REVIEW

Narancic and O’Connor, Microbiology 2019;165:129–137

DOI 10.1099/mic.0.000749

Plastic waste as a global challenge: are biodegradable plastics

the answer to the plastic waste problem?
Tanja Narancic1,2 and Kevin E. O’Connor1,2,*

Abstract
The strength, flexibility and light weight of traditional oil-derived plastics make them ideal materials for a large number of
applications, including packaging, medical devices, building, transportation, etc. However, the majority of produced plastics
are single-use plastics, which, coupled with a throw-away culture, leads to the accumulation of plastic waste and pollution,
as well as the loss of a valuable resource. In this review we discuss the advances and possibilities in the biotransformation
and biodegradation of oil-based plastics. We review bio-based and biodegradable polymers and highlight the importance of
end-of-life management of biodegradables. Finally, we discuss the role of a circular economy in reducing plastic waste
pollution.

PLASTICS AS POLLUTANTS incinerated [2]. In the USA, 53 % of total municipal solid

Traditional plastics derived from fossil oil are a family of
hundreds of materials with a wide range of properties. Their
strength, flexibility, light weight, and easy and low-cost pro-
duction make them ideal materials for application in a large
number of industrial and consumer products. Due to their
versatility, plastics are key materials in packaging, building,
transportation, medical devices, etc. In 2016, 335 million
tonnes of plastic were produced globally, reflecting their
popularity and widespread application [1]. Plastics help
society to reduce food waste by, for example, providing bet-
ter barrier properties and thus delaying food spoilage. They
also allow energy savings in transportation and thus
decrease greenhouse gas (GHG) emissions, e.g. plastic parts
in cars are estimated to reduce the environmental impact of
cars by fourfold compared to when other traditional materi-
als are used [1]. In addition, the plastics industry in the EU
employed 1.5 million people and generated a turnover of
e340 billion in 2015 [2].
However, the majority of the produced plastic is a single-
use plastic. Approximately 40 % of the plastic produced
goes to the packaging sector [1]. Thermoplastics polyethyl-
ene (PE), polyethylene terephthalate (PET), polypropylene
(PP) and polystyrene (PS) (Fig. 1) are the most frequently
used plastics in packaging and represented over 60 % of the
total plastics demand in Europe in 2016 [1]. This, coupled
with a single-use culture, carries two major consequences:
pollution and the loss of a valuable resource. In the EU,
70 % of the collected plastic waste ends in landfills or is
waste (MSW) is landfilled, with plastic waste representing
13 % of MSW [3]. The potential for recycling plastic waste
remains largely unexploited, with very low global rates of
recycling [4] and plastic recyclate representing only 6 % of
total demand for plastics [5]. The high price and low quality
of the recycled plastics relative to virgin plastics limits mar-
ket applications.
Most of the oil-based plastics are highly resistant to biodeg-
radation, which means that once they reach the environ-
ment they will inevitably accumulate, resulting in negative
environmental consequences. There is a long-term risk of
the release of hazardous chemicals from plastic waste in the
landfills and the contamination of ground waters [6]. Sin-
gle-use plastics make up 50 % of marine and beach litter in
Europe [7]. Plastic debris is a major pollutant in the world’s
oceans that kills hundreds of thousands of sea turtles, seals,
whales and seabirds due to ingestion or entanglement
[8–10]. There is also a growing concern about microplastic
pollution. While we do not exactly know how much of it is
out there, there are studies showing that environmentally
relevant concentrations of microplastics have a negative
effect on sea life [11, 12]. Moreover, microplastics have been
shown to impact on the biophysical properties of soil, bring-
ing about functional changes in soils [13]. Therefore, micro-
plastics might induce a global change of terrestrial
ecosystems, rather than exhibiting a direct lethal effect.
The waste management strategies proposed by the Euro-
pean Union (EU) and United States Environmental

Received 11 September 2018; Accepted 9 November 2018; Published 30 November 2018

Author affiliations: 1BEACON - Bioeconomy Research Centre, Ireland, University College Dublin, Belfield, Dublin 4, Ireland; 2UCD Earth Institute and
School of Biomolecular and Biomedical Science, University College Dublin, Belfield, Dublin 4, Ireland.
*Correspondence: Kevin E. O’Connor, kevin.oconnor@ucd.ie
Keywords: fossil-based plastic; plastic waste; biodegradable plastics; circular economy.

000749 ã 2019 The Authors

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 Narancic and O’Connor, Microbiology 2019;165:129–137

Fig. 1. Chemical structures of oil-derived, non-biodegradable polymers [polyethylene terephthalate (PET), polyethylene (PE), polypro-
pylene (PE), polystyrene (PS) and polyvinyl chloride (PVC)], bio-based non-biodegradable polyethylene-2,5-furandicarboxylate (PEF),
oil-based biodegradable polybutylene adipate terephthalate (PBAT) and polycaprolactone (PCLA), and bio-based and biodegradable pol-
yhydroxyalkanoate (PHA), polylactic acid (PLA) and thermoplastic starch (TPS). PHA: R=C1–C11. The presence of an ester bond (C–O)
provides a point of attack for hydrolytic enzymes, but not all plastics with an ester bond, e.g. PET, and PEF, are biodegradable. How-
ever hydrolytic enzymes such as cutinases are reported to degrade polymers such as PET [23, 26, 35–39] in vitro.

