Ghatge et al.

Appl Biol Chem (2020) 63:27

https://doi.org/10.1186/s13765-020-00511-3

REVIEW Open Access

Biodegradation of polyethylene: a brief

review
Sunil Ghatge1†, Youri Yang1†, Jae‑Hyung Ahn2* and Hor‑Gil Hur1*

Abstract 
Plastic waste management and recycling became a serious global issue as it affects living beings from all the eco‑
systems. Researchers investigated biodegradation of polyethylene (PE) by measuring changes in various physico-
chemical and structural characteristics using techniques like as fourier transform infrared spectroscopy (FTIR), scan‑
ning electron microscope (SEM), etc. However, these evidences are not enough to prove the exact biodegradation of
PE. In this review, we summarized microbial biodegradation of polyethylene and discussed recent developments for
the candidate microbial enzymes and their possible roles in PE degradation. In addition, we conversed the advanced
technologies correctly used for measuring PE degradation using isotope-labeled PE to figure out its metabolism into
the end products like as 13CO2.
Keywords:  Polyethylene, Biodegradation, Pre-treatment, Moth worms

Introduction plastics produced, and they are used for production of

Researches on various synthetic polymers have been plastic bags, disposable containers, bottles, packaging
accelerated since DuPont had succeeded in mass pro- materials, etc. [16]. An estimated more than 500 billion
duction and an exclusive sale of nylon in 1940. Versatile to 1 trillion plastic bags used globally disturb the eco-
properties of plastics, such as diverse application, con- system and ultimately result into serious environmental
venience, non-degradability, and low price, have led to issues of recycling these materials from the environment
the replacement of natural materials and common use as [4, 11, 20, 46, 91, 118].
household items since 1960. Worldwide the annual pro- The waste plastics that form particulate matter by UV
duction of non-degradable plastic ranges from 350 mil- irradiation and weathering increase surface area and
lion to 400 million tons out of that yearly, 5 to 13 million mobility and thereby incorporate easily into the food
tons of waste plastic are released into the ocean, which is chain, causing fatal injuries to all living organisms [14,
negatively affecting the ecological environment [47, 80]. 97]. In addition, the small size of suspended plastics
In fact, a survey on the Great Pacific Garbage Patch has causes the reduction of light transmission on the sea
estimated 80,000 tons of waste plastics have found in the surface, photosynthetic efficiency of micro-algae, and
Pacific latitudes, including Hawaii, with 54% of the waste the productivity of marine organisms. More seriously,
coming from North America and Asia [58]. Polyethylene the micro-plastic could act as a carrier to increase the
and polypropylene represent about 92% of the synthetic adsorption of hardly decomposable hydrophobic chemi-
cals such as polychlorinated biphenyl (PCB), when it is
introduced into the food chain [23, 86]. The groundwater
*Correspondence: [email protected]; [email protected] is also contaminated due to hazardous chemicals from
†
Sunil Ghatge and Youri Yang are contributed equally to this work
1
School of Earth Sciences and Environmental Engineering, Gwangju the plastic waste in landfills [72].
Institute of Science and Technology (GIST), Gwangju 61005, Republic Based on the annual production of various plastics such
of Korea as polyamide (PA), polyethylene (PE), polyethylene tere-
2
Agricultural Microbiology Division, National Institute of Agricultural
Sciences, Rural Development Administration, Wanju 55365, Republic phthalate (PET), polypropylene (PP), polystyrene (PS),
of Korea polyurethane (PU), and polyvinyl chloride (PVC), PE

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Ghatge et al. Appl Biol Chem (2020) 63:27 Page 2 of 14

and PP that have been widely used in Korea agriculture, polyethylene (LDPE) and high density polyethylene
comprise more than 60% of the total global plastic pro- (HDPE), have been used for biodegradation studies [86,
duction and the annual production of PE is 116 million 97] (Fig.  2). LDPE was prepared by the high pressure
tons [21]. In 2015, 6.7 million tons of plastic resins were polymerization of ethylene. The presence of branched
used in Korea, meaning that every single Korean con- chains is responsible for the low density of LDPE. Chemi-
sumed every single Korean consumed 132.7 kg of plastic cally, LDPE is inert at room temperature, however, it
per year. It represented that Korea is the second largest can be gradually attacked by strong oxidizing agents and
plastic consumer in the world. From the view on genera- some solvents, results in softening or swelling. It is intact
tion of waste plastics, about 10.1 million tons of waste for short period of time up to 95  °C and can be dura-
plastic generated in 2016 in Korea, indicating 196  kg of ble for longer hours at 80  °C. A degree of crystallinity
the waste per person per year (EUROMAP [29]). The of LDPE is within the range of 50–60% which provides
amount of waste plastics has been increasing every year several properties to the material such as opacity, tear
in Korea (Fig.  1). According to a survey by ministry of strength, tensile strength, rigidity and chemical resist-
environment in Korea, waste plastics from the agricul- ance, flexibility even at a low temperature [12, 33]. There
tural environment were 310,000 tons and among them,
200,000 tons were collected and 170,000 tons were recy-
cled (http://stat.me.go.kr). The non-recycled plastics left
in the agricultural environment without any management
will be getting worse due to a decrease in international oil
prices and increase of plastic waste.
PE is highly recalcitrant and inert material hence it is
very difficult to degrade in the environment even after
buried for several years as landfill. A polyethylene sheet
showed only partial degradation and negligible weight
loss when kept in moist soil for 12–32  years [77]. The
recalcitrance of PE is due to its water insolubility, the
hydrophobicity because of presence of linear backbone
of carbon atoms, degree of crystallinity, and its high
molecular weight [86, 111, 114]. PE such as low density Fig. 2  Structures of different types of polyethylene

