Environ Sci Pollut Res (2013) 20:4339–4355
DOI 10.1007/s11356-013-1706-x
REVIEW ARTICLE
Study of microbes having potentiality for biodegradation
of plastics
Swapan Kumar Ghosh & Sujoy Pal & Sumanta Ray
Received: 1 December 2012 / Accepted: 1 April 2013 / Published online: 24 April 2013
# Springer-Verlag Berlin Heidelberg 2013
Abstract Plastic is a broad name given to the different
types of organic polymers having high molecular weight
and is commonly derived from different petrochemicals.
Plastics are generally not biodegradable or few are degrad-
able but in a very slow rate. Day by day, the global demand of
these polymers is sharply increasing; however, considering
their abundance and potentiality in causing different environ-
mental hazards, there is a great concern in the possible
methods of degradation of plastics. Recently, there have been
some debates at the world stage about the potential degrada-
tion procedures of these synthetic polymers and microbial
degradation has emerged as one of the potential alternative
ways of degradation of plastics. Alternatively, some scientists
have also reported many adverse effects of these polymers in
human health, and thus, there is an immediate need of a
potential screening of some potential microbes to degrade
these synthetic polymers. In this review, we have taken an
attempt to accumulate all information regarding the chemical
nature along with some potential microbes and their enzymat-
ic nature of biodegradation of plastics along with some key
factors that affect their biodegradability.
Keywords Plastics . Environment . Pollution . Microbes .
Biodegradation . Enzymes
Introduction
Plastic is one of the most vital man-made product that has
been produced in huge quantity and is used widely for
Responsible editor: Philippe Garrigues
S. K. Ghosh (*) : S. Pal : S. Ray
Mycopathology Laboratory, Department of Botany, Ramakrishna
Mission Vivekananda Centenary College, P.O. Rahara,
Kolkata 700118 West Bengal, India
e-mail: swapan.krghosh@yahoo.com
different purposes in our everyday life, and gradually, the
global demand of this synthetic product is rapidly growing
day by day. The word plastic is derived from the Greek
work “plastikos” meaning the substance which can be
molded into any shape (Joel 1995). Chemically, plastic is a
long hydrocarbon chain polymer having high molecular
weight. Plastics are mainly derived from petrochemicals
which are further synthetically arranged by some chemical
processes to produce these long chain polymers (Shimao
2001). Now, plastics are mainly categorized into two
groups: one group is nonbiodegradable and another is bio-
degradable. Question may arise why nonbiodegradable plas-
tics are not degradable and why biodegradables are
degradables. Basically, plastics are usually long chains of
carbon and hydrogen atoms. The enzymes found in living
things can perform many chemical reactions, but they gen-
erally exploit some sort of imbalance of electric charge
within a molecule to do their job. A long chain of carbons
and hydrogens contains very balanced charges along its
length, making the molecule stable and difficult to change
with enzymes. Most biodegradable substances contain some
mixture of carbon and atoms like oxygen, nitrogen, sulfur,
and phosphorus, which create charge imbalances that en-
zymes can exploit. There are some bacteria that can break
down plastics. These bacteria usually contain enzymes
called oxygenases, which can add oxygen to a long carbon
chain. This destabilizes the local electric charge, and the
plastic can then be broken down. The oxygenase enzymes
too often are not found, however, because they can easily
destroy the molecules in the bacteria that carry them.
In India, plastic consumption grew exponentially in the
1990s. During the last decade, the total consumption of
plastics grew twice as fast as 12 % per annum (Shimao
2001) as the gross domestic product growth rate based on
purchasing power parities is 6 % per annum. The current
growth rate in Indian polymer consumption which is 16 %
per annum is clearly higher than that in China which has
10 % per annum and many other key Asian countries. The
4340
average Indian consumption of virgin plastics per capita
reached 3.2 kg in 2000/2001 (5 kg if recycled material is
included) from a mere 0.8 kg in 1990/1991. However, this is
only one-fourth of the consumption as compared to that of
China (12 kg/capita, 1998) and one-sixth of the world
average (18 kg/capita). This consumption led to more than
5,400 tonnes of plastic waste being generated per day in
2000/2001 (totaling about two million tonnes per annum)
(en.wikipedia.org).
The Middle East countries alone produce about 35 million
tons of polyethylene and polypropylene, and the capacity of
production of this man-made product is about to cross the 7
million tons per annum mark (data obtained from
CIEPT/Government of India). Asia alone is the world’s largest
plastic consumers, accounting about 35 % of the global con-
sumption which is followed by North America with about
26 % and Western Europe about 23 %. India (5 %) along with
Japan (6 %) comprises 11 % of the world plastic consumption
and about 40 % of the total consumption in Asia (European
Commission DG ENV 2011). According to the U.S. Environ-
mental Protection Agency (www.epa.gov), about 31 million
tons of plastic waste was produced in 2010, which represents
about 12.4 % of the total Mean Solid Waste, and out of that,
only 8 % waste was recycled. In contrast to that, to meet the
increasing demand of plastic, the production of these poly-
mers is rapidly increasing, and consequently, this man-made
chemical is causing high environmental pollution and also
notable human health hazards. Thus, almost every country in
this world is suffering from these problems.
India has witnessed a substantial growth in the consump-
tion of plastics and an increased production of plastic waste.
Polyolefins account for the major share of 60 % in the total
plastic consumption in India. Packaging is the major plastic-
consuming sector, with 42 % of the total consumption,
followed by consumer products and the construction indus-
try. The relationship observed between plastic consumption
and the gross domestic product for several countries was
used to estimate future plastic consumption (master curve).
Elasticities of the individual material growth with respect to
GDP were established for the past and for the next three
decades estimated for India, thereby assuming a develop-
ment comparable with that of Western Europe. On this basis,
the total plastic consumption is projected to grow by a factor
of 6 between 2000 and 2030. The consumption of various
end products is combined with their corresponding lifetimes
to calculate the total waste quantities. The weighted average
lifetime of plastic products was calculated as 8 years. Of the
total plastic waste generated, 47 % is currently recycled in
India; this is much higher than the share of recycling in most
of the other countries. The recycling sector alone employs as
many people as the plastic-processing sector, which em-
ploys about eight times more people than the plastic-
manufacturing sector. Due to the increasing share of long-
Environ Sci Pollut Res (2013) 20:4339–4355
life products in the economy, and consequently in the vol-
ume of waste generated, the share of recycling will decrease
to 35 % over the next three decades. The total waste avail-
able for disposal (excluding recycling) will increase at least
10-fold up to the year 2030 from its current level of 1.3
million tonnes (Wikipedia.org).
The increasing quantities of plastic waste and their effec-
tive and safe disposal have become a matter of public concern.
The increasingly visible consequences of indiscriminate
littering of plastic wastes (in particular, plastic packaging
wastes and discarded bags) have stimulated public outcry
and shaped policy. Littering also results in secondary prob-
lems such as drains becoming clogged and animal health
problems (both domesticated and wild). As a consequence,
many big cities, e.g., Kolkata, Mumbai, Bangalore, and Delhi,
are facing big threat of solid waste accumulation. Along with
that, it is becoming a serious threat as a soil and air pollutant
for if heated/burned, they produce some high amount of toxic
noxious gas (also some greenhouse gases) which are danger-
ous to human health, thus leading to severe air pollution.
When plastics are dumped in a field or in dumping areas,
there is evidential proof that they are causing a great change in
the pH of the soil followed by disturbance in the leaching of
the rain water and moisture, making the land bare and
unfertile. The biological degradation time of is very high,
and it takes thousands of years to degrade these long chain
polymers into simple hydrocarbons. Latest reports confirmed
that some plastic products are mimicking human hormones
(e.g., thyroxin and sex hormones), causing human health
hazards (Soto et al. 1991; Hao et al. 2011). It is also creating
a major problem in marine ecosystem.
For the last 30 years, scientists are trying to develop some
alternative ways other than the natural destruction to de-
grade these high molecular synthetic polymers, but yet now,
very few evidences are available where scientists are able to
develop some alternative ways to enhance the mode of
degradation and make it faster. Recent research suggests
that there have been a notable number of microorganisms
(especially some bacteria and fungi) which have the capac-
ity to degrade these synthetic polymers in much faster way
in comparison to the natural method by using some exo-
enzymes under stress conditions. The enzyme lipases from
Rhizopus arrhizus, Rhizopus delemar, Achromobacter sp.,
and Candida cylindracea and esterase from hog liver
showed activities on polyethylene adipate (PEA) and
poly(ε-caprolactone) (PCL) (Tokiwa and Suzuki 1977a, b).
Enzymatic degradation of PCL by Aspergillus flavus and
Penicillium funiculosum showed that faster degradation was
observed in the amorphous region (Cook et al. 1981). More-
over, a novel poly(3-hydroxybutyrate) (PHB) depolymerase
from a thermophilic Streptomyces sp. was also capable of
degrading poly(β-propiolactone) (PPL) (Calabia and
Tokiwa 2006). In comparative studies, the biodegradability
Environ Sci Pollut Res (2013) 20:4339–4355
of three polyalkylene succinate [polyethylene succinate
(PES), polybutylene succinate (PBS), and polypropylene
succinate (PPS)] with the same molecular weight was in-
vestigated using Rhizopus delemar lipase. PPS with low
melting temperature (Tm, 43–52 °C) had the highest biodeg-
radation rate followed by PES, owing to the lower crystallinity
of PPS compared to PES and PBS (Bikiaris et al. 2006).
The objectives of this review literature are to study the
nature and classification of plastic and its effect on the
ecosystem including the human health and to categorize
the potential microbes and their mechanisms of biodegrada-
tion of different plastic products. This work will provide a
ready reference for further scientific exploration of these
potential microbes.
Categories and classification of plastics
Chemically, there are various types of plastics which are
classified according to their chemical structure and their
properties.
Classification of plastic according to their thermal properties
Based on their thermal properties, plastics are classified into
two groups: thermoplastics and thermosetting polymers.
Thermoplastics
Thermoplastics are those types of plastics which cannot
undergo chemical changes in their composition when heat-
ed, and thus, they can undergo molding for several times.
Polyethylene (PE), polypropylene (PP), polystyrene (PS),
polyvinyl chloride (PVC), and polytetrafluoroethylene are
such examples. They are also known as common plastics
which range from 20,000 to 500,000 amu in molecular
weight. They have different numbers of repeating unit derived
from a simple monomer unit.
Thermosetting polymers
Thermosetting polymers are other types of plastics, which
when melt and been casted into a particular shape, remain
solid and after that they cannot be melt and modified again.
In the thermosetting polymers, the chemical change is irre-
versible and hence they are not recyclable too. Examples
include phenol–formaldehyde, polyurethanes, etc.
Classification of plastic according to their designing
properties
The other way of classification is based upon their relevance
of manufacturing process and designing. It is classified in
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different parameters like electrical conductivity, durability,
tensile strength, degradability, and thermal stability.
Classification of plastic according to their degradability
properties
The chemical properties of the plastics are also important
criteria for differentiating them into degradable and
nondegradable polymers. Usually, nonbiodegradable plas-
tics are known as synthetic plastics and they are derived
from petrochemicals. They have an unusual repeat of small
monomer units and thus have very high molecular weight.
In comparison, degradable plastics are made of starch and
thus are not of very high molecular weight. They usually
break down in interaction with UV, water, enzymes, and
gradual changes in pH. Biopol is an expensive biodegrad-
able plastic (which comprises polyhydroxybutyrate) avail-
able in market. Ecoflex, which is fully biodegradable, is
another biodegradable plastic polymer which is produced
by the German company BASF.
Types of plastics we use every day
Plastics are relatively very low cost, durable, and very easy
to manufacture. In Table 1, we have tried to briefly describe
some of the regular usable plastics and their application.
Effect of plastic on the environment and human health
With the rapid modernization and development of human
society, there has been a very sharp increase in the demand
of plastic-based products. Approximately 30–40 % of plas-
tics produced worldwide are used for different packaging
applications, and it is steeply increasing at the rate of about
12 % per annum (Sabir 2004). The reason behind such an
increase in the demand of plastic products lies in two im-
portant things: the durability and cost. For example, if we
compare a wooden chair with a plastic chair, the latter is
rather very cheap and highly durable. In this way, all plastic-
based products have very rapidly replaced almost every
natural product in the market, but due to their massive
production worldwide, there has been a sharp increase in
the generation of nondegradable solid plastic wastes. Thus,
slowly within the last 50 years, this has become one of the
great concerns for the modern human society. Many studies
also showed the striking effect of plastic waste on the
aquatic and marine ecosystem, and thus, it has become one
of the major problems for the modern environmentalist. To
get rid of such a menace, people usually put them in landfills
or burn it, but both these practices cause very serious threats
to the environment and the ecosystem. Burning plastics
usually produce some noxious gases like furans and dioxins
4342 Environ Sci Pollut Res (2013) 20:4339–4355