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Protection Agency(EPA) as the most preferable options
focus on the prevention and reduction of waste, followed by
reuse, recycling, energy recovery and disposal as the least
preferred option [14, 15]. An option that could be included
in these strategies is the management of biodegradable plas-
tics, as they can open up new end-of-life waste management
options that are not open to non-degradable plastics, such
as anaerobic digestion and composting. Food contamination
of current non-degradable plastics prevents or limits
options for recycling, whereas biodegradable plastics con-
taminated with food can be composted.
BIODEGRADATION OF OIL-BASED PLASTICS
The durability of plastic is the main obstacle to its degrada-
tion in the environment. However, some degradation does
occur. Degradation of oil-based plastic includes the com-
bined effect of biotic and abiotic factors [16]. Abiotic degra-
dation involves physical and chemical processes that cause
intramolecular changes in the polymer [17], including bond
cleavage, chemical transformation or the formation of new
chemical groups, and therefore change their mechanical,
optical and electrical properties [18]. The addition of photo-
or thermo-oxidizers to polymers can activate abiotic degra-
dation and is proposed to facilitate further degradation by
microbial metabolism [17].
Biodegradation is defined as the capacity of a micro-
organism or microbial consortium to use the polymer as a
sole source of carbon and energy (Fig. 2). While the biodeg-
radation of traditional plastics has been widely researched
[16–18], limited mineralization (conversion into biomass,
CO2, H2O or CH4) of oil-based plastics has been reported
[17]. In general, the biodegradation of oil-based plastics
requires the activity of several different micro-organisms
[18]. The extent of biodegradation is affected by polymer
characteristics and the micro-organisms present in certain
environment. Crystallinity, molecular weight, hydrophobic-
ity, the presence of functional groups, and the use of addi-
tives and plasticizers in the fabrication process affect the
biodegradability of polymers. For example, an increase in
crystallinity and molecular weight will negatively affect bio-
degradation. On the other hand, pre-treatments of polymer
such as irradiation increase biodegradation [17]. In general,
PET that has hydrolyzable bonds is more susceptible to bio-
degradation compared to PE, PS, PP and polyvinyl chloride
(PVC) [19–22]. The hydrolysis of PET by the fungi Fusar-
ium oxysporum LCH1 and Fusarium solani was demon-
strated via the release of terephthalic acid (TA), one of the
PET monomers [23]. However, it was not shown whether
TA supports the growth of these fungi. Similarly, a study by
Nimchua and co-workers showed that 22 out of 115 isolates
of microfungi had the capacity to modify PET films, but the
mineralization of the polymer was not shown [24]. Recently,
a bacterium, Ideonella sakaiensis, was isolated from a PET-
contaminated environmental sample and it was shown that
it not only degrades PET, but also assimilates the monomers
[25]. However, the PET material used in this study had a
very low crystallinity of 1.9 %, which does not reflect the
type of PET used for packaging, which has a much higher
crystallinity (30–40 %). The low crystallinity will make the
polymer much more susceptible to degradation. The main
challenges for the biodegradation of oil-based plastics by
micro-organisms are limited bioavailability and polymer
insolubility in water, which limits the access of the enzymes
involved in the breakage to the polymer [22].
The enzymes reported to be involved in the breakage of pol-
ymers are cutinases, laccases, lipases and enzymes involved
in lignin metabolism [17, 22, 26, 27]. Non-hydrolyzable pol-
ymers are significantly harder to break down compared to

Fig. 2. Degradation of plastics characterized by the presence of hydrolyzable bonds. The most frequently used packaging polymer
polyethylene terephthalate (PET) can be degraded by the activity of enzymes such as cutinases and lipases. However, there are no
examples of the complete mineralization of PET. Polymers such as bio-based polyhydroxyalkanoates (PHAs) and polylactic acid (PLA),
and oil-based polycaprolactone (PCL) can be completely mineralized by the activity of micro-organisms.