Fig. 1  Statistics of domestic waste plastics in Korea from 2004 to 2017 (adopted from Environmental Statistics Portal [25], Ministry of Environment
of Korea, Republic of Korea 2010) (http://stat.me.go.kr), Ministry of Environment of Korea)
 Ghatge et al. Appl Biol Chem (2020) 63:27 Page 3 of 14

are two different types of LDPE, linear low density poly- morphology of the original polymer and facilitate the
ethylene (LLDPE) and branched low density polyethylene polymer biodegradation [59]. Oxidation of PE with nitric
(BLDPE). They show different density, degree of branch- acid has been known to promote fragmentation of PE
ing and availability of functional groups on the surface. films followed by microbial degradation [68]. The bio-
LDPE films are transparent, free from odor and toxicity. degradation of PE involves use of microbes or microbial
They have better ductility, low water vapor permeability, communities that modify and consume the polymer as a
and heat seal ability [78, 109]. It is widely used for pack- source of energy leading to changes in its physico-chem-
aging food and non-food items, manufacturing trays and ical properties such as weight loss, structural deteriora-
plastic bags. LDPE is utilized for mulching agricultural tion, and eventually carbon fixation as a biomass [86, 97].
fields and constructing polyhouse. It is also utilized for However, the formation of a biofilm by polymer-degrad-
coating on paper, textiles, and other plastics [97]. Nota- ing microorganisms on PE was restricted due to a high
bly, PE accounts for 97% of total agricultural waste vinyl degree of hydrophobicity, a low specific surface area and
and about 200,000 tons of LDPE were consumed, consist- smooth surface topography [62, 86]. In addition, produc-
ing of 64% of all types of agricultural waste vinyl (Fig. 3) tive adsorption and catalytic performance of polymer-
(http://stat.me.go.kr). HDPE is a PE thermoplastic, pro- degrading enzymes have been shown incompatible with
duced by a catalytic process and having little branching. a hydrophobic polymer surface [27, 87, 88, 92]. Besides,
It has stronger intermolecular forces and greater tensile worms of the moth, Galleria mellonella and Plodia inter-
strength than LDPE. The higher density provides greater punctella, have been observed to degrade untreated
stability due to reduced bond length and compact pack- LDPE [13, 63].
aging. Due to its hardness, opacity, and durability at In most of these biodegradation studies, PE degrada-
higher temperatures (up to 120  °C), it is widely used in tion was determined by measuring weight loss, observ-
industrial and day-to-day applications such as produc- ing polymer structural changes under SEM, and chemical
tion of carry bags, milk jugs, detergent bottles, margarine modifications of functional groups using FTIR [86, 97].
tubs, garbage containers, water pipes, etc. [8]. It has been criticized that the weight loss and surface
The extensive usage of both LDPE and HDPE poses topography changes are probably derived from the degra-
severe environmental threats to the terrestrial and dation of various additives, which often contribute to sig-
marine ecosystems, as experienced like blockages of PE nificant fraction of the PE. Hence, the results from many
in the intestines of fish, birds and marine mammals [10, of these studies need to be cross checked using more
99]. In addition, several hundreds of different species advanced physico-chemical, biochemical, and molecu-
from different ecosystems are on the edge of becoming lar biological technologies [21, 115]. Furthermore, the
endangered due to the ingestion of this waste [95, 104]. exact biochemical mechanisms and enzymes involved in
Inevitable use of PE to increase agricultural productiv- PE breakdown are still unknown. From the view on the
ity in small territory has caused a significant social issue. establishment of waste vinyl management, the researches
According to a report by Chinese Academy of Agricul- on finding microorganisms effectively degrading plas-
tural Sciences in 2014, mulching waste vinyl reduced tics and analyzing their physiology and application are
the movement of essential materials in soil such as air, essential.
moisture, and nutrients and the mobility of soil organ- In this review, we described the effect of pretreatment
isms including earthworms [60]. This led to a decrease in and additives for enhancing biodegradability of PE and
soil quality, physiological disorders in plant growth such summarized recent studies on biodegradation of PE by
as seed germination and root growth, which would which various bacteria, fungi, and moth worms along with gut
would decrease agricultural productivity in the end. Thus, microorganisms from worms. In addition, we discussed
the use and waste treatment for plastics have become a candidate microbial enzymes involved in PE biodegrada-
global problem. Therefore, it is of inevitable necessity to tion and various advanced technologies used for evaluat-
minimize PE and other plastics and to develop efficient ing the actual biodegradation of PE.
methods for plastic degradation and recycling.
Extensive research has been carried out for degrada- Pretreatments and effects of additives
tion of PE either by physico-chemical or microbial meth- for enhancing biodegradability of PE
ods or combination of both [14, 86, 97]. Physico-chemical Thermo‑UV pretreatment
technologies include thermal and UV treatment or com- Thermo-UV pretreatment was used to induce partial
bination of both, which reduces the polymer chain size photolysis of the PE film and to simulate weathering of
and form oxidized groups such as carboxyl, carbonyl PE that occurs in nature (e.g., PE used for soil mulch-
and hydroxyl, on the surface of polymer [19, 44, 57, 68]. ing or polyhouse cover). PE samples were treated in a
These treatments modify the crystallinity and surface QUV accelerated weathering tester developed by QLAB,
Ghatge et al. Appl Biol Chem (2020) 63:27 Page 4 of 14