Table 1 List of various types of plastic we use everyday

Types Use/application

Polyester (PES) Fibers, textiles

Polyethylene terephthalate (PET)
Polyethylene (PE)
High-density polyethylene (HDPE)
Polyvinyl chloride (PVC)
Polyvinylidene chloride (PVDC)
(Saran)
Low-density polyethylene (LDPE)
Polypropylene (PP)
Polystyrene (PS)
High impact polystyrene (HIPS)
Polyamides (PA) (nylons)
Acrylonitrile butadiene styrene (ABS)
Polycarbonate (PC)
Polycarbonate/acrylonitrile butadiene
styrene (PC/ABS)
Polyurethanes (PU)
Melamine formaldehyde (MF)
Plastarch material
Phenolics (PF) or (phenol
formaldehydes)
Polyetheretherketone (PEEK)
Polyetherimide (PEI) (Ultem)
Polylactic acid (PLA)
Polymethyl methacrylate (PMMA)
Polytetrafluoroethylene (PTFE)
Urea–formaldehyde (UF)
Carbonated drinks bottles, peanut butter jars, plastic film, microwavable packaging
Wide range of inexpensive uses including supermarket bags, plastic bottles
Detergent bottles and milk jugs
Plumbing pipes and guttering, shower curtains, window frames, flooring
Food packaging
Outdoor furniture, siding, floor tiles, shower curtains, clamshell packaging
Bottle caps, drinking straws, yogurt containers, appliances, car fenders (bumpers), plastic pressure pipe
systems
Packaging foam, food containers, plastic tableware, disposable cups, plates, cutlery, CD and cassette
boxes
Refrigerator liners, food packaging, vending cups
Fibers, toothbrush bristles, fishing line, under-the-hood car engine moldings
Electronic equipment cases (e.g., computer monitors, printers, keyboards), drainage pipe
Compact discs, eyeglasses, riot shields, security windows, traffic lights, lenses
A blend of PC and ABS that creates a stronger plastic. Used in car interior and exterior parts and mobile
phone bodies
Cushioning foams, thermal insulation foams, surface coatings, printing rollers (currently the sixth or
seventh most commonly used plastic material, for instance, the most commonly used plastic found in
cars)
One of the aminoplasts and used as a multicolorable alternative to phenolics, for instance, in moldings
(e.g., break-resistance alternatives to ceramic cups, plates, and bowls for children) and the decorated
top surface layer of the paper laminates (e.g., Formica)
Biodegradable and heat-resistant thermoplastic composed of modified cornstarch
High-modulus, relatively heat-resistant, and excellent fire-resistant polymer; used for insulating parts in
electrical fixtures, paper laminated products (e.g., Formica), and thermally insulation foams. It is a
thermosetting plastic, with the familiar trade name Bakelite that can be molded by heat and pressure
when mixed with filler-like wood flour or can be cast in its unfilled liquid form or cast as foam (e.g.,
Oasis). Problems include the probability of moldings naturally being dark colors (red, green, brown),
and as thermoset, it is difficult to recycle
Strong, chemical, and heat-resistant thermoplastic; biocompatibility allows for use in implant
applications and aerospace moldings. One of the most expensive commercial polymers
A high-temperature, chemically stable polymer that does not crystallize
A biodegradable thermoplastic that can be converted into a variety of aliphatic polyesters derived from
lactic acid which in turn can be made by fermentation of various agricultural products such as
cornstarch, once made from dairy products
Contact lenses, glazing (best known in this form by its various trade names around the world, e.g.,
Perspex, Oroglas, PLEXIGLAS), aglets, fluorescent light diffusers, rear light covers for vehicles. It
forms the basis of artistic and commercial acrylic when suspended in water with the use of other
agents
Heat-resistant, low-friction coatings, and used in things like nonstick surfaces for frying pans, plumber’s
tape, and water slides. It is more commonly known as Teflon
One of the aminoplasts and used as a multicolorable alternative to phenolics. Used as a wood adhesive
(for plywood, chipboard, hardboard) and electrical switch housings