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hydrolyzable polymers. It is proposed that non-hydrolyzable
PE could be oxidized and then degraded by b-oxidation,
similarly to n-alkanes of 10–50 carbon atoms [27]. The
enzymes involved in this process are intracellular, and
therefore reduction in PE mass is required prior to the intra-
cellular oxidation of PE. There are a few examples of bacte-
rial and fungal laccases and peroxidases that have been
shown to reduce the molecular mass of PE [28–30]. In addi-
tion, the pre-treatment of PE, such as pyrolysis, can give rise
to n-alkanes that are metabolized by micro-organisms [31].
PS is another example of highly recalcitrant plastic. While a
study by Otake and colleagues reported that no PS biodegra-
dation was observed in soil in 32 years [32], another study
showed very limited degradation (1 %) in a 90-day period
[33]. It was recently reported that the microbiome present
in the gut of mealworms is necessary for the biodegradation
of PS and PE, as well as the mixture of these two polymers
[34]. A decrease in the residual, i.e. non-degraded, polymer
was observed simultaneously with the increase in gas frac-
tion, suggesting that the larger fraction of the ingested plas-
tic was mineralized and/or incorporated into mealworm
biomass [34].
The hydrolases normally involved in the biodegradation of
biological macromolecules such as proteins and cellulose
are also reported to attack hydrolyzable bonds in fossil-
based polymers such as the ester linkage in PET between
terephthalic acid and ethylene glycol [22]. Several hydrolytic
enzymes with PET-degrading capacity have been reported
in the literature, such as TfCut1 and TfCut2 from Thermobi-
fidus fusca KW3 [26], LC-cutinase isolated from leaf-branch
compost using a metagenomic approach [35], cutinase-like
enzyme from Saccharomonospora viridis AHK190 [36], HiC
from Thermomyces insolens [37] and lipase B from Candida
antarctica [38]. While these enzymes can hydrolyze PET,
their activity is low and the process temperatures are high
[39]. For example, HiC, designated as the most active fungal
hydrolase to date, shows nearly complete biodegradation of
low-crystallinity PET (7 %) in 96 h at 70  C [37]. Various
approaches have been used to increase the activity of these
hydrolyzes. The addition of Ca2+ to cutinase-like enzyme
induced large conformational changes and thereby stabiliza-
tion of the binding of the substrate and increased thermo-
stability [36]. Site-directed mutagenesis was used to alter the
Ca2+ binding site, which resulted in increased thermostabil-
ity and PET degradation capacity [40]. Furthermore, using
a combination of enzymes such as TfCut2 and LC-cutinase
showed enhanced PET hydrolysis [41].
A PETase originating from I. sakaiensis (IsPETase) was
recently characterized and the mechanism of PET degrada-
tion to monohydroxyethyl terephthalate (MHET) was pro-
posed [42]. PET degradation involves the secretion of
IsPETase from the bacterium and its attachment to the sur-
face of PET via a flat hydrophobic surface of the enzyme
containing the substrate-binding cleft [42]. This is followed
by PET hydrolysis resulting in two PET chains, one with a
TA terminus and the second with a 2-hydroxyethyl (HE)
132
terminus. These two chains are further digested to produce
MHET and TA [42]. IsPETase shares 52 % amino acid iden-
tity with TfCut2 and both enzymes have the same catalytic
triad consisting of Ser–His–Asp [42, 43], suggesting that the
PET degradation mechanism is similar. The conversion of
the IsPETase active site to a narrower, cutinase-like active
site via site-directed mutagenesis led to the creation of a
PETase variant carrying two amino acid substitutions
(S238F and W159H) that performs better than the IsPETase
with regard to reduction of PET crystallinity and product
release [43]. Molecular modelling revealed that this is likely
due to the closer positioning of the PET carbonyl group for
the attack by catalytically active Ser [43].
BIOPLASTICS AND BIODEGRADATION
Bio-based polymers (bioplastics) are partly or completely
derived from biomass. Bioplastics can contribute to reduc-
tions in GHG emissions as a result of the consumption of
CO2 during the growth of crops used to supply the sugars,
present in the biomass, for bioplastic manufacture. How-
ever, the bio-based origin does not mean that bio-based
plastic is also biodegradable. Biodegradability is a feature
related to the chemical nature of the polymer and the envi-
ronmental conditions where the plastic is located. One hun-
dred per cent bio-based polyethylene (bio-PE) and
bio-based polyethylene terephthalate (bio-PET) are not bio-
degradable [44, 45]. Their chemical structure does not differ
from oil-based PET and PE. Polyethylene-2,5-furandicar-
boxylate (PEF; Fig. 1) is bio-based, but, like PET, it is not
biodegradable. The enzyme IsPETase, which was shown to
hydrolyze PET, also hydrolyzes PEF in laboratory condi-
tions, but the degree of degradation is limited [43]. Biode-
gradable plastics such as thermoplastic starch (TPS),
polylactic acid (PLA) and polyhydroxyalkanoate (PHA) can
be derived from renewable resources and are thus bio-based
and biodegradable plastics, but there are other biodegrad-
ables derived from fossil carbon, e.g. polycaprolactone
(PCL) and polybutylene adipate terephthalate (PBAT) [45]
(Fig. 1). The low melting temperature of 60  C [46] and the
abundance of PCL-degrading organisms [47, 48] are pro-
posed as key features allowing for the biodegradation of
PCL.
Whether a material will completely biodegrade (i.e. be min-
eralized) in a certain environment depends on its crystallin-
ity, the presence of additives, the presence of adequate
micro-organisms, temperature, moisture and the pH of the
environment [49]. This suggests that a plastic might be bio-
degradable in one but not another environment. It is there-
fore of paramount importance to assess the behaviour of
different biodegradable plastics in a range of environments
and determine whether plastics show complete biodegrada-
tion and under what conditions.
One of the major bioplastics on the market, polylactic acid
(PLA) is produced through a combination of the fermenta-
tion of bio-based sugars (to produce lactic acid) and chemis-
try to convert the lactic acid or lactide to PLA. A major
