis via a RR (2023) Enrichment of native plastic-associated biofilm communities to
novel artificial biosynthetic pathway. Biotechnol Bioeng 114(11):2571–2580. enhance polyester degrading activity. Environ Microbiol 25(12):2698–2718.
https://doi.org/10.1002/bit.26364 https://doi.org/10.1111/1462-2920.16466
Chen CC, Dai L, Ma L, Guo RT (2020) Enzymatic degradation of plant biomass and Ibrahim S, Gupta RK, War AR, Hussain B, Kumar A, Sofi T, Noureldeen A, Darwish
synthetic polymers. Nat Rev Chem 4(3):114–126. https://doi.org/10.1038/ H (2021) Degradation of chlorpyriphos and polyethylene by endosymbiotic
s41570-020-0163-6 bacteria from citrus mealybug. Saudi J Biol Sci 28(6):3214–3224. https://doi.
Chen C-C, Han X, Li X, Jiang P, Niu D, Ma L, Liu W, Li S, Qu Y, Hu H, Min J, Yang org/10.1016/j.sjbs.2021.03.058
Y, Zhang L, Zeng W, Huang J-W, Dai L, Guo R-T (2021) General features to Imai S, Ichikawa K, Muramatsu Y, Kasai D, Masai E, Fukuda M (2011) Isolation and
enhance enzymatic activity of poly(ethylene terephthalate) hydrolysis. Nat characterization of Streptomyces, Actinoplanes, and Methylibium strains that
Catal 4(5):425–430. https://doi.org/10.1038/s41929-021-00616-y are involved in degradation of natural rubber and synthetic poly(cis-1,4-iso-
Chow J, Perez-Garcia P, Dierkes R, Streit WR (2023) Microbial enzymes will offer prene). Enzyme Microb Technol 49(6–7):526–531. https://doi.org/10.1016/j.
limited solutions to the global plastic pollution crisis. Microb Biotechnol enzmictec.2011.05.014
16(2):195–217. https://doi.org/10.1111/1751-7915.14135 Ioakeimidis C, Fotopoulou KN, Karapanagioti HK, Geraga M, Zeri C, Papathanas-
Cribari MA, Unger MJ, Unarta IC, Ogorek AN, Huang X, Martell JD (2023) Ultrahigh- siou E, Galgani F, Papatheodorou G (2016) The degradation potential of
throughput directed evolution of polymer-degrading enzymes using yeast pet bottles in the marine environment: an atr-ftir based approach. Sci Rep
display. J Am Chem Soc 145(50):27380–27389. https://doi.org/10.1021/ 6:23501. https://doi.org/10.1038/srep23501
jacs.3c08291 Jiang J, Luo Y, Fei P, Zhu Z, Peng J, Lu J, Zhu D, Wu H (2024) Effect of adaptive labo-
Cui C, Jiang M, Zhang C, Zhang N, Jin FJ, Li T, Lee HG, Jin L (2023) Assembly strate- ratory evolution of engineered Escherichia coli in acetate on the biosynthesis
gies for rubber-degrading microbial consortia based on omics tools. Front of succinic acid from glucose in two-stage cultivation. Bioresour Bioprocess
Bioeng Biotechnol 11:1326395. https://doi.org/10.3389/fbioe.2023.1326395 11(1):34. https://doi.org/10.1186/s40643-024-00749-5
Cui Y, Chen Y, Sun J, Zhu T, Pang H, Li C, Geng WC, Wu B (2024) Computational Kelly JJ, London MG, McCormick AR, Rojas M, Scott JW, Hoellein TJ (2021) Waste-
redesign of a hydrolase for nearly complete pet depolymerization at water treatment alters microbial colonization of microplastics. PLoS ONE
industrially relevant high-solids loading. Nat Commun 15(1):1417. https://doi. 16(1):e0244443. https://doi.org/10.1371/journal.pone.0244443
org/10.1038/s41467-024-45662-9 Khandare SD, Chaudhary DR, Jha B (2021) Marine bacterial biodegradation of low-
Dąbrowska GB, Garstecka Z, Olewnik-Kruszkowska E, Szczepańska G, Ostrowski density polyethylene (ldpe) plastic. Biodegradation 32(2):127–143. https://
M, Mierek-Adamska A (2021) Comparative study of structural changes of doi.org/10.1007/s10532-021-09927-0
polylactide and poly(ethylene terephthalate) in the presence of Trichoderma Khatua S, Simal-Gandara J, Acharya K (2024) Myco-remediation of plastic pollution:
Viride. Int J Mol Sci 22:7. https://doi.org/10.3390/ijms22073491 current knowledge and future prospects. Biodegradation 35(3):249–279.