Fig. 3  Statistics on the treatment of waste vinyl from agriculture (Top) and types of agricultural waste vinyl (Bottom) in Korea from 2004 to 2017
(adopted from Environmental Statistics Portal [25], Ministry of Environment of Korea, Republic of Korea 2010) (http://stat.me.go.kr), Ministry of
Environment of Korea)

Homestead, FL. Herein, PE was alternatively exposed to incubated with Rhodococcus ruber strain C208 showed
cycles of UV and humidity: five cycles per d of UV expo- that formation of the carbonyl residues on the surface
sure (four of 4 h each, one of 3 h at 70 °C) separated by of photooxidized PE plays an important role in initia-
1-h intervals (50  °C). The pretreated PE film was tested tion of biodegradation [38, 101]. In another study, Bacil-
for biodegradation and biofilm formation by Rhodococ- lus amyloliquefaciens strain isolated from composed
cus ruber (C208). FTIR spectra of UV-photooxidized PE plastic was studied for assessing deterioration effect
 Ghatge et al. Appl Biol Chem (2020) 63:27 Page 5 of 14

of thermo-irradiation pretreatment. Herein LDPE and however, it is not known how microbes participate in
LLDPE films were exposed to gamma rays followed by polymer chain cleavage and what kind of enzyme system
thermal treatment at 150  °C and 90  °C for 7 d, respec- is involved in this process.
tively. Furthermore, LDPE films were additionally
exposed to UV irradiation in an oven at 60 °C for 7 d [73]. Photo‑catalysis using titanium dioxide (­ TiO2)
When the pretreated LLDPE was incubated with the iso- TiO2 is an ecofriendly photocatalyst that absorbs light in
lated bacterium for 40–60 day, the dry weight of LLDPE the UV region. Hence, T ­ iO2-incorporated polymer films
was slightly decreased by 1.1  ±  0.3 to 3.2  ±   1.3%, with efficiently absorb UV light. T ­ iO2 mediated photocataly-
flattening of carbonyl band (1300–1100 cm−1) in the sis involves the absorption of photons of suitable energy
FTIR spectra, indicating a biodeterioration. Electrospray leading to the generation of electrons and holes which
ionization-mass spectrometry (ESI-MS) analysis showed promote the formation of free radicals, resulting in the
the release of 3-hydroxybutyrate oligomers only in the oxidation of the polymer, followed by its degradation.
medium containing pretreated LLDPE and not with Thankam thomas and Sandhyarani et  al. (2013) investi-
native LLDPE. These oligomers disappeared after incuba- gated photocatalytic degradation of LDPE incorporated
tion with Bacillus amyloliquefaciens indicating metabo- with titania nanoparticles after treatment with solar
lism of low molecular LLDPE fractions. radiation. The composite PE film showed weight loss of
Balasubramanian et al. [9] used combination of physi- 68% after exposer to the solar radiation for 200 h which
cal, chemical and biological treatment with Aspergillus is a significantly higher compared with a study where
terreus  MF12 for enhancing HDPE degradation. HDPE the similar rate of weight loss was observed after 400  h
film initially heated at 50  °C for 72  h followed by expo- [108, 125]. FTIR and SEM analysis revealed the presence
sure to UV (312 nm) and humidity. Secondly, HDPE film of carbonyl groups and the creation of holes at the inter-
was chemically treated by immersing into ­KMnO4/HCl at face, respectively indicating the degradation of the LDPE.
a concentration of 0.25/0.5 mol l−1 at 45 °C for 8 h [32] Another study showed that solar irradiation of a PE film
and 10% citric acid for 8 h at 45 °C. The HDPE degrada- blended with copper phthalocyanine (CuPc) modified
tion gradually increased from 9.4  ±  0.1 to 20.8  ±   0.1% ­TiO2 ­(TiO2/CuPc) photocatalyst resulted in significant
between the physico-chemical and biological treatments, weight loss rate, rough surface texture, higher amount of
indicating synergism between biotic and abiotic factors generated ­CO2, compared to the original PE film [126].
for HDPE degradation by A. terreus MF12 [9]. Surface photovoltage spectroscopy (SPS) analysis sug-
gested that CuPc promoted charge separation of ­TiO2.
Treatment with pro‑oxidants Reactive oxygen species generated on the surface of ­TiO2
All commercial prepared PE films contain a small amount or ­TiO2/CuPc particles are responsible for enhanced
of a stabilizer that prevents oxidation during processing degradation of PE. Recently, Fa et  al. [31] synthesized
and significantly prolongs its lifetime [15]. Supporting ­TiO2-FeSt3 ferric stearate-polyethylene (TFPE) compos-
pro-oxidant additives to enhance the photo- and thermo- ite film and studied photo-degradation by treating UV
oxidation of PE films may lead to radical reactions that irradiation for 240 h and/or thermo-degradation at 70 °C
result in polymer chain cleavage [54, 55]. Conventional for 30  d. FTIR spectroscopy confirms the formation of
pro-oxidants include transient metal ions such as iron, carbonyl and hydroxyl group which assist in biodegrada-
manganese, titanium, cobalt in the form of stearate. Iron tion of PE films. The tensile strength and elongation at
and titanium complexes as a source of radicals initiate break of TFPE film reduced to 60% and 97.7%, respec-
photo-oxidation, while manganese and cobalt catalyze tively [31].
peroxidation without light. Study on biodegradation of
PE (LDPE) treated with pro-oxidant, manganese stearate Biodegradation of PE
followed by UV irradiation and treatment with Aspergil- The biodegradation of recalcitrant PE has been investi-
lus oryzae resulted in 62 and 51%, decrease in percentage gated by many researchers [28, 40, 56, 62, 98, 100, 127].
elongation and tensile strength, respectively. Further- Still, the complex biodegradation mechanism of PE is not
more, FTIR analysis confirmed the formation of more yet fully understood. It is suggested that various abiotic
carbonyl and carboxylic groups after treatment with and biotic factors play a vital role in the biodegradation
pro-oxidant over UV treated film, which was completely of PE in the environment [62, 100]. Biodegradation stud-
degraded after incubation with A. oryzae thus confirm- ies have been accomplished either using pure cultures
ing role of pro-oxidants in enhancing PE biodegradation that are able to degrade PE [2, 5, 8, 34, 38, 41, 54, 81, 84,
[52]. Although it is clear that lower molecular weight 101, 110, 113, 117, 119] or using complex microbial com-
products were formed by the catalytic action of pro- munities from various terrestrial (soil from landfill sites,
oxidants, which were consumed by the microorganisms, composting) and marine habitats [1, 3, 6, 18, 49, 61, 71,
Ghatge et al. Appl Biol Chem (2020) 63:27 Page 6 of 14