 Narancic and O’Connor, Microbiology 2019;165:129–137
application area for PLA is packaging [50]. PLA is highly
crystalline, which gives it desirable properties for applica-
tions. However, higher temperatures are required to allow
biodegradation. While the micro-organisms and enzymes
involved in PLA degradation are known [51] (Fig. 2), non-
enzymatic hydrolysis of PLA is a major degradation route
[52]. We have recently analysed various bio-based and fos-
sil-based biodegradable plastics for biodegradation across
seven environments, and we showed that PLA reaches inter-
national standards for biodegradation in industrial com-
posting and anaerobic digestion conditions [53]. We have
observed a very slow rate of PLA degradation in soil, based
on which we estimate that it would take over three decades
for PLA to biodegrade in soil [53].
PLA’s mechanical properties can be improved by blending
with polymers such as starch, poly(propylene-carbonate),
methacrylate-butadiene-styrene, natural rubber, PS and
PHA, and by making PLA composites with different fibres,
etc. [54, 55]. The fate of the new composite created by
blending could be completely different from that of the neat
polymer. We have demonstrated that the addition of PCL to
PLA not only improves the material properties of the blend,
but also makes this material home-compostable [53]. The
home-compostability of PLA-based plastics is an exciting
prospect, as it would provide an option to avoid the collec-
tion and transportation of packaging material to an indus-
trial composting facility, thereby reducing the GHG
emissions associated with kerb-side collection and also offer
consumers more choice for the management of their waste.
Therefore, the smart design of plastics can open up new
possibilities for the post-consumer management of
bioplastics.
However, biodegradable plastics must be managed and can-
not been seen as an easy solution to the current plastic
pollution problems. The blend of PLA/PCL that is home-
compostable shows no biodegradation in soil- or water-
based environments, highlighting the importance of the
effect of the environment on the biodegradability of biode-
gradable plastics [53] and the need to manage our waste.
Thus, assessing individual bioplastics and their blends for
biodegradation in various environments is vital, as the
uncontrolled release of biodegradable plastics into environ-
ments already endangered by plastic pollution, such as
marine or freshwater, will present similar problems to those
caused by non-degradable polymers [53].
Another example of bio-based and biodegradable plastics is
polyhydroxyalkanoates (PHA). PHA is a family of bacterial
polyesters that are usually accumulated by bacteria in
response to stress [56]. They are grouped into short-chain-
length (scl) polymers, containing (R) 3-hydroxyalkanoic
acids with four or five carbon atoms, with polyhydroxybuty-
rate (PHB) as a typical example, and medium-chain-length
(mcl) polymers of (R) 3-hydroxyalkanoic acids containing
6–12 carbon atoms [57]. PHAs have higly diverse material
propeties due in part to the monomer composition. There
are over 150 different known PHA monomers reported in
133
the scientific literature. For example, PHB is highly crystal-
line and brittle [56]. A closely related copolymer, poly(3-
hydroxybutyrate-co-3-hydroxyvalerate) [P(3HB-co-3HV)],
is more flexible because of reduced crystallinity and is suit-
able for many commercial applications [58]. MclPHAs such
as polyhydroxyoctanoate (PHO) are of low crystallinity and
behave as a rubber-like material [56].
The key hydrolytic enzymes involved in the microbial deg-
radation of PHA are depolymerases [59] (Fig. 2). PHB bio-
degradation has been studied extensively [48, 60, 61].
Bioplastics mainly composed of PHB or co-polymers of
PHB can be degraded to completion in soil and sludges
[61–63]. In addition, PHB along with TPS is the only poly-
mer that has been shown to completely degrade in various
environments according to international standard biodegra-
dation tests [American Standard Testing Method (ASTM),
International Standards Organization (ISO) and European
Norms (EN)] [53]. PHO, a reference mclPHA, met the
industrial composting standards but showed poor biodegra-
dation in soil [53, 64], freshwater and sea water conditions
[53].
The time frame for biodegradation is critically important in
defining the suitability of a plastic for a particular end-of-
life management technology or its likely fate in the environ-
ment. If the micro-organisms and therefore key enzymes
involved in biodegradation are present, but the rate of deg-
radation is so low that it does not differ significantly from
non-degradable counterparts, then the limited biodegrad-
ability does not offer a benefit to the environment or the
management of biodegradable waste. According to currently
employed international standards (ISO and ASTM) the pro-
posed time frame for biodegradation in water environments
is 56 days and in soil up to 2 years [53]. In managed envi-
ronments, such as industrial composting, home composting
and anaerobic digestion the proposed time frame is
180 days, 1 year and 15 days, respectively [53].
The major focus of society and policy-makers for the end-
of-life management of biodegradable bioplastics has been
industrial composting [65, 66]. However, anaerobic diges-
tion (AD), also known as biogasification, a biotechnological
approach to renewable biomass-derived energy production,
may also be an end-of-life management option, but it has
been largely unexplored. In this naturally occurring process
micro-organisms break down biodegradable material in an
enclosed system, producing methane-rich biogas and a
nutrient-rich fertilizer known as ‘digestate’ [67]. Currently
AD is targeting the organic fraction of MSW [68]. The
majority of bioplastics we tested in our recent study are
degraded in AD [53]. Therefore, the biodegradable plastics
used for food packaging could become a part of an inte-
grated waste management strategy for the organic fraction
of MSW. However, the residence time of biodegradable
plastics in AD is longer than the current residence times of
industrial AD reactors and so the design of AD reactors
and/or engineering of biodegradable plastics to decrease res-
idence times should be investigated.
 Narancic and O’Connor, Microbiology 2019;165:129–137