DelRe C, Jiang Y, Kang P, Kwon J, Hall A, Jayapurna I, Ruan Z, Ma L, Zolkin K, Li T, https://doi.org/10.1007/s10532-023-10053-2
Scown CD, Ritchie RO, Russell TP, Xu T (2021) Near-complete depolymeriza- Kotova IB, Taktarova YV, Tsavkelova EA, Egorova MA, Bubnov IA, Malakhova DV,
tion of polyesters with nano-dispersed enzymes. Nature 592(7855):558–563. Shirinkina LI, Sokolova TG, Bonch-Osmolovskaya EA (2021) Microbial degra-
https://doi.org/10.1038/s41586-021-03408-3 dation of plastics and approaches to make it more efficient. Microbiology
Devi D, Gupta KK, Chandra H, Sharma KK, Sagar K, Mori E, de Farias PAM, Coutinho 90(6):671–701. https://doi.org/10.1134/S0026261721060084
HDM, Mishra AP (2023) Biodegradation of low-density polyethylene Lin H, Mu X, Huang J, Jiang H, Niu J, Shu Z (2019) Comparative analysis of polyester
(ldpe) through application of indigenous strain Alcaligenes faecalis isj128. hydrolysis activity among three lipolytic enzymes. J Chem Technol Biotechnol
Environ Geochem Health 45(12):9391–9409. https://doi.org/10.1007/ 94(8):2522–2528. https://doi.org/10.1002/jctb.6042
s10653-023-01590-z Lusher AL, Hernandez-Milian G, Berrow S, Rogan E, O’Connor I (2018) Incidence of
Dhanraj ND, Hatha AAM, Jisha MS (2022) Biodegradation of petroleum based and marine debris in cetaceans stranded and bycaught in Ireland: recent findings
bio-based plastics: approaches to increase the rate of biodegradation. Arch and a review of historical knowledge. Environ Pollut 232:467–476. https://doi.
Microbiol 204(5):258. https://doi.org/10.1007/s00203-022-02883-0 org/10.1016/j.envpol.2017.09.070
Duval C (2014) Plastic waste and the environment. Environ Impact Polym. https:// Meng X, Yang L, Liu H, Li Q, Xu G, Zhang Y, Guan F, Zhang Y, Zhang W, Wu N, Tian
doi.org/10.1002/9781118827116.ch2 J (2021) Protein engineering of stable ispetase for pet plastic degradation
Europe P (2023) Plastics – the fast facts 2023. Plastics Europe, https://plasticseu- by premuse. Int J Biol Macromol 180:667–676. https://doi.org/10.1016/j.
rope.org/knowledge-hub/plastics-the-fast-facts-2023/ ijbiomac.2021.03.058
Fecker T, Galaz-Davison P, Engelberger F, Narui Y, Sotomayor M, Parra LP, Ramírez- Meyer Cifuentes IE, Degenhardt J, Neumann-Schaal M, Jehmlich N, Ngugi DK,
Sarmiento CA (2018) Active site flexibility as a hallmark for efficient pet Öztürk B (2023) Comparative biodegradation analysis of three compostable
degradation by i.sakaiensis petase. Biophys J 114(6):1302–1312. https://doi. polyesters by a marine microbial community. Appl Environ Microbiol
org/10.1016/j.bpj.2018.02.005 89(12):e0106023. https://doi.org/10.1128/aem.01060-23
Finzi-Quintão CM, Novack KM, Bernardes-Silva AC, Silva TD, Moreira LES, Braga LEM Montazer Z, Habibi Najafi MB, Levin DB (2019) Microbial degradation of low-
(2019) Biodegradation of moringa oleifera’s polymer blends. Environ Technol density polyethylene and synthesis of polyhydroxyalkanoate polymers. Can J
40(4):508–517. https://doi.org/10.1080/09593330.2017.1397763 Microbiol 65(3):224–234. https://doi.org/10.1139/cjm-2018-0335
García-Meseguer R, Ortí E, Tuñón I, Ruiz-Pernía JJ, Aragó J (2023) Insights into the Mostafa NA, Farag AA, Abo-dief HM, Tayeb AM (2018) Production of biodegrad-
enhancement of the poly(ethylene terephthalate) degradation by fast-petase able plastic from agricultural wastes. Arab J Chem 11(4):546–553. https://doi.