which are some dangerous greenhouse gases and play an
important role in ozone layer depletion. In fact, dioxins
cause serious problems in the human endocrine hormone
activity, thus becoming a major concern for the human
health too (NoPE 2002; Pilz et al. 2010). Dioxins also cause
very serious soil pollution, causing a great concern for the
scientific community worldwide.
Some plastics which are produced for packing foods are
made with the assistance of a substance called bisphenol A
(BPA), which is a synthetic chemical compound
[(CH3)2C(C6H4OH)2]. This plastic has an excellent property
of heat stability, making it tremendously useful in many
applications. Studies show that BPA can interfere with the
regulation of both development and reproduction, through its
Environ Sci Pollut Res (2013) 20:4339–4355
interaction with estrogen. In 1993, some scientific groups
reported on the interference estrogenic activity of BPA that
was released from polycarbonate flasks during the autoclaving
of media. There was also evidence that BPA could act on
MCF-7 human breast cancer cells as an estrogen, stimulating
cellular proliferation and inducing progesterone receptors.
BPA could bind to estrogen receptors, and the estrogenic
effects induced by BPA were blocked by the estrogen antag-
onist tamoxifen, thus supporting the notion that the estrogenic
activity of BPA was mediated via the estrogen receptor
(Krishnan et al. 1993). Soto and colleagues (1991) at Tufts
University showed very similar results with another compo-
nent of plastics, p-nonylphenol. This alkylphenolic substance,
which is used as a plastic additive and surfactant, could be
released from polyvinyl chloride and polystyrene plastics
even without autoclaving. Soto et al. (1995) also developed
an “E-SCREEN” to assay the estrogenic activity of unknown
substances or mixtures based upon their ability to stimulate
MCF-7 cell proliferation. A link between chemicals called
phthalates and thyroid hormone levels was confirmed by the
University of Michigan in the first large-scale and nationally
representative study of phthalates and BPA in relation to
thyroid function in humans (Science Daily: July 11, 2011). It
has also been noted that long exposure to BPA shows a
significant effect on the sex hormones (progesterone) in fe-
males (Hao et al. 2011).
BPA is now declared a controversial plastic product because
it exerts weak, but detectable, hormone-like properties, raising
great concerns about its presence in consumer products and
foods contained in such products. In 2008, several govern-
ments questioned about its safety, prompting some retailers
to withdraw polycarbonate products. In 2010, reports from the
U.S. Food and Drug Administration raised further concerns
regarding exposure of fetuses, infants, and young children
(U.S. FDA 2010). In September 2010, Canada became the
first country to declare BPA as a toxic substance (Mittelstaedt
2010; Canada Gazette Part II 2010). In the European Union
and Canada, BPA use is banned in baby bottles. There has not
been any such major report on other types of plastics, but there
is a great potentiality that these polymers may have some other
types of effects on the human health which is yet to be
discovered.
There is an evidential proof that every year, thousands of
aquatic and marine life suffer due to dumping of solid
plastic wastes in the sea and river. According to the World
Wide Fund for Nature (formally known as the World Wild-
life Fund) organization, it is estimated that about 100,000
marine mammals are killed due to plastic debris around the
world. It occurs because the floating plastic debris is often
mistaken as food, causing blockage in the digestive system
of the animals. Recently, in August 2000, an autopsy of a
dead Bryde’s whale near Cairns, Australia revealed that the
stomach of the whale was filled with plastic rubbishes which
4343
are thrown in the sea (WWF-Australia 2010). Sometimes, the
endocrine- and sex hormone-disrupter chemicals follow the
food chain (as sea food and sea fish is consumed in very high
amount), thus showing an indirect effect on human health.
Potential degradation procedures of plastics
Degradation in broad terms is defined as any process which
results into breaking of large complex molecules into smaller
molecules. Any type of physical or chemical changes can result
into degradation of these long polymers into their monomers.
There are several ways implemented in degrading the plastic.
Thermal degradation
It is a process where heat is used to degrade the plastic
polymers into their simpler form. It is also known as a
thermo-oxidative reaction as it requires oxygen. In this
process, the long polymers are broken into smaller mono-
meric units (radicals) which further react with oxygen to
produce peroxide radicals.
Photoreactive degradation
It is another type of process where high-intensity photon
particles are used to degrade the large polymers into smaller
ones. In this process, high-energy radiations like UV are
used to react with the photoreactive groups to break these
long chain polymers.
Biochemical/microbial degradation (biodegradation)
It is a new concept in which some live microorganisms are
used, which produce some biologically active enzymes to
degrade the long polymers, and further, these microorgan-
isms use these polymers as carbon and energy sources.
What modern scientists are thinking about degradability
of plastic polymers?
There is an evidential proof that the current techniques of
plastic polymer degradation are not so much effective;
hence, scientists are looking for some alternative ways
where they can use microbes to degrade these long chain
synthetic polymers into their respective monomers. This
process may also be termed as reverse flow to produce
simple hydrocarbons because these polymers are usually
produced from different petrochemical products. Thus, we
can be able to go back to those simple monomers which can
be an alternative source of energy and may even be the next
generation fuel. Alternatively, the demand for petrochemical
4344
products will surely be reduced as we can again recycle
these monomers to reproduce these polymers. Hence, there
is a huge demand in exploring these microbes which can
grow in different conditions and, under specific stress con-
dition, may be directed to grow and use those carbon poly-
mers as their energy source, thus degrading these synthetic
polymers. As our world is a great natural source of these
varied types of microbes, scientists are trying to explore and
utilize them for such activities. Some scientists are now
engaged in research in these directions, and an attempt has
been made to focus their works in this review article.
Some of the well-known microbes which have been
utilized and have the capacity to degrade plastic polymers
into their respective simple monomeric units are shown in
Table 2.
The information in Table 2 showed that many fungi are
capable of degrading various categories of plastic. PHB and
polyesters are degraded by many fungi genera such as
Acremonium, Cladosporium, Debaryomyces, Emericellopsis,
Eupenicillium, Fusarium, Mucor, Paecilomyces, Penicillium,
Pullularia, Rhodosporidium, and Verticillium. Similarly, PCL
is degraded by Aspergillus, Aureobasidium, Chaetomium,
Cryptococcus, Fusarium, Rhizopus, Penicillium, and
T he r m oa s c us. P EA is de gr ade d b y A s p e rg i l l u s,
Aureobasidium, Penicillium, Pullularia. On the other hand,
polylactic acid (PLA) is subjected to degradation by only two
genera of fungi (Penicillium roqueforti and Tritirachium
album). Mogil'nitskii et al. (1987) reported that Aspergillus
niger van Tieghem F-1119 had the ability to degrade PVC
plastic. Table 2 also indicated that there is a maximum
number of plastic-degrading fungal genera belonging to
Deuteromycota, whereas there is only a minimum number of
genera belonging to Zygomycota (Mucor sp. and Rhizopus sp.).
Different kinds of plastics and microbes
which can degrade these plastics
Aliphatic polyesters from fossil resources
Polyethylene adipate (PEA)
PEA ([−OCH2CH2OOC(CH2)4CO–]n) is a prepolymer of
polyurethane. PEA-degrading microorganisms were
screened and isolated using PEA (number average molecu-
lar weight (Mn) 3,000) as the sole source of carbon. Among
the isolated PEA-degrading microorganisms, Penicillium sp.
strain 14–3 exhibited the strongest activity. PEA was de-
graded in 120 h at high cell concentrations. This strain can
degrade not only PEA but also aliphatic polyesters such as
PES, PBS, and polybutylene adipate (PBA) (Tokiwa and
Suzuki 1974). The enzyme responsible for the degradation
of PEA has been purified and is considered to be a kind of
Environ Sci Pollut Res (2013) 20:4339–4355
lipase with broad substrate specificity. The purified enzyme
has a molecular weight of 25 kDa and could degrade various
kinds of aliphatic polyesters, such as PPL and PCL, but
not poly( D L -3-methylpropiolactone) or poly( D L -3-
hydroxybutyrate) (Tokiwa and Suzuki 1977a, b). This enzyme
can also hydrolyze plant oils, triglycerides, and methyl esters
of fatty acids. Given that the purified enzyme of Penicillium
sp. strain 14–3 has properties that are similar to lipase, some
commercially available lipases and esterases were used to
confirm if it was capable of degrading PEA. Results showed
that lipases from Rhizopus arrhizus, Rhizopus delemar,
Achromobacter sp., and Candida cylindracea and esterase
from hog liver showed activities on PEA and PCL (Tokiwa
and Suzuki 1977a, b).
Poly(ε-caprolactone) (PCL)
PCL ([−OCH2CH2CH2CH2CH2CO–]n) is a biodegradable
synthetic partially crystalline polyester with low melting
point (60 °C) and with a glass transition temperature (Tg)
of −60 °C. It is prepared by ring-opening polymerization of
ε-caprolactone. PCL has been shown to be degraded by the
action of aerobic and anaerobic microorganisms that are
widely distributed in various ecosystems. Furthermore, the
degradation of high molecular weight PCL was investigated
using Penicillium sp. strain 26–1 (ATCC 36507) isolated
from soil. PCL was almost completely degraded in 12 days.
This strain can also assimilate unsaturated aliphatic and
alicyclic polyesters, but not aromatic polyesters (Tokiwa et
al. 1976). A thermotolerant PCL-degrading microorganism
which was identified as Aspergillus sp. strain ST-01 was
isolated from soil. PCL was completely degraded by this
strain after 6 days of incubation at 50 °C (Sanchez et al.
2000). PCL and PHB were degraded under anaerobic con-
dition by new species of microorganisms belonging to the
genus Clostridium (Abou-Zeid et al. 2001). PCL can be
degraded by lipases and esterases (Tokiwa and Suzuki
1977a, b). The degradation rate of PCL is dependent on its
molecular weight and degree of crystallinity. Enzymatic
degradation of PCL by Aspergillus flavus and Penicillium
funiculosum showed that faster degradation was observed in
the amorphous region (Cook et al. 1981).
Poly(β-propiolactone) (PPL)
PPL ([−OCH2CH2CO–]n) is a chemosynthetic biodegrad-
able aliphatic polyester with good mechanical properties.
Many PPL-degrading microorganisms are widely distribut-
ed in various environments, and majority of these microor-
ganisms belong to Bacillus sp. (Nishida et al. 1998). PPL-
degrading microorganisms were isolated from different eco-
systems, and out of 13 isolates, nine of these strains were
identified as Acidovorax sp., Variovorax paradoxus, and
Environ Sci Pollut Res (2013) 20:4339–4355 4345