74, 76, 79]. In addition, it was reported that bioaugmenta- and biosurfactant suggesting their importance in poly-
tion with tailored microbial consortia could facilitate the mer oxidation and biodegradation [5]. Tribedi and Sil
biodegradation of PE [107]. Furthermore, significantly [110] showed that the addition of mineral oil to the LDPE
faster biodegradation of PE was observed by waxworms. degradation medium of Pseudomonas sp. strain AKS2
stimulated the hydrophobic interaction to form biofilms
Bacterial biodegradation of PE on polymer surfaces and degraded 5 ± 1% of the original
More than 20 bacterial genera have been shown to PE material for 45 d, whereas Tween 80 had an adverse
degrade different types of PE. Those include various effect to the biofilm formation. A thermophilic bacte-
Gram-negative and Gram-positives species belonging to rium Brevibacillus borstelensis isolated from soil has
the genera Pseudomonas, Ralstonia, Stenotrophomonas, been reported to utilize BLDPE as the sole carbon and
Klebsiella, Acinetobactor, etc. and Rhodococcus, Staphy- energy source, by which 30% of the molecular weight of
lococcus, Streptococcus, Streptomyces, Bacillus, etc. [21, PE film was reduced during an incubation period of 30 d
39, 86, 97].. Most of these bacterial strains possess the [41]. [38] isolated a biofilm producing Rhodococcus ruber
ability to deteriorate surface and/or form a biofilm on (C208) strain which degraded PE at a rate of 0.86% per
PE. Table 1 summarizes bacterial strains associated with week. A hydrophobic cell surface (e.g., mycolic acid layer)
PE biodegradation. Studies on diverse activities of the of this strain may play an important role in biofilm for-
genus Pseudomonas have been carried out to investigate mation on PE surface [38, 101]. Awasthi et al. [7] reported
their capabilities to degrade and metabolize a variety of that HDPE after thermal treatment was degraded by
synthetic plastics polymers and the by-products. Pseu- Klebsiella pneumoniae. This strain was able to strongly
domonas species have the unique ability to degrade and adhere to HDPE surfaces, leading to increasing biofilm
metabolize the polymers with extracellular oxidative and/ thickness with decreasing weight and tensile strength
or hydrolytic enzyme activities, which facilitate uptake of the HDPE film by 18.4% and 60%, respectively, in 60
and degradation of the polymer fragments, and control d. Possible biodegradation of an HDPE film exhibits the
interaction between biofilms and polymer surfaces [117]. SEM and atomic force microscopy (AFM) images of sub-
Complete degradation of PE in water was observed after surface corrosion, cracks, and surface roughness pro-
treatment with P. fluorescens in presence of surfactant duced by bacteria.

Table 1  Bacterial strains associated with PE degradation

Bacterial strain Substrate Country of origin References

Streptomyces badius, S. setonii, starch-PE USA [81]

Arthrobacter paraffineus LDPE, HDPE Sweden [2]
Brevibacillus borstelensis LDPE Israel [41]
Nocardia asteroids LDPE, HDPE France, Czech Republic [54]
Rhodococcus rhodochrous LDPE, HDPE France, Czech Republic [54]
Bacillus halodenitrificans LDPE India [91]
Bacillus sphericus LDPE, HDPE India [105]
Arthrobacter sp. HDPE India [8]
Pseudomonas sp. HDPE India [8]
Staphylococcus epidermidis LDPE India [17]
Rhodococcus rhodochrous LDPE, HDPE, LLDPE France, Belgium, Italy [34]
Arthrobacter viscosus, Acinetobacter baumannii LDPE Poland [74]
Bacillus amyloliquefaciens, B. cereus B. circulans, B. mycoides, B. pumilus, B.
thuringienesis, M. luteous, M. lylae
Pseudomonas fluorescens, Paenibacillus macerans, Rahnella aquatilis LDPE Poland [74]
Staphylococcus cohnii LDPE Poland [74]
Staphylococcus xylosus LDPE Poland [74]
Microbacterium paraoxydans LDPE India [84]
Pseudomonas aeruginosa LDPE India [84]
Rhodococcus ruber C208 LDPE Israel [93]
Pseudomonas sp. AKS2 LDPE India [110]
Klebsiella pneumonia HDPE India [7]
 Ghatge et al. Appl Biol Chem (2020) 63:27 Page 7 of 14