from computational modeling. J Am Chem Soc 145(35):19243–19255. org/10.1016/j.arabjc.2015.04.008
https://doi.org/10.1021/jacs.3c04427 Nawaz A, Hasan F, Shah AA (2015) Degradation of poly(ε-caprolactone) (pcl) by a
Giacomucci L, Raddadi N, Soccio M, Lotti N, Fava F (2019) Polyvinyl chloride newly isolated Brevundimonas sp. Strain mrl-an1 from soil. FEMS Microbiol
biodegradation by Pseudomonas citronellolis and Bacillus flexus. N Biotechnol Lett 362(1):1–7. https://doi.org/10.1093/femsle/fnu004
52:35–41. https://doi.org/10.1016/j.nbt.2019.04.005 Nawong C, Umsakul K, Sermwittayawong N (2018) Rubber gloves biodegradation
Gómez-Méndez LD, Jiménez-Borrego LC, Pérez-Flórez A, Poutou-Piñales RA, by a consortium, mixed culture and pure culture isolated from soil samples.
Pedroza-Rodríguez AM, Salcedo-Reyes JC, Vargas A, Bogoya JM (2021) Ldpe Braz J Microbiol 49(3):481–488. https://doi.org/10.1016/j.bjm.2017.07.006
transformation by exposure to sequential low-pressure plasma and tio(2)/uv Nguyen LH, Nguyen HD, Tran PT, Nghiem TT, Nguyen TT, Dao VL, Phan TN, To
photocatalysis. Molecules 26(9). https://doi.org/10.3390/molecules26092513 AK, Hatamoto M, Yamaguchi T, Kasai D, Fukuda M (2020) Biodegradation of
Hajighasemi M, Nocek BP, Tchigvintsev A, Brown G, Flick R, Xu X, Cui H, Hai T, natural rubber and deproteinized natural rubber by enrichment bacte-
Joachimiak A, Golyshin PN, Savchenko A, Edwards EA, Yakunin AF (2016) rial consortia. Biodegradation 31(4–6):303–317. https://doi.org/10.1007/
Biochemical and structural insights into enzymatic depolymerization of poly- s10532-020-09911-0
lactic acid and other polyesters by microbial carboxylesterases. Biomacromol- Olewnik-Kruszkowska E, Burkowska-But A, Tarach I, Walczak M, Jakubowska E
ecules 17(6):2027–2039. https://doi.org/10.1021/acs.biomac.6b00223 (2020) Biodegradation of polylactide-based composites with an addition of
Harshvardhan K, Jha B (2013) Biodegradation of low-density polyethylene by a compatibilizing agent in different environments. Int Biodeterior Biodegrad
marine bacteria from pelagic waters, arabian sea, India. Mar Pollut Bull 147. https://doi.org/10.1016/j.ibiod.2019.104840
77(1–2):100–106. https://doi.org/10.1016/j.marpolbul.2013.10.025 Park SY, Kim CG (2019) Biodegradation of micro-polyethylene particles by
bacterial colonization of a mixed microbial consortium isolated from
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 15 of 16
a landfill site. Chemosphere 222:527–533. https://doi.org/10.1016/j. Srivastava P, Saji J, Manickam N (2024) Biodegradation of polyethylene terephthal-
chemosphere.2019.01.159 ate (pet) by Brucella Intermedia iitr130 and its proposed metabolic pathway.