Table 2 List of fungal strains and the types of plastic which they degrade

Fungal strain Polyesters hydrolyzed Group Reference

Acremonium sp. PHB, poly[3HB-co-(10 mol%) 3HV] Deuteromycota Mergaert et al. (1993)
Aspergillus fischeri PCL Deuteromycota Benedict et al. (1983a, b)
A. flavus ATCC PCL Deuteromycota Benedict et al. (1983a, b)
9643
A. flavus QM380 PEA, PPA, PBA Deuteromycota Darby and Kaplan (1968)
Aspergillus niger PVC Deuteromycota Mogil'nitskii et al. (1987)
van Tieghem F-
1119
A. fumigatus M2A PHB, poly[3HB-co-(7–77 mol%) 3HV], PHV, poly[3HB-co-(13– Deuteromycota Scherer et al. (1999)
61 mol%) 4HB], PES, PEA, PBA, PES/A, PES, PBS/A
A. fumigatus PHB, poly[3HB-co-(10 mol%) 3HV] Deuteromycota Mergaert et al. (1993, 1994)
A. fumigatus LAR 9 PHB, Sky-Green Deuteromycota Kim et al. (2000a, b, c)
A. fumigatus ST-01 PHB, PCL, PBS, PBS/A Deuteromycota Sanchez et al. (2000)
A. fumigatus Pdf1 PHB, poly(3HB-co-3HV), PHV Deuteromycota Iyer et al. (2000)
A. niger ATCC 9642 Sky-Green Deuteromycota Kim et al. (2000a, b, c)
A. niger QM386 PEA, PPA, PBA Deuteromycota Darby and Kaplan (1968)
A. penicilloides PHB Deuteromycota Mergaert et al. (1992)
A. ustus T-221 PHB Deuteromycota Gonda et al. (2000)
A. ustus M-224 PHB Deuteromycota Gonda et al. (2000)
A. ustus LAR 25 Sky-Green Deuteromycota Kim et al. (2000a, b, c)
A. versicolor PEA, PPA, PBA Deuteromycota Darby and Kaplan (1968)
QM432
Aureobasidium PCL Deuteromycota Fields et al. (1974)
pullulans
A. pullulans PEA, PPA, PBA Deuteromycota Darby and Kaplan (1968)
QM279c
Aureobasidium PCL, PU Deuteromycota Fields et al. (1974); Howard
pullulans (2002)
A. pullulans PEA, PPA, PBA Deuteromycota Darby and Kaplan (1968)
QM279c
Candida PHB Deuteromycota Gonda et al. (2000)
guilliermondii
Cephalosporium sp. PHB Deuteromycota Matavulj and Molitoris (1992)
Chaetomium PCL Ascomycota Benedict et al. (1983a, b)
globosum ATCC
6205
C. globosum PEA, PPA, PBA Ascomycota Darby and Kaplan (1968)
QM459
Cladosporium sp. PHB Deuteromycota Matavulj and Molitoris (1992)
Curvularia PE Deuteromycota Howard (2002)
senegalensis
Cryptococcus PCL Basidiomycota Benedict et al. (1983a, b)
laurentii
Curvularia Sky-Green Deuteromycota Kim et al. (2000a, b, c)
protuberate LAR
12
Debaryomyces PHB Ascomycota Gonda et al. (2000)
hansenii
Emericellopsis PHB, poly[3HB-co-(30 mol%) 3HV] Ascomycota Kim et al. (2002a, b, c)
minima W2
Eupenicillium sp. PHB Ascomycota McLellan and Halling (1988)
IMI 300465
Fusarium sp. PCL Deuteromycota Benedict et al. (1983a, b)
F. moniliforme PCL, cutin Deuteromycota Murphy et al. (1996)
F. oxysporum F1-3 Poly[3HB-co-(12 mol%) 3HV] Deuteromycota Sang et al. (2002)
F. solani LAR 11 PHB Deuteromycota Kim et al. (2000a, b, c)
4346 Environ Sci Pollut Res (2013) 20:4339–4355