Fig. 3. Making plastic circular. Plastics are currently produced in a linear economy of ‘take, make and dispose’, with limited recycling.
This must be turned into a circular production cycle where plastics are re-used, recycled and upcycled through their conversion to
valuable products such as biodegradable polymers [polyhydroxyalkanoates (PHAs)]. Introducing biodegradable plastics to this model
through the upcycling of waste would allow the return of carbon to nature in a managed, safe and sustainable way.

Biogas generated via AD can contribute to renewable energy
targets. It is projected that 80 % of total US electricity
demand could be supplied by renewable electricity technol-
ogies in 2050 [69] and EU countries have agreed on a
renewable energy target of at least 27 % of final energy con-
sumption in the EU by 2030 as part of the EU’s energy and
climate goals for 2030 [70]. In addition, AD technologies
and the downstream purification of the produced gases
have advanced significantly in recent years [71]. The adap-
tation of AD technology can produce fatty acids [72], which
are valuable chemicals with multiple markets [73]. Finally,
CO2, which is co-produced with methane by AD, is emerg-
ing as an exciting building block to make polymers and
chemicals [74–76] in a circular economy [77].
CIRCULAR ECONOMY
The circular economy focuses on the retention of value in
the material cycle through the re-use and recycling of a
materials such as plastic and preventing their uncontrolled
release into the environment [77, 78]. The circular economy
focuses on the recycling of plastics (Fig. 3).
A current gap in circular economy thinking with respect to
plastics is the role of biodegradable plastics in the circular
economy. Firstly, biodegradable plastics such as PLA can be
mechanically recycled but also converted by biological pro-
cesses where carbon can be returned to nature in a safe and
sustainable way, e.g. by composting, which is central to a
circular economy. We have also reported on the
combination of pyrolysis and microbiology to convert non-
degradable plastics into biodegradable plastics, offering an
unconventional route for non-degradable plastics to cross
over from the technical half of the circular economy to the
biological half [31, 79–81]. If improvements to enzyme
activity are made then one can envisage enzyme technolo-
gies entering the technical half (i.e. pyrolysis and depolyme-
rizing plastics) of the circular economy in the future [82].
The generated monomers could be used to make more bio-
degradable plastics, which would represent a completely
biological recycling of plastics in a circular economy. The
integration of hydrolytic enzymes into a microbial chassis
would result in a custom microbial platform that is capable
of converting plastic into biodegradable counterparts in a
single cell [83, 84].
CONCLUDING REMARKS
The answer to the plastic waste problem requires a major
shift in human behaviour, which, in combination with tech-
nological solutions, can address the plastic waste problem,
but the latter will be largely ineffective without the former.
The throw-away attitude of society must change, and we
should not look upon biodegradable plastics as a technologi-
cal solution that excuses us from our environmental respon-
sibility. The predicted long residence times of most
biodegradable plastics in nature clearly indicate that these
plastics must be managed by society and not released into
the environment. Biodegradable plastics thus do not offer
society a ‘continue to throw away’ solution, but rather they