Pathak VM, Navneet (2017) Review on the current status of polymer degradation: https://doi.org/10.1007/s10532-024-10070-9. Biodegradation
a microbial approach. Bioresour Bioprocess 4(1):15. https://doi.org/10.1186/ Sun J, Prabhu A, Aroney STN, Rinke C (2022a) Insights into plastic biodegrada-
s40643-017-0145-9 tion: community composition and functional capabilities of the superworm
Peixoto J, Silva LP, Kruger RH (2017) Brazilian cerrado soil reveals an untapped (Zophobas Morio) microbiome in styrofoam feeding trials. Microb Genom
microbial potential for unpretreated polyethylene biodegradation. J Hazard 8(6). https://doi.org/10.1099/mgen.0.000842
Mater 324 (Pt B 634–644. https://doi.org/10.1016/j.jhazmat.2016.11.037 Sun Y, Hu J, Yusuf A, Wang Y, Jin H, Zhang X, Liu Y, Wang Y, Yang G, He J (2022b)
Pomata D, La Nasa J, Biale G, Barlucchi L, Ceccarini A, Di Filippo P, Riccardi C, A critical review on microbial degradation of petroleum-based plastics:
Buiarelli F, Modugno F, Simonetti G (2024) Plastic breath: quantification of quantitatively effects of chemical addition in cultivation media on biodeg-
microplastics and polymer additives in airborne particles. Sci Total Environ radation efficiency. Biodegradation 33(1):1–16. https://doi.org/10.1007/
932:173031. https://doi.org/10.1016/j.scitotenv.2024.173031 s10532-021-09969-4
Prakash T, Yadav SR, Bürger M, Jendrossek D (2024) Cleavage of natural rubber by Suzuki N, Suda D, Ngan NTT, Gibu N, Huong NL, Anh TK, Kasai D (2022) Charac-
rubber oxygenases in gram-negative bacteria. Appl Microbiol Biotechnol terization of latex-clearing protein and aldehyde dehydrogenases involved
108(1):191. https://doi.org/10.1007/s00253-023-12940-3 in the utilization of poly(cis-1,4-isoprene) by Nocardia farcinica Nbrc 15532.
Prasopdee T, Smitthipong W (2020) Effect of fillers on the recovery of rubber foam: Microorganisms 10:12. https://doi.org/10.3390/microorganisms10122324
from theory to applications. Polym (Basel) 12(11). https://doi.org/10.3390/ Swiontek Brzezinska M, Walczak M, Kalwasinska A, Richert A, Swiatczak J, Deja-
polym12112745 Sikora E, Burkowska-But A (2020) Biofilm formation during biodegradation
Qi X, Wu Y, Zhang ST, Yin CF, Ji M, Liu Y, Xu Y, Zhou NY (2024) The unique salt of polylactide, poly (3,4 hydroxybutyrate) and poly(epsilon-caprolactone) in
bridge network in glacpetase: a key to its stability. Appl Environ Microbiol activated sludge. Int J Biol Macromol 159:539–546. https://doi.org/10.1016/j.
90(3):e0224223. https://doi.org/10.1128/aem.02242-23 ijbiomac.2020.05.107
Rai P, Mehrotra S, Priya S, Gnansounou E, Sharma SK (2021) Recent advances in the Tournier V, Topham CM, Gilles A, David B, Folgoas C, Moya-Leclair E, Kamionka E,
sustainable design and applications of biodegradable polymers. Bioresour Desrousseaux ML, Texier H, Gavalda S, Cot M, Guemard E, Dalibey M, Nomme
Technol 325:124739. https://doi.org/10.1016/j.biortech.2021.124739 J, Cioci G, Barbe S, Chateau M, Andre I, Duquesne S, Marty A (2020a) An engi-
Rajendran N, Han J (2022) Integrated polylactic acid and biodiesel production neered pet depolymerase to break down and recycle plastic bottles. Nature
from food waste: process synthesis and economics. Bioresour Technol 580(7802):216–219. https://doi.org/10.1038/s41586-020-2149-4
343:126119. https://doi.org/10.1016/j.biortech.2021.126119 Tournier V, Topham CM, Gilles A, David B, Folgoas C, Moya-Leclair E, Kamionka E,
Richter PK, Blázquez-Sánchez P, Zhao Z, Engelberger F, Wiebeler C, Künze G, Frank Desrousseaux ML, Texier H, Gavalda S, Cot M, Guémard E, Dalibey M, Nomme
R, Krinke D, Frezzotti E, Lihanova Y, Falkenstein P, Matysik J, Zimmermann W, J, Cioci G, Barbe S, Chateau M, André I, Duquesne S, Marty A (2020b) An engi-
Sträter N, Sonnendecker C (2023) Structure and function of the metage- neered pet depolymerase to break down and recycle plastic bottles. Nature
nomic plastic-degrading polyester hydrolase phl7 bound to its product. Nat 580(7802):216–219. https://doi.org/10.1038/s41586-020-2149-4
Commun 14(1):1905. https://doi.org/10.1038/s41467-023-37415-x Urbanek AK, Strzelecki MC, Mirończuk AM (2021) The potential of cold-adapted
Sarkar B, Gupta AM, Mandal S (2021) Insights from the comparative genome microorganisms for biodegradation of bioplastics. Waste Manag 119:72–81.