Fungal biodegradation of PE fungal strains leading to biofilm formation and morpho-
In addition to bacteria, several fungal genera including logical changes on LDPE and HDPE surfaces, including
Aspergillus, Cladosporium, Fusarium, Penicillium, Phan- cracks, pits, and undulations [75].
erochaete have been reported for PE degradation [21, 39,
86, 97]. In general, fungi are thought to be more efficient Role of Waxworm and gut microbiome in biodegradation
than bacteria for the degradation of PE because they are of PE
capable of attaching to the hydrophobic surface of the Larvae of Galleria mellonella and Plodia interpunctella,
polymers [51, 96, 97], producing extracellular enzymes have been reported to degrade LDPE without pretreat-
targeting insoluble fibers, and surviving in stressful ment [13, 63]. The worms could soften thin-film PE
growth conditions [98]. Table 2 summarizes some fungal shopping bags and metabolize them to ethylene glycol.
strains capable of degrading PE. Weight loss measure- Because there is a structural similarity between beeswax
ment is a commonly used method to analyze biodegra- and PE, the biochemical machinery for beeswax metabo-
dation of PE. For examples, biodegradation of LDPE by lism of G. mellonella makes it useful for PE metabolism.
A. niger and A. japonicas in laboratory conditions have About 100 worms of G. mellonella can cause a weight
been found to decrease the dry weight by 5.8% and 11.1% loss of 92 mg from a commercial PE shopping bag within
per month, respectively [82]. Das and Kumar [66] stud- 12  h. [115] criticized that the waxworm research lacked
ied microbial deterioration of LDPE by Aspergillus and the necessary information to support the claims of the
Fusarium sp. Among them Fusarium sp. FSM-10 and original G. mellonella report. Although the waxworm
Aspergillus sp. FSM-3 showed maximum weight reduc- researches lacked information about the biodegradation
tion about 8–9%, while only 5% weight loss was observed mechanism of PE [115], cutting holes on the surface of
by Aspergillus sp. FSM-5 after 60 d of incubation. Usha PE by waxworms and FTIR analysis of degraded PE indi-
et  al. [112] isolated strains belonging to Aspergillus fla- cated that PE pieces break down with the carbon–carbon
vus and A. nidulans through enrichment culture show- bond cleavage by mechanical force or enzymatic diges-
ing the clearing zone around their colonies on PE agar tion or both. Recently, biodegradation of PE by Entero-
plates. Kathiresan [50] isolated PE degrading fungi from bacter sp. D1 from the guts of wax moth G. mellonella
mangrove soils. Yamada-Onodera et  al. [119] reported has been investigated [85]. The authors performed AFM
degradation of additives free PE by P. simplicissimum and SEM analysis to show that the strain D1 was able to
YK. Esmaeili et al. [26] isolated A. niger from soils of PE form colonies around a PE film after14 d of incubation
wastes landfills using mineral medium with PE powder as and disrupted the PE film surface. The treatment of a PE
a sole carbon source. From SEM and AFM analysis of the film with the strain D1 has highlighted that the appear-
PE surface, several strains of Chrysonilia, Aspergillus, and ance of carbonyl and ether functional groups on the
Penicillium species have been isolated using synthetic FTIR spectra was concomitant with the release of oxi-
medium [67]. Among the isolated fungi, P. chrysogenum dative cleavage products containing alcohol, ester and
NS10 (KU559907), P. oxalicum NS4 (KU559906) have acid groups, analyzed by high-performance liquid chro-
been evaluated for HDPE and LDPE degradation using matography-mass spectrometry. This study indicates the
the response surface methodology to optimize the growth involvement of strain D1 in PE degradations. However, it
media for increasing the mycelium weight. The AFM and is still unclear that PE is degraded by G. mellonella lar-
SEM analysis is widely used to ratify PE degradation by vae or by the gut flora. Recently, complete genomic and

Table 2  Fungal strains associated with PE degradation

Fungal strain Substrate Country of origin References

Fusarium redolens LDPE C14 labelled; UV treated Sweden [1, 49]