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 Narancic and O’Connor, Microbiology 2019;165:129–137
can offer humankind new end-of-life management options
for plastic waste that provide value through bio-based fertil-
izer (compost), biogas (renewable energy) or chemicals such
as fatty acids.
Funding information
T. N. acknowledges support under EU Horizon 2020 research and
innovation programme 633962 for the project P4SB.
Conflicts of interest
The authors declare that there are no conflicts of interest.
References
1. Plastics Europe. Plastics - The facts 2017. 2017.
2. European Commission. Questions & answers: a European strat-
egy for plastics. Strasbourg 2018.
3. EPA US. Advancing sustainable materials management. Fact
Sheet 2016.
4. System Initiative on Environment and Natural Resource Security.
The New Plastics Economy: Catalysing action. World Economic
Forum 2016.
5. European Commission. A European strategy for plastics in a cir-
cular economy; 2018 Contract No. SWD 2018;16.
6. North EJ, Halden RU. Plastics and environmental health: the road
ahead. Rev Environ Health 2013;28:1–8.
7. Hanke G. Marine beach litter in Europe – Top items: Joint
Research Centre. European Commission 2016.
8. Rochman CM, Browne MA, Halpern BS, Hentschel BT, Hoh E et al.
Policy: Classify plastic waste as hazardous. Nature 2013;494:169–
171.
9. Rochman CM, Hoh E, Kurobe T, Teh SJ. Ingested plastic transfers
hazardous chemicals to fish and induces hepatic stress. Sci Rep
2013;3:3263.
10. Wilcox C, van Sebille E, Hardesty BD. Threat of plastic pollution to
seabirds is global, pervasive, and increasing. Proc Natl Acad Sci
USA 2015;112:11899–11904.
11. Sussarellu R, Huvet A, Lap egue S, Quillen V, Lelong C et al. Addi-
tive transcriptomic variation associated with reproductive traits
suggest local adaptation in a recently settled population of the
Pacific oyster, Crassostrea gigas. BMC Genomics 2015;16:808.
12. Sussarellu R, Suquet M, Thomas Y, Lambert C, Fabioux C et al.
Oyster reproduction is affected by exposure to polystyrene micro-
plastics. Proc Natl Acad Sci USA 2016;113:2430–2435.
13. de Souza Machado AA, Lau CW, Till J, Kloas W, Lehmann A et al.
Impacts of microplastics on the soil biophysical environment.
Environ Sci Technol 2018;52:9656–9665.
14. EC. GREEN PAPER: on a european strategy on plastic waste in the
environment. 2013.
15. Epa U, Year F. EPA Strategic Plan. 2014.
16. Wei R, Zimmermann W. Microbial enzymes for the recycling of
recalcitrant petroleum-based plastics: how far are we? Microb
Biotechnol 2017;10:1308–1322.
17. Sivan A. New perspectives in plastic biodegradation. Curr Opin
Biotechnol 2011;22:422–426.
18. Shah AA, Hasan F, Hameed A, Ahmed S. Biological degradation of
plastics: a comprehensive review. Biotechnol Adv 2008;26:246–
265.
19. Webb HK, Arnott J, Crawford RJ, Ivanova EP. Plastic degradation
and its environmental implications with special reference to poly
(ethylene terephthalate). Polymers-Basel 2013;5:1–18.
20. Zheng Y, Yanful EK, Bassi AS. A review of plastic waste biodegra-
dation. Crit Rev Biotechnol 2005;25:243–250.
21. Tokiwa Y, Calabia BP, Ugwu CU, Aiba S. Biodegradability of plas-
tics. Int J Mol Sci 2009;10:3722–3742.
135
22. Krueger MC, Harms H, Schlosser D. Prospects for microbiological
solutions to environmental pollution with plastics. Appl Microbiol
Biotechnol 2015;99:8857–8874.
23. Nimchua T, Punnapayak H, Zimmermann W. Comparison of the
hydrolysis of polyethylene terephthalate fibers by a hydrolase
from Fusarium oxysporum LCH I and Fusarium solani f. sp. pisi.
Biotechnol J 2007;2:361–364.
24. Nimchua T, Eveleigh DE, Sangwatanaroj U, Punnapayak H.
Screening of tropical fungi producing polyethylene terephthalate-
hydrolyzing enzyme for fabric modification. J Ind Microbiol
Biotechnol 2008;35:843–850.
25. Yoshida S, Hiraga K, Takehana T, Taniguchi I, Yamaji H et al. A
bacterium that degrades and assimilates poly(ethylene tere-
phthalate). Science 2016;351:1196–1199.
26. Wei R, Oeser T, Zimmermann W. Synthetic polyester-hydrolyzing
enzymes from thermophilic actinomycetes. Adv Appl Microbiol
2014;89:267–305.
27. Restrepo-Flórez J-M, Bassi A, Thompson MR. Microbial degrada-
tion and deterioration of polyethylene – A review. Int Biodeterior
Biodegradation 2014;88:83–90.
28. Ojha N, Pradhan N, Singh S, Barla A, Shrivastava A et al. Evalua-
tion of HDPE and LDPE degradation by fungus, implemented by
statistical optimization. Sci Rep 2017;7:39515.
29. Fujisawa M, Hirai H, Nishida T. Degradation of polyethylene and
Nylon-66 by the laccase-mediator system. J Polym Environ 2001;
9:103–108.
30. Santo M, Weitsman R, Sivan A. The role of the copper-binding
enzyme – laccase – in the biodegradation of polyethylene by the
actinomycete Rhodococcus ruber. Int Biodeterior Biodegradation
2013;84:204–210.
31. Guzik MW, Kenny ST, Duane GF, Casey E, Woods T et al. Conver-
sion of post consumer polyethylene to the biodegradable polymer