analysis of natural rubber degrading nocardia species. Bioinformation https://doi.org/10.1016/j.wasman.2020.09.031
17(10):880–890. https://doi.org/10.6026/97320630017880 Vahabi H, Laoutid F, Mehrpouya M, Saeb MR, Dubois P (2021) Flame retardant poly-
Satti SM, Shah AA (2020) Polyester-based biodegradable plastics: an approach mer materials: an update and the future for 3d printing developments. Mater
towards sustainable development. Lett Appl Microbiol 70(6):413–430. https:// Sci Engineering: R: Rep 144. https://doi.org/10.1016/j.mser.2020.100604
doi.org/10.1111/lam.13287 Vargas-Suarez M, Fernandez-Cruz V, Loza-Tavera H (2019) Biodegradation of poly-
Schmaltz E, Melvin EC, Diana Z, Gunady EF, Rittschof D, Somarelli JA, Virdin J, acrylic and polyester polyurethane coatings by enriched microbial communi-
Dunphy-Daly MM (2020) Plastic pollution solutions: emerging technolo- ties. Appl Microbiol Biotechnol 103(7):3225–3236. https://doi.org/10.1007/
gies to prevent and collectmarineplastic pollution. Environ Int 144:106067. s00253-019-09660-y
https://doi.org/10.1016/j.envint.2020.106067 Vivi VK, Martins-Franchetti SM, Attili-Angelis D (2019) Biodegradation of pcl and
Schmitt G, Birke J, Jendrossek D (2019) Towards the understanding of the pvc: Chaetomium Globosum (atcc 16021) activity. Folia Microbiol (Praha)
enzymatic cleavage of polyisoprene by the dihaem-dioxygenase roxa. AMB 64(1):1–7. https://doi.org/10.1007/s12223-018-0621-4
Express 9(1):166. https://doi.org/10.1186/s13568-019-0888-0 Wang J, Jain R, Shen X, Sun X, Cheng M, Liao JC, Yuan Q, Yan Y (2017) Rational engi-
Serrano-Aguirre L, Prieto MA (2024) Can bioplastics always offer a truly sustainable neering of diol dehydratase enables 1,4-butanediol biosynthesis from xylose.
alternative to fossil-based plastics? Microb Biotechnol 17(4):e14458. https:// Metab Eng 40:148–156. https://doi.org/10.1016/j.ymben.2017.02.003
doi.org/10.1111/1751-7915.14458 Wang M, Liu P, Gu Z, Cheng H, Li X (2019a) A scientometric review of resource recy-
Sharma V, Siedenburg G, Birke J, Mobeen F, Jendrossek D, Prakash T (2018) cling industry. Int J Environ Res Public Health 16(23). https://doi.org/10.3390/
Metabolic and taxonomic insights into the gram-negative natural rubber ijerph16234654
degrading bacterium Steroidobacter cummioxidans sp. Nov., strain 35y. PLoS Wang X, Chen J, Tang X, Wang J, Zhu L, Zhang W, Wang H, Li Y, Zhang Q (2019b)
ONE 13(5):e0197448. https://doi.org/10.1371/journal.pone.0197448 Biodegradation mechanism of polyesters by hydrolase from Rhodopseudo-
Shi K, Su T, Wang Z (2019) Comparison of poly(butylene succinate) biodegradation monas palustris: an in silico approach. Chemosphere 231:126–133. https://
by Fusarium solani cutinase and Candida Antarctica lipase. Polym Degrad Stab doi.org/10.1016/j.chemosphere.2019.05.112
164:55–60. https://doi.org/10.1016/j.polymdegradstab.2019.04.005 Wang N, Guan F, Lv X, Han D, Zhang Y, Wu N, Xia X, Tian J (2020) Enhancing secre-
Shilpa, Basak N, Meena SS (2022) Microbial biodegradation of plastics: challenges, tion of polyethylene terephthalate hydrolase petase in Bacillus subtilis wb600
opportunities, and a critical perspective. Front Environ Sci Eng 16(12):161. mediated by the sp(amy) signal peptide. Lett Appl Microbiol 71(3):235–241.