Verticillium lecanii LDPE Sweden [49]
Phanerochaete chrysosporium LDPE/starch Turkey [76]
Penicillum simplicissimum HDPE UV treated Japan [119]
Aspergillus niger Thermal treated LDPE USA [113]
Penicillum pinophilum Powdered LDPE USA [113]
Cladosporium cladosporioides Degradable polyethylene green film France, UK [14]
Glioclodium virens Thermal treated LDPE Mexico [64]
Aspergillus flavus LDPE and HDPE film France, Czech Republic [54]
Ghatge et al. Appl Biol Chem (2020) 63:27 Page 8 of 14

transcriptomic data of G. mellonella have been used to polymer: functional groups on the surface, hydropho-
explore the beeswax and PE metabolism [53]. Notably, bicity/hydrophilicity, crystallinity, molecular weight dis-
the long chain fatty acids from beeswax and PE were tribution, surface topography, mechanical properties,
detected even in the absence of gut microbiota indicat- and mass balance (Table  3). The methods used to study
ing that the PE metabolism can occur without help of gut these changes have been extensively reviewed in previous
microflora. studies [86, 97]. Hence we will just briefly discuss each
In another study, two PE-degrading strains, named of these 7 characteristics for understanding PE degrada-
Bacillus sp. YP1 and Enterobacter asburiae YT1, were tion. FTIR spectroscopy is used to study the formation of
isolated from the gut of P. interpunctella larvae [123]. various functional groups on the surface of PE after the
They reduced the hydrophobicity and caused surface dis- abiotic and biotic oxidation by thermo-UV treatment
ruption of PE film after 28 d of incubation. After 60 d of and microbial degradation. For example, UV and nitric
incubation with these strains approximately 10.7 ± 0.2% acid treatment to PE led to an increase in absorbance of
and 6.1 ± 0.3% of the PE films (100  mg) were degraded, infrared at 1710–1715  cm−1 (corresponding to carbonyl
respectively. These findings demonstrated the impor- group) and 1640  cm−1 and 830–880  cm−1 (correspond-
tance of gut microbes of moth larvae for PE biodegra- ing to –C=C–), which was then reduced after incubation
dation. Yang et  al. [120] explored biodegradation of PE with microbial consortia [43]. Similarly, Harshvardhan
and plastic mixtures by yellow mealworms i.e. larvae of and Jha [42] reported PE biodegradation with an increase
Tenebrio molitor. Up to 49.0 ± 1.4% of the ingested PE in the index of carbonyl bond, the keto carbonyl bond,
was metabolized to C ­ O2 after incubation with larvae. and the vinyl bond, calculated using FT-IR spectra. These
PE-fed mealworms showed 40.1 ± 8.5% reduction in the functional groups at the surface of PE are considered
molecular weights of egested polymer. The gut micro- important because oxidized groups cause an increase in
biome study with next generation sequence analysis has the hydrophilicity which in turns results in the efficient
shown the abundance of Citrobacter sp. and Kosakonia attachment of microbes to the PE surface thereby pro-
sp. attached to PE. Both of them are members of the moting the biodegradation [2, 110]. The hydrophilicity is
Enterobacteriaceae family which is known to contain PE usually estimated by measuring the contact angle of the
degrading bacteria’s [123]. In addition, Citrobacter sp. surface with water. A small contact angle with water indi-
(aerobic), and Kosakonia sp. (facultative anaerobic) can cates a high hydrophilicity of the oxidized PE surface [91,
utilize oxygen which suggests their involvement in plastic 105].
degradation [37, 56, 98, 111]. Further studies are required Crystallinity is another important parameter to predict
to reveal the mechanism of enzymatic degradation in the the extent of biodeterioration of the polymer, measured
guts of mealworms and waxworms, which will facilitate with the help of Differential Scanning Calorimetry (DSC)
the biodegradation of a variety of plastic materials. and FTIR analysis. Generally, the amorphous region
is easily accessible and degraded by microorganisms,
Effects of microbial activity on PE resulting in an initial increase in crystallinity [5, 44, 83].
The effect of microbial colonization on the surface of PE After attack (or dwell) in the amorphous regions, micro-
was studied by monitoring seven different characteris- organisms will start degrading the crystalline region and
tics with respect to the degree of biodegradation of the increase in the proportion of larger crystals [2, 105]. Size

Table 3  Techniques used for characterization of PE biodegradation

Techniques used Changes in PE Measured characteristics References

FTIR Functional groups on the surface Keto-carbonyl index (I1715/I1565); Ester-carbonyl index [3, 6, 34, 38, 41, 74,
(I1740/I1465); Vinyl-bound index (I1640/I1465); 83, 93, 94, 105,
Double bound index (I908/I1465); 113]
C-O stretching (I1100)
Contact angle drop deposition Hydrophobicity/hydrophilicity Contac angle with water, Surface energy [6, 91, 105]
FTIR/DSC/XRD Crystallinity Crystallinity; Melting temperature; Relative crystallinity; Lamel‑ [2, 76]
lar thickness
HT-SEC/GPC Molecular weight distribution Changes in the molecular weight [38];
[41];
SEM/AFM Surface topography Molecular weight distribution topography [6, 105]
Instron Mechanical properties Tensile strength; Strain energy;  % Elongation and extension [8, 14, 18]
Gravimetric ­CO2 evolution Consumption of the polymer Weight loss [34, 74, 93]
 Ghatge et al. Appl Biol Chem (2020) 63:27 Page 9 of 14