polyhydroxyalkanoate. Appl Microbiol Biotechnol 2014;98:4223–
4232.
32. Otake Y, Kobayashi T, Asabe H, Murakami N, Ono K. Biodegrada-
tion of low-density polyethylene, polystyrene, polyvinyl chloride,
and urea formaldehyde resin buried under soil for over 32 years.
J Appl Polym Sci 1995;56:1789–1796.
33. Kaplan DL, Hartenstein R, Sutter J. Biodegradation of polystyrene,
poly(metnyl methacrylate), and phenol formaldehyde. Appl Environ
Microbiol 1979;38:551–553.
34. Brandon AM, Gao SH, Tian R, Ning D, Yang SS et al. Biodegrada-
tion of polyethylene and plastic mixtures in mealworms (Larvae of
Tenebrio molitor) and effects on the gut microbiome. Environ Sci
Technol 2018;52:6526–6533.
35. Sulaiman S, Yamato S, Kanaya E, Kim JJ, Koga Y et al. Isolation
of a novel cutinase homolog with polyethylene terephthalate-
degrading activity from leaf-branch compost by using a metage-
nomic approach. Appl Environ Microbiol 2012;78:1556–1562.
36. Miyakawa T, Mizushima H, Ohtsuka J, Oda M, Kawai F et al.
Structural basis for the Ca(2+)-enhanced thermostability and
activity of PET-degrading cutinase-like enzyme from Saccharomo-
nospora viridis AHK190. Appl Microbiol Biotechnol 2015;99:4297–
4307.
37. Ronkvist Åsa M, Xie W, Lu W, Gross RA. Cutinase-catalyzed
hydrolysis of poly(ethylene terephthalate). Macromolecules 2009;
42:5128–5138.
38. Carniel A, Valoni Érika, Nicomedes J, Gomes Adac, Castro Amde.
Lipase from Candida antarctica (CALB) and cutinase from Humicola
insolens act synergistically for PET hydrolysis to terephthalic acid.
Process Biochem 2017;59:84–90.
39. Wei R, Zimmermann W. Biocatalysis as a green route for recy-
cling the recalcitrant plastic polyethylene terephthalate. Microb
Biotechnol 2017;10:1302–1307.
40. Then J, Wei R, Oeser T, Barth M, Belis ario-Ferrari MR et al. Ca2+
and Mg2+ binding site engineering increases the degradation of
polyethylene terephthalate films by polyester hydrolases from
Thermobifida fusca. Biotechnol J 2015;10:592–598.
 Narancic and O’Connor, Microbiology 2019;165:129–137
41. Barth M, Honak A, Oeser T, Wei R, Belisario-Ferrari MR et al.
A dual enzyme system composed of a polyester hydrolase and
a carboxylesterase enhances the biocatalytic degradation of
polyethylene terephthalate films. Biotechnol J 2016;11:1082–
1087.
42. Joo S, Cho IJ, Seo H, Son HF, Sagong HY et al. Structural insight
into molecular mechanism of poly(ethylene terephthalate) degra-
dation. Nat Commun 2018;9:382.
43. Austin HP, Allen MD, Donohoe BS, Rorrer NA, Kearns FL et al.
Characterization and engineering of a plastic-degrading aro-
matic polyesterase. Proc Natl Acad Sci USA 2018;115:E4350–
E4357.
44. Reddy MM, Vivekanandhan S, Misra M, Bhatia SK, Mohanty AK.
Biobased plastics and bionanocomposites: current status and
future opportunities. Prog Polym Sci 2013;38:1653–1689.
45. Bioplastics E. What are bioplastics? www.european-bioplastics.
org/bioplastics/. [accessed 2018].
46. Woodruff MA, Hutmacher DW. The return of a forgotten polymer
—Polycaprolactone in the 21st century. Prog Polym Sci 2010;35:
1217–1256.
47. Nishida H, Tokiwa Y. Distribution of poly(b-hydroxybutyrate) and
poly("-caprolactone)aerobic degrading microorganisms in differ-
ent environments. J Environ Polym Degrad 1993;1:227–233.
48. Suyama T, Tokiwa Y, Ouichanpagdee P, Kanagawa T, Kamagata
Y. Phylogenetic affiliation of soil bacteria that degrade aliphatic
polyesters available commercially as biodegradable plastics. Appl
Environ Microbiol 1998;64:5008–5011.
49. Mohee R, Unmar G. Determining biodegradability of plastic mate-
rials under controlled and natural composting environments.
Waste Manag 2007;27:1486–1493.
50. European Bioplastics. Applications for bioplastics. 2018. www.
european-bioplastics.org/market/applications-sectors/.
51. Kawai F. Polylactic Acid (PLA)-Degrading Microorganisms and
PLA Depolymerases. In: Cheng HN and Gross RA (editors). Green
Polymer Chemistry: Biocatalysis and Biomaterials American Chemi-
cal Society. pp. 405–414.
52. Tsuji H. Polylactides. In: Steinbüchel A and Doi Y (editors).
Biopolymers: Polyesters III. Weinheim, Germany: Wiley-VCH; 2002.
pp. 129– 177.
53. Narancic T, Verstichel S, Reddy Chaganti S, Morales-Gamez L,
Kenny ST et al. Biodegradable plastic blends create new possibili-
ties for end-of-life management of plastics but they are not a
panacea for plastic pollution. Environ Sci Technol 2018;52:10441–
10452.
54. Sangeetha VH, Deka H, Varghese TO, Nayak SK. State of the art
and future prospectives of poly(lactic acid) based blends and com-
posites. Polym Compos 2018;39:81–101.
55. Chen CC, Chueh JY, Tseng H, Huang HM, Lee SY. Preparation and
characterization of biodegradable PLA polymeric blends.
Biomaterials 2003;24:1167–1173.
56. Rehm BH. Bacterial polymers: biosynthesis, modifications and
applications. Nat Rev Microbiol 2010;8:578–592.
57. Sudesh K, Abe H, Doi Y. Synthesis, structure and properties of
polyhydroxyalkanoates: biological polyesters. Prog Polym Sci
2000;25:1503–1555.