Table 2 (continued)

Fungal strain Polyesters hydrolyzed Group Reference

F. solani strain 77-2- PCL, cutin Deuteromycota Murphy et al. (1996)

3
F. solani ATCC PCL, cutin Deuteromycota Murphy et al. (1996)
38136
Mucor sp. PHB Zygomycota Matavulj and Molitoris (1992)
Paecilomyces Poly[3HB-co-(12 mol%) 3HV] Deuteromycota Sang et al. (2002)
farinosus F4-7
P. farinosus LAR 10 PHB, Sky-Green Deuteromycota Kim et al. (2000a, b, c)
P. lilacinus D218 PHB, PCL Deuteromycota Oda et al. (1995)
P. lilacinus F4-5 Poly[3HB-co-(12 mol%) 3HV] Deuteromycota Sang et al. (2002)
P. marquandii PHB Deuteromycota Mergaert et al. (1992)
P. simplicissimum PE Ascomycota Yamada-Onodera et al. (2001)
YK
Penicillium PHB Deuteromycota Mergaert et al. (1992)
adametzii
P. argillaceum IFO PCL Deuteromycota Sanchez et al. (2000)
31071
P. chermisinum PHB Deuteromycota Mergaert et al., 1995
P. chrysosporium Poly[3HB-co-(7 mol%) 3HV] Deuteromycota Renstad et al. (1999)
P. daleae PHB Deuteromycota Mergaert et al. (1992)
P. dupontii IFO PCL Deuteromycota Sanchez et al. (2000)
31798
P. funiculosum PHB, PCL Deuteromycota Brucato and Wong (1991); Oda et
ATCC 9644 al. (1995)
P. funiculosum IFO PHB, PHV, poly[3HB-co-(7, 14 %) 4HB], poly[3HB-co-(7, 27, Deuteromycota Miyazaki et al. (2000)
6345 45, 71 %) 3HV]
P. funiculosum PEA, PPA, PBA Deuteromycota Darby and Kaplan (1968)
QM301
P. funiculosum PCL Deuteromycota Benedict et al. (1983a, b)
ATCC 11797
P. funiculosum LAR PHB Deuteromycota Kim et al. (2000a, b, c)
18
P. janthinellum PHB Deuteromycota Mergaert et al. (1995)
P. minioluteum LAR PHB Deuteromycota Kim et al. (2000a, b, c)
14
P. orchrochloron PHB Deuteromycota Mergaert et al. (1992)
P. pinophilium PHB Deuteromycota Han et al. (1998)
ATCC 9644
P. pinophilium LAR PHB Deuteromycota Kim et al. (2000a, b, c)
15
P. restricum PHB Deuteromycota Mergaert et al. (1992)
P. roqueforti PLA Deuteromycota Torres et al. (1996); Pranamuda
et al. (1997)
P. simplicissimum PHB Deuteromycota McLellan and Halling (1988)
IMI 300465
P. simplicissimum PHB, Sky-Green Deuteromycota Kim et al. (2000a, b, c)
LAR 13
P. simplicissimum PHB Deuteromycota Mergaert et al. (1995)
P. simplicissimum Poly[3HB-co-(7 mol%) 3HV] Deuteromycota Renstad et al. (1999)
Penicillium sp. strain PEA, PCL, polyalkylene dicarboxylic acids Deuteromycota Tokiwa and Suzuki (1977a, b)
14-3
Penicillium sp. strain PHB, PCL, polyalkylene dicarboxylic acids Deuteromycota Tokiwa et al. (1976)
26-1
P. verruculosum Mater-Bi Deuteromycota Kim et al. (2000a, b, c)
LAR 17
Phanerochaete Poly[3HB-co-(7 mol%) 3HV] Basidiomycota Renstad et al. (1999); Lee et al.
chrysosporium (1991)
Environ Sci Pollut Res (2013) 20:4339–4355
Table 2 (continued)
Fungal strain Polyesters hydrolyzed
Physarum PMA
polycephalum
Polyporus circinatus PHB
Pullularia pullulans PEA, PPA, PBA
QM279c
Rhizopus delemar PPA, PET copolymers with dicarboxylic acids
R. arrhizus PCL, polyalkylene dicarboxylic acids
Rhodosporidium PHB
sphaerocarpum
Thermoascus PHB, PCL, PBS
aurantiacus IFO
31910
Tritirachium album PLA
ATCC 22563
Verticillium PHB
leptobactrum
Sphingomonas paucimobilis. PHB was also degraded by
these isolates (Kobayashi et al. 1999). Rhizopus delemar
can also degrade PPL (Tokiwa and Suzuki 1977a, b). More-
over, a novel PHB depolymerase from a thermophilic Strep-
tomyces sp. was also capable of degrading PPL (Calabia and
Tokiwa 2006).
Polybutylene succinate (PBS) and polyethylene succinate
(PES)
PBS ([−O(CH2)4OOC(CH2)2CO–]n) and PES
([−O(CH2)2OOC(CH2)2CO–]n) are aliphatic synthetic polyes-
ters with high melting points of 112–114 and 103–106 °C,
respectively. They are synthesized from dicarboxylic acids
(e.g., succinic and adipic acid) and glycols (e.g., ethylene
glycol and 1,4-butanediol). Their mechanical properties are
comparable to polypropylene and low-density polyethylene
(LDPE). PBS-degrading microorganisms are widely distribut-
ed in the environment, but their ratio to the total microorgan-
isms is lower than PCL degraders. The degradation of PBS by
Amycolatopsis sp. HT-6 was investigated, and results showed
that this strain can degrade not only PBS but also PHB and
PCL (Pranamuda et al. 1995). Several thermophilic actinomy-
cetes from the Japan Collection of Microorganisms were
screened for their capability of degrading PBS. Microbispora
rosea, Excellospora japonica, and Excellospora viridilutea
formed a clear zone on agar plates containing emulsified
PBS. Microbispora rosea was able to degrade 50 % (w/v) of
PBS film after 8 days of cultivation in a liquid medium (Jarerat
and Tokiwa 2001a, b). PES is a chemosynthetically aliphatic
polyester which is prepared either by ring-opening polymeri-
zation of succinic anhydride with ethylene oxide or by poly-
condensation of succinic acid and ethylene glycol (Maeda et
4347
Group Reference
Myxomycota Korherr et al. (1995)
Basidiomycota Matavulj and Molitoris (1992)
Deuteromycota Darby and Kaplan (1968)
Zygomycota Walter et al. (1995); Nagata et al.
(1997); Fukuzaki et al. (1989)
Zygomycota Tokiwa et al. (1986)
Basidiomycota Gonda et al. (2000)
Ascomycota Sanchez et al. (2000)
Deuteromycota Jarerat and Tokiwa (2001a, b)
Deuteromycota Mergaert et al. (1994)
al. 1993). In contrast with microbial polyesters which are
susceptible to degradation in various environments, the de-
gradability of PES was found to be strongly dependent on
environmental factors (Kasuya et al. 1997). Moreover, PES-
degrading microorganisms have limited distribution in the
environment in comparison with PHB- and PCL-degrading
microorganisms. A thermophilic Bacillus sp. TT96, a PES
degrader, was isolated from soil. This bacterium can also form
clear zones on PCL and PBS plates, but not on PHB
(Tansengco and Tokiwa 1998). A number of mesophilic
PES-degrading microorganisms were isolated from aquatic
and soil environments. Phylogenetic analysis revealed that
the isolates belong to the genera Bacillus and Paenibacillus.
Among the isolates, strain KT102 which is related to Bacillus
pumilus was chosen since it could degrade PES film at the
fastest rate among the isolates. This strain can degrade PES,
PCL, and olive oil, but not PBS, PHB, and PLA (Tezuka et al.
2004). In addition, several fungi were isolated from various
ecosystems and the isolates formed clear zones around the
colony on agar plates containing PES. A strain NKCM1003
belonging to Aspergillus clavatus was selected, and it can
degrade PES film at a rate of 21 μg/cm2/h (Ishii et al. 2007).
Comparative studies on the biodegradability of three
polyalkylene succinate (PES, PBS, and PPS) with the same
molecular weight were investigated using Rhizopus delemar
lipase. PPS with low Tm (43–52 °C) had the highest biodeg-
radation rate followed by PES, owing to the lower crystallinity
of PPS compared to PES and PBS (Bikiaris et al. 2006).
Aliphatic–aromatic copolyesters (AACs)
It has been reported that aliphatic–aromatic copolyester
(AAC), which consisted of PCL and aromatic polyester such
4348
as polyethylene terephthalate (PET), polybutylene tere-
phthalate, and polyethylene isophthalate (PEIP), was hydro-
lyzed by Rhizopus delemar lipase (Tokiwa and Suzuki
1981). The susceptibility of these AACs to hydrolysis by
Rhizopus delemar lipase decreased rapidly with an increase
in aromatic polyester content. The susceptibility to lipase of
AAC [which consisted of PCL and PEIP, and the latter
being used as a low Tm (103 °C) aromatic polyester] was
greater than those of other AAC. It was assumed that the
rigidity of the aromatic ring in the AAC chains influenced
their biodegradability with this lipase. Another synthetic
AAC-containing adipic acid and terephthalic acid can also
be attacked by microorganisms (Witt et al. 1995). Kleeberg
et al. evaluated the biodegradation of AAC synthesized from
1,4-butanediol, adipic acid, and terephthalic acid.
Thermobifida fusca (known previously as Thermomonospora