exclusion chromatography and time of flight mass spec- have been involved in the biodegradation of PE [56,
trometry (TOF–MS) analysis gives an idea about molec- 86]. These include manganese peroxidase (MnP, EC
ular weight distribution of the PE after biodegradation. 1.11.1.13), lignin peroxidases (LiP, EC 1.11.1.14), and
An increase in the average of molecular weight is wit- laccases (EC 1.10.3.2.) [116]. A copper dependent lac-
nessed after initial degradation of low molecular weight case from R. ruber strain C208 was reported to degrade
chains [5, 41, 81, 119]. UV pretreated PE films [93]. Laccase mediated oxida-
SEM and AFM analyses are commonly employed to tive cleavage of amorphous region of PE films results
investigate surface topography of PE films during biodeg- in formation of easily accessible carbonyl groups and
radation. Biofilm formation on the surface of the poly- significant decrease in weight of a PE film. Fujisawa
mers [14, 34, 38, 54, 101, 110] and penetration of hyphal et  al. [36] showed reduction of the molecular weight
structures [44, 83, 113] have been generally observed of a PE membrane after treatment with laccase from
after incubation with microbes. The structural changes Trametes versicolor in presence of 1-hydroxybenzo-
in the formation of pits, holes, and erosions have been triazole as a mediator. Degradation of a high molecular
observed under SEM indicating surface destruction of weight PE membrane by P. chrysosporium ME-446 and
PE. Alterations in crystallinity and the average molecular an isolate IZU-154 have been described and MnP from
weight as a result of oxidation modify the chemical and this white-rot fungus was found to be the key enzyme
mechanical properties of PE. Universal mechanical test- responsible for PE degradation [45]. A partially puri-
ing system (UMTS) is preferentially used for studying fied MnP stimulated degradation of PE in presence of
changes in the mechanical properties of a polymer [74, various surfactants [24, 45]. The most active MnP from
79, 105]. However, this method is prone to underestimate IZU-154 has been characterized in regard to the oxida-
the local surface related damage caused by the micro- tion of 2,6-dimethoxyphenol [65] and the degradation
organisms. Microorganisms utilize PE as a sole carbon of nylon-66 [22]. Enhanced extracellular secretion of
source and metabolize it to C ­ O2 during respiration and laccases and MnP from B. cereus was observed when
hence measurement of released ­CO2 can be linked to the the strain was incubated with UV-irradiated PE [102].
amount of polymer consumed. Some studies reported However, the same PE film treated with a partially puri-
a decrease in the weight of samples measured either by fied laccase and a MnP from P. simplicissimum showed
gravimetric measurements [6, 41, 74, 101, 105] or by ­CO2 negligible weight loss [103]. The LiP activity in the
emissions from the samples [1, 3, 49, 96]. Progressive concentrated culture supernatants of lignocellulose
­CO2 emissions in the samples are measured to define the degrading Streptomyces species has been reported to
total degradation of the polymer along with its rate. responsible for degradation of a heat treated PE [81].
Similarly, up to 70% degradation of a pre-oxidized high
Microbial enzymes involved in PE biodegradation molecular weight PE sample has been reported after 15
The biodegradation of plastics is a complex process d of treatment with P. chrysosporium strain MTCC-787.
involving various abiotic and biotic factors [28, 40, 56, The extracellular peroxidases play a vital role in the bio-
62, 98, 100, 127]. The cooperative action of abiotic fac- degradation of PE by this strain [70].
tors and microorganisms causes fragmentation of the AlkB family alkane hydroxylases (AH) (EC 1.14.15.3)
bulk polymer to increase accessible surfaces for biodeg- can degrade hydrocarbon oligomers by terminal or
radation. Some extracellular enzymes carry out further sub-terminal oxidation [89]. Yoon et al. [124] reported
fragmentation of the polymers [62, 100] (Fig. 4). Several action of a recombinant AH from Pseudomonas sp. E4
lignin degrading enzymes also participate in the break- at 37  °C for 80 d converted 20% of the low molecular
down of the PE thermoplastic [86, 100]. After the initial weight PE sample to C ­ O2. Expression of the complete
cleavage of the polymer into oligomers of 10–50 carbon AH system of P. aeruginosa strain E7 in E. coli showed
atoms which can be transported into the cell for further 30% degradation of a PE sample [48]. These studies uti-
metabolism [62, 86]. Biodegradation of PE is restricted lized crude or partially purified enzymes and required
by the absence of hydrolysable functional groups in the long treatment time. Notably, the use of tailored micro-
backbone [56, 86]. The carbonyl and hydroxyl groups bial consortia has shown promising degradation of PS
formed from various pretreatment such as thermo- and PE compared to the use of single microorganisms
UV irradiation or addition of oxidizing agents could be [26, 69, 91, 121]. Recently, transcriptome analysis of G.
adopted to stimulate biodegradation further [34, 54] a, mellonella fed on beeswax similar to PE showed, upreg-
b). Hence most of the biodegradation studies on PE have ulation of genes encoding carboxylesterase, lipase, and
been carried out using substrate peroxidation [86]. enzymes related to fatty acid metabolism. However, the
Microbial enzymes capable of degrading lignin detailed mechanisms of these enzymes have not been
polymer containing oxidizable C–C bonds [35, 106] investigated.
Ghatge et al. Appl Biol Chem (2020) 63:27 Page 10 of 14