58. Kunioka M, Tamaki A, Doi Y. Crystalline and thermal properties of
bacterial copolyesters: poly(3-hydroxybutyrate-co-3-hydroxyvaler-
ate) and poly(3-hydroxybutyrate-co-4-hydroxybutyrate).
Macromolecules 1989;22:694–697.
59. Jendrossek D, Handrick R. Microbial degradation of polyhydrox-
yalkanoates. Annu Rev Microbiol 2002;56:403–432.
60. Verma SL, Marschner P. Compost effects on microbial biomass
and soil P pools as affected by particle size and soil properties. J
Soil Sci Plant Nut 2013;13:313–328.
61. Volova TG, Prudnikova SV, Vinogradova ON, Syrvacheva DA,
Shishatskaya EI. Microbial Degradation of Polyhydroxyalkanoates
136
with Different Chemical Compositions and Their Biodegradability.
Microb Ecol 2017;73:353–367.
62. Sridewi N, Bhubalan K, Sudesh K. Degradation of commercially
important polyhydroxyalkanoates in tropical mangrove ecosystem.
Polym Degrad Stab 2006;91:2931–2940.
63. Wang Y-W, Mo W, Yao H, Wu Q, Chen J et al. Biodegradation stud-
ies of poly(3-hydroxybutyrate-co-3-hydroxyhexanoate). Polym
Degrad Stab 2004;85:815–821.
64. Lim SP, Gan SN, Tan IK. Degradation of medium-chain-length pol-
yhydroxyalkanoates in tropical forest and mangrove soils. Appl
Biochem Biotechnol 2005;126:23–33.
65. McCarthy BEC. Developments in the end-of-life management of
plastics. The International Institute for Industrial Environmental
Economics IIIEE Lund 2005.
66. European Bioplastics. Driving the evolution of plastics. 2016.
67. Santi G, Proietti S, Moscatello S, Stefanoni W, Battistelli A. Anaer-
obic digestion of corn silage on a commercial scale: Differential
utilization of its chemical constituents and characterization of the
solid digestate. Biomass and Bioenergy 2015;83:17–22.
68. Gerber van Doren L, Posmanik R, Bicalho FA, Tester JW, Sills
DL. Prospects for energy recovery during hydrothermal and
biological processing of waste biomass. Bioresour Technol 2017;
225:67–74.
69. Bazilian M, Mai T, Baldwin S, Arent D, Miller M et al. Decision-
making for High Renewable Electricity Futures in the United
States. Energy Strategy Reviews 2014;2:326–328.
70. EC. Renewable energy progress report. 2017.
71. Zhao Q, Leonhardt E, MacConnell C, Frear C, Chen S. Purification
Technologies for Biogas Generated by Anaerobic Digestion. Center
for Sustaining Agriculture and Natural Resources 2010.
72. Khan MA, Ngo HH, Guo WS, Liu Y, Nghiem LD et al. Optimization
of process parameters for production of volatile fatty acid, biohy-
drogen and methane from anaerobic digestion. Bioresour Technol
2016;219:738–748.
73. IndustryARC. Fatty Acid Market: By Bond (Unsaturated Fatty Acid.
and Saturated Fatty Acid), By Length of Chain (Short-chain fatty
acids (SCFA), Medium-chain fatty acids (MCFA), Long-chain fatty
acids (LCFA) and Very long chain fatty acids (VLCFA)) & By
Region-Forecast. 2016:Contract No.: CMR 0377.
74. Hu B, Guild C, Suib SL. Thermal, electrochemical, and photochem-
ical conversion of CO2 to fuels and value-added products. Journal
of CO2 Utilization 2013;1:18–27.
75. Revelles O, Beneroso D, Men endez JA, Arenillas A, García JL
et al. Syngas obtained by microwave pyrolysis of household
wastes as feedstock for polyhydroxyalkanoate production in Rho-
dospirillum rubrum. Microb Biotechnol 2017;10.
76. LanzaTech. www.lanzatech.com/innovation/technical-overview/.
2017.
77. World Economic Forum. Ellen MacArthurFoundation and McKin-
sey & Company. The New Plastics Economy — Rethinking the
Futureof Plastics; 2016. www.ellenmacarthurfoundation.org/
publications.
78. The Ellen MacArthur Foundation. Circular Economy System Dia-
gram. [accessed 2018]: https://www.ellenmacarthurfoundation.
org/circular-economy/interactive-diagram.
79. Kenny ST, Runic JN, Kaminsky W, Woods T, Babu RP et al. Devel-
opment of a bioprocess to convert PET derived terephthalic acid
and biodiesel derived glycerol to medium chain length polyhydrox-
yalkanoate. Appl Microbiol Biotechnol 2012;95:623–633.
80. Kenny ST, Runic JN, Kaminsky W, Woods T, Babu RP et al. Up-
cycling of PET (polyethylene terephthalate) to the biodegradable
plastic PHA (polyhydroxyalkanoate). Environ Sci Technol 2008;42:
7696–7701.
81. Goff M, Ward PG, O’Connor KE. Improvement of the conversion of
polystyrene to polyhydroxyalkanoate through the manipulation of
the microbial aspect of the process: a nitrogen feeding strategy
 Narancic and O’Connor, Microbiology 2019;165:129–137

for bacterial cells in a stirred tank reactor. J Biotechnol 2007;132:
283–286.
82. Franden MA, Jayakody LN, Li WJ, Wagner NJ, Cleveland NS et al.
Engineering Pseudomonas putida KT2440 for efficient ethylene
glycol utilization. Metab Eng 2018;48:197–207.
83. From plastic waste to plastic value using Pseudomonas putida
synthetic biology. www.p4sb.eu.
84. Wierckx N, Prieto MA, Pomposiello P, de Lorenzo V, O’Connor K
et al. Plastic waste as a novel substrate for industrial biotechnol-
ogy. Microb Biotechnol 2015;8:900–903.

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