fusca) isolated from compost, showed 20-fold higher degra-
dation rates than that usually observed in a common compost
test (Kleeberg et al. 1998). A thermophilic hydrolase from
Thermobifida fusca was found to be inducible not only by
AAC but also by esters. This enzyme was classified as a serine
hydrolase with high similarity to triacylglycerol lipase from
Streptomyces albus G and triacylglycerol acylhydrolase from
Streptomyces sp. M11 (Kleeberg et al. 2005).
Aliphatic polyesters from renewable resources
Poly(3-hydroxybutyrate) (PHB)
PHB ([−O(CH3)CHCH2CO–]n) is a natural polymer pro-
duced by many bacteria as a means to store carbon and
energy. This polymer has attracted research and commercial
interest worldwide because it can be synthesized from re-
newable low-cost feedstocks and the polymerizations are
operated under mild process conditions with minimal envi-
ronmental impact. Furthermore, it can be biodegraded in
both aerobic and anaerobic environments, without forming
any toxic products.
Chowdhury reported for the first time the PHB-degrading
microorganisms from Bacillus, Pseudomonas, and Streptomy-
ces species (Chowdhury 1963). From then on, several aerobic
and anaerobic PHB-degrading microorganisms have been
isolated from soil (Pseudomonas lemoigne, Comamonas sp.,
Acidovorax faecalis, Aspergillus fumigatus, and Variovorax
paradoxus), activated and anaerobic sludge (Alcaligenes
faecalis, Pseudomonas, Ilyobacter delafieldi), seawater, and
lakewater (Comamonas testosterone, Pseudomonas stutzeri)
(Lee 1996). The percentage of PHB-degrading microorgan-
isms in the environment was estimated to be 0.5–9.6 % of the
total colonies (Suyama et al. 1998a, b). Majority of the PHB-
degrading microorganisms were isolated at ambient or
mesophilic temperatures, and very few of them were capable
of degrading PHB at higher temperature. Tokiwa et al.
Environ Sci Pollut Res (2013) 20:4339–4355
emphasized that composting at high temperature is one of
the most promising technologies for recycling biodegradable
plastics and thermophilic microorganisms that could degrade
polymers play an important role in the composting process
(Tokiwa et al. 1992). Thus, microorganisms that are capable of
degrading various kinds of polyesters at high temperatures are
of interest. A thermophilic Streptomyces sp. isolated from soil
can degrade not only PHB but also PES, PBS, and
poly[oligo(tetramethylene succinate)-co-(tetramethylene car-
bonate)]. This actinomycete has higher PHB-degrading activity
than thermotolerant and thermophilic Streptomyces strains from
culture collections (Calabia and Tokiwa 2004). A
thermotolerant Aspergillus sp. was able to degrade 90 % of
PHB film after 5 days of cultivation at 50 °C (Sanchez et al.
2000). Furthermore, several thermophilic polyester-degrading
actinomycetes were isolated from different ecosystems.
Out of 341 strains, 31 isolates were PHB, PCL, and
PES degraders and these isolates were identified as
members of the genera Actinomadura, Microbispora, Strep-
tomyces, Thermoactinomyces, and Saccharomonospora
(Tseng et al. 2007).
Polylactic acid (PLA)
PLA ([−O(CH3)CHCO–]n) is a biodegradable and biocom-
patible thermoplastic which can be produced by fermenta-
tion from renewable resources. It can also be synthesized
either by condensation polymerization of lactic acid or by
ring-opening polymerization of lactide in the presence of a
catalyst (Carothers and Hill 1932). Ecological studies on the
abundance of PLA-degrading microorganisms in different
environments have confirmed that PLA degraders are not
widely distributed, and thus, it is less susceptible to micro-
bial attack compared to other microbial and synthetic ali-
phatic polymers (Suyama et al. 1998a, b; Pranamuda et al.
1997; Tansengco and Tokiwa 1998). The degradation of
PLA in soil is slow and takes a long time to start (Uruyama
et al. 2002; Ohkita and Lee 2006). Microbial degradation of
PLA using Amycolatopsis sp. was first reported by
Pranamuda et al. (1997). Since then, a number of research
studies dealing with microbial and enzymatic degradation of
PLA have been published (Tokiwa and Calabia 2006).
Many strains of genus Amycolatopsis and Saccharothrix
were able to degrade both PLA and silk fibroin. Several
proteinous materials such as silk fibroin, elastin, gelatin, and
some peptides and amino acids were found to stimulate the
production of enzymes from PLA-degrading microorgan-
isms (Pranamuda et al. 2001; Jarerat and Tokiwa 2001a, b;
Jarerat and Tokiwa 2003a, b; Jarerat et al. 2004). Williams
(1981) investigated the enzymatic degradation of PLA using
proteinase K, bromelain, and pronase. Among these en-
zymes, proteinase K from Tritirachium album was the most
effective for PLA degradation. Proteinase K and other serine
Environ Sci Pollut Res (2013) 20:4339–4355
proteases are capable of degrading L-PLA and DL-PLA, but
not D-PLA. Furthermore, proteinase K preferentially hy-
drolyzes the amorphous part of L-PLA and the rate of
degradation decreases with an increase in the crystalline
part (Reeve et al. 1994; McDonald et al. 1996). Fukuzaki
et al. reported that the degradation of PLA oligomers
was accelerated by several esterase-type enzymes, espe-
cially Rhizopus delemar lipase (Fukuzaki et al. 1989).
The purified PLA depolymerase from Amycolatopsis sp.
was also capable of degrading casein, silk fibroin,
Suc–(Ala)3–pNA, but not PCL, PHB, and Suc–(Gly)3–
pNA (Pranamuda et al. 2001). Their studies showed that
PLA depolymerase was a kind of protease and not a
lipase. It was reported that α-chymotrypsin can degrade
PLA and PEA with lower activity on poly[(butylene
succinate)-co-adipate] (PBS/A). Moreover, several serine
proteases such as trypsin, elastase, and subtilisin were
able to hydrolyze L-PLA (Lim et al. 2005).
Different blends of L-PLA/PCL (75/25, 50/50, 25/75)
were prepared, and enzymatic degradation was observed
using proteinase K or Pseudomonas lipase. Proteinase K
was able to degrade the amorphous domain of PLA, but
not the crystalline part of L-PLA or PCL. On the contrary,
Pseudomonas lipase can degrade both the amorphous and
crystalline part of PCL, but not L-PLA (Liu et al. 2000).
Tribedi et al. have reported a new isolated strain of Pseudo-
monas sp. (AKS2) from soil and observed its efficacy in the
biodegradation of PES and found that the degradation was
mediated by esterase activity (Tribedi et al. 2011).
Polymer blends
Blends of polyester with other polymers
The blending of biodegradable polymers is an approach of
reducing the overall cost of the material and modifying the
desired properties and degradation rates. Compared to the
copolymerization method, blending is a much easier and
faster way to achieve the desired properties. Iwamoto and
Tokiwa (1994a, b) developed blend plastics by combining
PCL with conventional plastics, such as LDPE, PP, PS,
nylon 6 (NY), PET, and PHB, and evaluated their level of
enzymatic degradabilities. The blends of PCL/LDPE and
PCL/PP retained the high biodegradability of PCL. In con-
trast, the degradability of the PCL part in the blends of PCL
and PS, PCL and PET, and PCL and PHB dropped off
remarkably. In the case of blends of PCL and NY or PS,
the biodegradability of PCL did not change so much. Thus,
it seems that the higher the miscibility of PCL and conven-
tional plastics, the harder the degradation of PCL on their
blends by Rhizopus arrhizus lipase (Iwamoto and Tokiwa
1994a, b). Furthermore, it was found that degradabilities of
PCL/LDPE (Tokiwa et al. 1990) and PCL/PP (Iwamoto and
4349
Tokiwa 1994a, b) blends by the lipase could be controlled,
depending on their phase structure. Different blends of
PHB have been performed with biodegradable and
nonbiodegradable polymers and polysaccharides. The mis-
cibility, morphology, and biodegradability of PHB blends
with PCL, PBA, and polyvinyl acetate (PVAc) were inves-
tigated. PHB/PCL and PHB/PBA blends were immiscible in