https://doi.org/10.1007/s11783-022-1596-6 https://doi.org/10.1111/lam.13312
Skariyachan S, Megha M, Kini MN, Mukund KM, Rizvi A, Vasist K (2015) Selec- Wei R, Oeser T, Schmidt J, Meier R, Barth M, Then J, Zimmermann W (2016)
tion and screening of microbial consortia for efficient and ecofriendly Engineered bacterial polyester hydrolases efficiently degrade polyethyl-
degradation of plastic garbage collected from urban and rural areas of ene terephthalate due to relieved product inhibition. Biotechnol Bioeng
bangalore, India. Environ Monit Assess 187(1):4174. https://doi.org/10.1007/ 113(8):1658–1665. https://doi.org/10.1002/bit.25941
s10661-014-4174-y Wei R, von Haugwitz G, Pfaff L, Mican J, Badenhorst CPS, Liu W, Weber G, Austin HP,
Soong YV, Abid U, Chang AC, Ayafor C, Patel A, Qin J, Xu J, Lawton C, Wong HW, Bednar D, Damborsky J, Bornscheuer UT (2022) Mechanism-based design of
Sobkowicz MJ, Xie D (2023) Enzyme selection, optimization, and production efficient pet hydrolases. ACS Catal 12(6):3382–3396. https://doi.org/10.1021/
toward biodegradation of post-consumer poly(ethylene terephthalate) at acscatal.1c05856
scale. Biotechnol J 18(12):e2300119. https://doi.org/10.1002/biot.202300119 Wiśniewska P, Wang S, Formela K (2022) Waste tire rubber devulcanization tech-
Sova N, Savchenko B, Beloshenko V, Slieptsov A, Vozniak I (2023) Sorption proper- nologies: state-of-the-art, limitations and future perspectives. Waste Manag
ties of pet copolyesters and new approach for foaming with filament extru- 150:174–184. https://doi.org/10.1016/j.wasman.2022.07.002
sion additive manufacturing. Polym (Basel) 15(5). https://doi.org/10.3390/ Xing RZ, Zhao ZQ, Zhao WQ, Chen Z, Chen JF, Zhou SG (2021) Biodegradation of
polym15051138 polystyrene by Geobacillus stearothermophilus. Huan Jing Ke Xue 42(6):3056–
3062. https://doi.org/10.13227/j.hjkx.202009298
Wu et al. Bioresources and Bioprocessing (2024) 11:79 Page 16 of 16
Yan T, Balzer AH, Herbert KM, Epps TH 3rd, Korley LTJ (2023) Circularity in polymers: depolymerization. ACS Catal 14(5):3627–3639. https://doi.org/10.1021/
addressing performance and sustainability challenges using dynamic acscatal.4c00400
covalent chemistries. Chem Sci 14(20):5243–5265. https://doi.org/10.1039/ Zhigang X, Kuangdi X (2022) Plastic deformation. The ecph encyclopedia of
d3sc00551h mining and metallurgy. Springer Nature Singapore, Singapore https://link.
Zhang S, He Y, Yin Y, Jiang G (2019) Fabrication of innovative thermoplastic springer.com/referencework/10.1007/978-981-99-2086-0
starch bio-elastomer to achieve high toughness poly(butylene succinate) Zhong-Johnson EZL, Dong Z, Canova CT, Destro F, Cañellas M, Hoffman MC,
composites. Carbohydr Polym 206:827–836. https://doi.org/10.1016/j. Maréchal J, Johnson TM, Zheng M, Schlau-Cohen GS, Lucas MF, Braatz RD,
carbpol.2018.11.036 Sprenger KG, Voigt CA, Sinskey AJ (2024) Analysis of poly(ethylene tere-
Zhang Y, Pedersen JN, Eser BE, Guo Z (2022) Biodegradation of polyethylene and phthalate) degradation kinetics of evolved ispetase variants using a surface
polystyrene: from microbial deterioration to enzyme discovery. Biotechnol crowding model. J Biol Chem 300(3):105783. https://doi.org/10.1016/j.
Adv 60:107991. https://doi.org/10.1016/j.biotechadv.2022.107991 jbc.2024.105783
Zhao SL, Xi PF, Guo FY, Deng S, Jia JL (2020) Evaluation and screening of dioxin
control technology in waste incineration flue gas. Huan Jing Ke Xue
41(9):3985–3992. https://doi.org/10.13227/j.hjkx.202002170 Publisher’s Note
Zheng Y, Li Q, Liu P, Yuan Y, Dian L, Wang Q, Liang Q, Su T, Qi Q (2024) Springer Nature remains neutral with regard to jurisdictional claims in
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