Fig. 4  Mechanism of abiotic and biotic degradation of polyethylene

Recent developments in the analytical methods of additives [21, 86, 97]. In this regard, a robust, reliable
to correctly estimate PE degradation method has been introduced to assess biodegradability
Many researchers investigated PE degrading microor- of PE via the quantification of ­CO2 using gas chroma-
ganisms using commercially available polymers that tography as a result of bacterial degradation and respira-
possibly contain various chemical additives. The extent tion [90]. Herein, the soil bacterium R. rhodochrous was
of degradation was estimated by calculating weight loss grown in a defined aqueous medium with PE as the only
and functional group changes on the surface of polymer carbon source and the production of C ­ O2 was directly
by FTIR. However, it is very obscure that the weight loss related to the mineralization of the added carbon source
and surface structure changes result from degradation of via bacterial respiration. At stationary phase, no signifi-
additives, which often contribute a major portion of the cant difference in the release of C
­ O2 between cells grown
polymer. Hence, more substantial changes are needed to with no carbon source and with LDPE. This suggested
distinguish the actual degradation of PE and minimiz- that bioavailability of carbon was limited in bacterial
ing chances of artifacts originated from the degradation growth on LDPE hence showing limited biodegradation.
 Ghatge et al. Appl Biol Chem (2020) 63:27 Page 11 of 14

Furthermore, the effect of UV pretreatment on biodeg- biodegradation of PE in order to minimize artifacts

radability of LDPE was studied by incubating R. rho- formed from degradation of additives rather than PE.
dochrous with UV pretreated LDPE and native LDPE Hence, upcoming research should be performed using
for 35 d and released ­CO2 was measured over the time. additive-free PE. In addition, 13C-polyethylene degrada-
The biodegradation of UV treated LDPE was threefold tion is proposed to show the formation of 13C-labeled
greater than non-treated LDPE. Authors also tested metabolites including ­CO2 emissions, progressively dur-
growth of alkanes degrading marine bacteria Alcanivo- ing an incubation period of time. Further investigations
rax borkumensis on LDPE. Negligible difference between on the mechanism of enzymatic degradation will high-
­CO2 generated by A. borkumensis on LDPE and the sam- light the pathway for an efficient biodegradation of PE at
ple without LDPE indicating that the strain was not able molecular levels.
to utilize LDPE.
In another study, 1320 oxidized oligomers from PE
Abbreviations
films before and after biodegradation with R. rho- PE: Polyethylene; FTIR: Fourier transform infrared spectroscopy; SEM: Scan‑
dochrous have been characterized by MS and nuclear ning electron microscope; PA: Polyamide; PET: Polyethylene terephthalate;
magnetic resonance (NMR) spectroscopy [30]. The strain PP: Polypropylene; PS: Polystyrene; PU: Polyurethane; PVC: Polyvinyl chloride;
LDPE: Low density polyethylene; HDPE: High density polyethylene; LLDPE:
can assimilate 95% of the soluble oligomers after 240 d. Linear low density polyethylene; BLDPE: Branched low density polyethylene;
Notably, longer molecules degraded quickly than the ESI–MS: Electrospray ionization-Mass spectrometry; TiO2: Titanium dioxide;
smaller ones, suggesting that both extracellular chain CuPc: Copper phthalocyanine; SPS: Surface photovoltage spectroscopy;
TFPE: TiO2-FeSt3 ferric stearate-polyethylene; AFM: Atomic force microscopy;
cleavage and intracellular β-oxidation mechanisms play HPLC-MS: High-performance liquid chromatography-mass spectrometry; DSC:
an important role in PE biodegradation. In addition, Differential Scanning Calorimetry; TOF-MS: Time of flight mass spectrometry;
there are several reports on assessing biodegradability of UMTS: Universal mechanical testing system; MnP: Manganese peroxidase;
LiP: Lignin peroxidases; AH: Alkane hydroxylases; NMR: Nuclear magnetic
plastic based on carbon tracing from polymers into C ­ O2 resonance; PBAT: Poly(butylene adipate-co-terephthalate); CP/MAS NMR:
and biomass. Zumestein et  al. [128] used 13C-labeled Cross-polarization/magic angle spinning nuclear magnetic resonance.
polymer poly(butylene adipate-co-terephthalate) (PBAT),
Authors’ contributions
and investigated biodegradability in the soil. Various SG and YY written the review and equally contributed for this manuscript;
soil microorganisms and filamentous fungi were found HGH and JHA revised the manuscript and share the corresponding author‑
to utilize carbon from each monomer unit of PBAT as a ship. All authors read and approved the final manuscript.
carbon and energy source. Biodegradation and minerali- Funding
zation of PS by Tenebrio molitor have been studied using This study was carried out with the support (PJ01497402) of National Institute
13 of Agricultural Sciences, Rural Development Administration, Republic of Korea.
C-labeled PS [121, 122]. The analysis of fecula egested
from Styrofoam-feeding larvae was performed using 13C Availability of data and materials
cross-polarization/magic angle spinning nuclear mag- All data sets presented in this study are included in the published article.
netic resonance (CP/MAS NMR) spectroscopy, which
Ethics approval and consent to participate
showed that 47.7% of the gulped Styrofoam carbon was Not applicable.
transformed into C­ O2 and 49.2% residue was excreted as
fecula with only 0.5% incorporation into biomass. Thus, Consent for publication
Not applicable.
tests with labeled PS indicated the degradation of PS
into 13CO2 and incorporation into lipids [120]. Recently, Competing interests
a mass balance study showed that PE-fed mealworms of The authors declare that they have no competing interests.
Tenebrio molitor converted 49.0 ± 1.4% of the ingested Received: 19 March 2020 Accepted: 6 May 2020
PE into ­CO2 [120]. In addition, 1H-NMR data were used
to determine the chemical modifications in the residual
polymer from the excreta of PE-fed mealworms, in com-
parison with bran-fed mealworms. References
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