the amorphous state while PHB/PVAc are miscible. Enzy-
matic degradation of these blends was carried out using
PHB depolymerase from Alcaligenes faecalis T1. Results
showed that the weight loss of the blends decreased linearly
with an increase in the amount of PBA, PVAc, or PCL
(Kumagai and Doi 1992). Koyama and Doi 1997 studied
the miscibility, morphology, and biodegradability of
PHB/PLA blend. The spherulites of the blends decreased
with an increase in the content of the PLA, and the rate of
enzymatic surface erosion also decreased with increasing
PLA content in the blend. It was evident that polymer blends
containing PHB usually showed improved properties and
biodegradability when compared with pure PHB (Kumagai
and Doi 1992). Different blends of L-PLA/PCL (75/25,
50/50, 25/75) were prepared, and enzymatic degradation was
observed using proteinase K or Pseudomonas lipase. Protein-
ase K was able to degrade the amorphous domain of PLA, but
not the crystalline part of L-PLA or PCL. On the contrary,
Pseudomonas lipase can degrade both the amorphous and
crystalline part of PCL, but not L-PLA (Liu et al. 2000).
Blends of polyester with starch
Blends of synthetic polymers and starch offer cost–perfor-
mance benefits because starch is renewable, cheap, and
available. In this case, the starch blend can be in the form
of granules or gelatinized starch or even starch which has
been modified chemically to a thermoplastic. It is generally
known that blends of PCL and granular starch exhibit a high
degree of biodegradation (Tokiwa et al. 1990). Takagi et al.
developed PCL/gelatinized starch blends using corn starch
acetates and evaluated their biodegradabilities by an en-
zyme, α-amylase. Their biodegradabilities rapidly de-
creased with an increase in PCL content (Takagi et al. 1994).
Polycarbonates
Aliphatic polycarbonates are known to have greater resistance
to hydrolysis than aliphatic polyesters. The distribution of
PEC (Mn 50,000)-degrading microorganisms seems to be
limited, although PPC (M n 50,000) appears to be
nonbiodegradable. Suyama et al. isolated polyhexamethylene
carbonate (PHC, Mn 2000)-degrading microorganisms which
were phylogenetically diverse. Roseateles depolymerans 61A
formed di(6-hydroxyhexyl) carbonate and adipic acid from
4350
PHC, and di(4-hydroxybutyl) carbonate and succinic acid
from polybutylene carbonate (PBC, Mn 2,000) (Suyama et
al. 1998a, b). Pranamuda et al. found that Amycolatopsis sp.
HT-6 degraded high molecular weight PBC (Mn 37,000). In a
liquid culture containing 150 mg of PBC film, 83 mg of film
was degraded after 7 days of cultivation (Pranamuda et al.
1999). Suyama et al. (1998a, b) reported that a cholesterol
esterase from Candida cylindracea, lipoprotein lipase from
Pseudomonas sp., and lipase from Candida cylindracea,
Chromobacterium viscosum, porcine pancreas, Pseudomonas
sp., and Rhizopus arrhizus degraded PBC (Mn 2000). Lipase
and lipoprotein lipase from Pseudomonas sp. could also de-
grade high molecular weight PBC (Mn 30,000). Lipoprotein
lipase from Pseudomonas sp. produced 1,4-butanediol, CO2,
and di(4-hydroxybutyl) carbonate from PBC (Suyama and
Tokiwa 1997).
Crabbe et al. reported on the degradation of a polyester-
type polyurethane (ES-PU) and the secretion of an enzyme-
like factor with esterase properties by Curvularia
senegalensis, a fungus isolated from soil (Crabbe et al.
1994). Subsequently, Nakajima-Kambe et al. showed that
Comamonas acidovorans strain TB-35 was able to degrade
ES-PU made from polydiethylene adipate (Mn 2,500 and
2,690) and TDI. A purified ES-PU-degrading enzyme from
Comamonas acidovorans TB-35, a type of esterase, hydro-
lyzed the ES-PU and released diethylene glycol and adipic
acid (Nakajima-Kambe et al. 1995). However, it seems that
no microbe can degrade polyurethane completely, and there-
fore, it is difficult to clarify the fate of residues after degra-
dation of ES-PU by both microorganisms and enzymes.
Furthermore, it is difficult to determine whether ET-PU
itself was degraded by microbes to any significant extent.
Polyethylene (PE)
PE is a stable polymer and consists of long chains of
ethylene monomers. PE cannot be easily degraded with
microorganisms. However, it was reported that lower mo-
lecular weight PE oligomers (MW=600–800) were partially
degraded by Acinetobacter sp. 351 upon dispersion, while
high molecular weight PE could not be degraded (Tsuchii et
al. 1980). Furthermore, the biodegradability of low-density
PE/starch blends was enhanced with a compatibilizer (Bikiaris
and Panayiotou 1998). Biodegradability of PE can also be
improved by blending it with biodegradable additives and
photoinitiators or by copolymerization (Griffin 2007;
Hakkarainen and Albertsson 2004). The initial concept of
blending PE with starch was established in the UK to produce
paper-like PE bag. A few years later, the idea to blend PE with
starch and photoinitiators was conceived in the USA as a way
of saving petroleum, though its biodegradability was also
taken into account. Environmental degradation of PE pro-
ceeds by synergistic action of photo- and thermo-oxidative
Environ Sci Pollut Res (2013) 20:4339–4355
degradation and biological activity (i.e., microorgan-
isms). When PE is subjected to thermo- and photo-
oxidization, various products such as alkanes, alkenes,
ketones, aldehydes, alcohols, carboxylic acid, keto acids,
dicarboxylic acids, lactones, and esters are released. Blending
of PE with additives generally enhances auto-oxidation, re-
duces the molecular weight of the polymer, and then makes it
easier for microorganisms to degrade the low molecular
weight materials. It is worthy to note that despite all these
attempts to enhance the biodegradation of PE blends, the
biodegradability with microorganisms on the PE part of the
blends is still very low.
Polypropylene (PP)
PP is a thermoplastic which is commonly used for plastic
moldings, stationary folders, packaging materials, plastic
tubs, nonabsorbable sutures, diapers, etc. PP can be degrad-
ed when it is exposed to ultraviolet radiation from sunlight.
Furthermore, at high temperatures, PP is oxidized. The
possibility of degrading PP with microorganisms has been
investigated (Cacciari et al. 1993).
Polystyrene (PS)
PS is a synthetic hydrophobic polymer with high molecular
weight. PS is recyclable but not biodegradable. Although it
was reported that PS film was biodegraded with an actino-
mycete strain, the degree of biodegradation was very low
(Mor and Silvan 2008). At room temperature, PS exists in
solid state. When it is heated above its glass transition
temperature, it flows and then turns back to solid upon
cooling. PS being a transparent hard plastic is commonly
used as disposable cutleries, cups, plastic models, and pack-
ing and insulation materials.
Table 3 revealed 15 bacterial genera which have the ca-
pacity to degrade various types of plastics. Among them
Pseudomonas is dominant. It can degrade polythene, PVC,
PHB, poly(3-hydroxybutyrate-co-3-mercaptopropionate), and
poly(3-droxypropionate). Bacillus brevis can degrade only
polycaprolactone while Streptomyces can degrade PHB,
poly(3-hydoxybutyarate-co-3-hydroxyvalerate), and starch
or polyester. Ochrobactrum TD is also able to degrade PVC.
Majority of the strains that are able to degrade PHB
belong to different taxa such as Gram-positive and Gram-
negative bacteria, Streptomyces, and fungi (Mergaert and
Swings 1996). It has been reported that 39 bacterial strains
of the classes Firmicutes and Proteobacteria can degrade
PHB, PCL, and PBS, but not PLA (Suyama et al. 1998a,
b). Other bacterial species identified having the properties of
degrading plastics were Bacillus sp., Staphylococcus sp.,
Streptococcus sp., Diplococcus sp., Micrococcus sp., Pseu-
domonas sp., and Moraxella sp. (Kathiresan 2003).
Environ Sci Pollut Res (2013) 20:4339–4355 4351

Table 3 List of microbial strains and the types of plastic which they degrade

Plastic Microorganism Reference

Polyethylene Brevibacillus borstelensis Hadad et al. (2005)

Rhodococcus rubber Sivan et al. (2006); Gilan et al. (2004)
Pseudomonas chlororaphis Zheng et al. (2005)
Comamonas acidovorans TB-35 Akutsu et al. (1998)
Polyvinyl chloride Pseudomonas putida AJ Anthony et al. (2004)
Ochrobactrum TD Mogil'nitskii et al. (1987)
Pseudomonas fluorescens B – 22
BTA copolyester Thermomonospora fusca Kleeberg et al. (1998)
Some biodegradable/natural plastics and their degrading microorganisms
Poly(3-hydroxybutyrate-co-3-mercaptopropionate) Schlegelella thermodepolymerans Elbanna et al. (2004)
Poly(3-hydroxybutyrate) Pseudomonas lemoignei Jendrossek et al. (1995)
Poly(3-hydroxybutyrate-co-3-mercaptopropionate) Pseudomonas indica K2 Elbanna et al. (2004)
Poly(3-hydroxybutyrate), poly(3-hydroxybutyrate- Streptomyces sp. SNG9 Mabrouk and Sabry (2001)
co-3-hydroxyvalerate)
Poly(3-hydroxybutyrate-co-3-hydroxypropionate) Ralstonia pikettii T1 Wang et al. (2002)
Acidovorax sp. TP4
Poly (3-hydroxybutyrate), poly(3- Alcaligenes faecalis Kasuya et al. (1999)
hydroxypropionate), poly(4-hydroxybutyrate), Pseudomonas stutzeri
polyethylene succinate, polyethylene adipate Comamonas acidovorans
Poly(3-hydroxybutyrate) Alcaligenes faecalis Kita et al. (1997)
Schlegelella thermodepolymerans
Caenibacterium thermophilum Romen et al. (2004)
Poly(3-hydroxybutyrate-co-3-hydroxyvalerate) Clostridium botulinum Abou-Zeid et al. (2001)
Clostridium acetobutylicum
Polycaprolactone Clostridium botulinum Abou-Zeid et al. (2001)
Clostridium acetobutylicum
Amycolatopsis sp.
Bacillus brevis
Polymer blends and its degrading microorganisms
Starch/polyester Streptomyces Lee et al. (1991)

Mechanism of biodegradation of plastics by microbes
The mechanisms of action of microorganisms on polymers
are influenced by two different processes:
1. Direct action: In this case, the deterioration of plastics
serves as a nutritive substance for the growth of the
microorganisms.
2. Indirect action: The influence of the metabolic products
of the microorganisms, e.g., discoloration or further
deterioration.
Biodegradation of a polymeric material is a chemical
degradation brought by the action of naturally occurring
microorganisms such as bacteria and fungi via enzymat-
ic action into metabolic products of microorganisms (e.g.,
H2O, CO2, CH4, biomass, etc.) (David et al. 1994; Chandra
and Rustgi 1998; Lenz 1993; Mohanty et al. 2000). Lipase ,
proteinase K, pronase, hydrogenase, etc. are important
enzymes secreted by microbes for plastic biodegrada-
tion. Among these enzymes, proteinase K from
Tritirachium album was the most effective for PLA
degradation. Many strains of the genera Amycolatopsis
and Saccharothrix were able to degrade PLA. Fukuzaki
et al. reported that the degradation of PLA oligomers
was accelerated by several esterase-type enzymes, espe-
cially Rhizopus delemar lipase (Fukuzaki et al. 1989).
Several serine proteases such as trypsin, elastase, and
subtilisin were able to hydrolyze L-PLA (Lim et al.
2005). In contrast to the biodegradation of polymers,
where a near complete conversion of the material com-
ponents takes place, only a change in the polymer
structure or the plastic composition is observed in many
cases in polymer biodeterioration or biocorrosion (Gu
2003). The ultimate result in the both the cases are a complete
loss of structural integrity as a result of a drastic decrease in
molecular weight.
4352
Conclusion
As it is the plastic era, we cannot think beyond it, but
plastics become one of the major problems for the modern
environmentalist. To get rid of such a menace, people usu-
ally put them in landfills or burn them, but both these
practices cause very serious threats to the environment and
the ecosystem. Burning plastics usually produces some nox-
ious gases like furans and dioxins, which are some danger-
ous greenhouse gases and play an important role in ozone
layer depletion. In fact, dioxins cause serious problems in
the human endocrine hormone activity, thus becoming a
major concern for the human health too (NoPE 2002; Pilz
et al. 2010). Dioxins also cause very serious soil pollution,
causing a great concern for the scientific community world-
wide. Hence, under such circumstances degradation of plastic
by microbes is one of the eco-friendly and innovative
methods. Many fungal genera (e.g., Acremonium,
Cladosporium, Debaryomyces, Emericellopsis,
Eupenicillium, Fusarium, Mucor, Paecilomyces, Penicillium,
Pullularia, Rhodosporidium, Verticillium, Aspergillus,
Aureobasidium, Chaetomium, Cryptococcus, Fusarium, Rhi-
zopus, Penicillium, Thermoascus, Penicillium roqueforti,
Tritirachium album, etc.) and bacterial genera (Brevibacillus,
Streptomyces, Amycolatopsis, Clostridium, Schlegelella,
Pseudomonas, etc.) have been reported to degrade various
kinds of plastics (PEA, PPA, PBA, PCL, PVC, PHB, PU,
etc.). This review may give brief information regarding
the nature and biodegradation of plastic by means of
different microbes. It is expected that this review work
will encourage young scientists to find out one or more
microbial strain(s) from nature for the potential biodeg-
radation of plastic wastes.
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