Study of Microbes Having Potentiality For Biodegradation of Plastics

Environ Sci Pollut Res (2013) 20:43394355
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 degradable 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 environmental 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 degradation 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 enzymatic 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 biodegradable. Question may arise why nonbiodegradable plastics 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 generally 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 enzymes 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
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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 worlds largest
plastic consumers, accounting about 35 % of the global consumption 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. Environmental 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 polymers 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 consumption 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 plasticconsuming sector, with 42 % of the total consumption,
followed by consumer products and the construction industry. 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 development 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 employs about eight times more people than the plasticmanufacturing sector. Due to the increasing share of long-
life products in the economy, and consequently in the volume of waste generated, the share of recycling will decrease
to 35 % over the next three decades. The total waste available 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 effective 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 problems 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 dangerous 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 degrade 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 capacity to degrade these synthetic polymers in much faster way
in comparison to the natural method by using some exoenzymes 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(-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). Moreover, a novel poly(3-hydroxybutyrate) (PHB) depolymerase
from a thermophilic Streptomyces sp. was also capable of
degrading poly(-propiolactone) (PPL) (Calabia and
Tokiwa 2006). In comparative studies, the biodegradability
of three polyalkylene succinate [polyethylene succinate

(PES), polybutylene succinate (PBS), and polypropylene
succinate (PPS)] with the same molecular weight was investigated using Rhizopus delemar lipase. PPS with low
melting temperature (Tm, 4352 C) had the highest biodegradation 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 biodegradation 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.
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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 plastics 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 biodegradable plastic (which comprises polyhydroxybutyrate) available 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
Classification of plastic according to their thermal properties

Based on their thermal properties, plastics are classified into
two groups: thermoplastics and thermosetting polymers.
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.
Thermoplastics
Effect of plastic on the environment and human health
Thermoplastics are those types of plastics which cannot
undergo chemical changes in their composition when heated, 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 irreversible and hence they are not recyclable too. Examples
include phenolformaldehyde, 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
With the rapid modernization and development of human

society, there has been a very sharp increase in the demand
of plastic-based products. Approximately 3040 % of plastics 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 important 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 plasticbased 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
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Table 1 List of various types of plastic we use everyday

Types
Use/application
Polyester (PES)
Polyethylene terephthalate (PET)
Polyethylene (PE)
High-density polyethylene (HDPE)
Polyvinyl chloride (PVC)
Polyvinylidene chloride (PVDC)
(Saran)
Low-density polyethylene (LDPE)
Polypropylene (PP)
Fibers, textiles
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
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)
Ureaformaldehyde (UF)
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, plumbers
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
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 antagonist 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 component 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 females (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 governments 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 Wildlife 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 Brydes whale near Cairns, Australia revealed that the
stomach of the whale was filled with plastic rubbishes which
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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 monomeric 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 microorganisms 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
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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 condition, may be directed to grow and use those carbon polymers 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 molecular weight (Mn) 3,000) as the sole source of carbon. Among
the isolated PEA-degrading microorganisms, Penicillium sp.
strain 143 exhibited the strongest activity. PEA was degraded 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
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 -3hydroxybutyrate) (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 143 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(-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
-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 261 (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 condition 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(-propiolactone) (PPL)
PPL ([OCH2CH2CO]n) is a chemosynthetic biodegradable aliphatic polyester with good mechanical properties.
Many PPL-degrading microorganisms are widely distributed in various environments, and majority of these microorganisms belong to Bacillus sp. (Nishida et al. 1998). PPLdegrading microorganisms were isolated from different ecosystems, and out of 13 isolates, nine of these strains were
identified as Acidovorax sp., Variovorax paradoxus, and
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Table 2 List of fungal strains and the types of plastic which they degrade
Fungal strain
Polyesters hydrolyzed
Group
Reference
Acremonium sp.
Aspergillus fischeri
A. flavus ATCC
9643
A. flavus QM380
Aspergillus niger
van Tieghem F1119
A. fumigatus M2A
PHB, poly[3HB-co-(10 mol%) 3HV]

PCL
PCL
Deuteromycota
Deuteromycota
Deuteromycota
Mergaert et al. (1993)

Benedict et al. (1983a, b)
PEA, PPA, PBA

PVC
Deuteromycota
Deuteromycota
Darby and Kaplan (1968)

Mogil'nitskii et al. (1987)
PHB, poly[3HB-co-(777 mol%) 3HV], PHV, poly[3HB-co-(13

61 mol%) 4HB], PES, PEA, PBA, PES/A, PES, PBS/A
PHB, Sky-Green
PHB, PCL, PBS, PBS/A
Deuteromycota
Scherer et al. (1999)
Deuteromycota
Deuteromycota
Deuteromycota
Mergaert et al. (1993, 1994)

Kim et al. (2000a, b, c)
Sanchez et al. (2000)
PHB, poly(3HB-co-3HV), PHV

Sky-Green
PEA, PPA, PBA
PHB
PHB
PHB
Sky-Green
PEA, PPA, PBA
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota
Iyer et al. (2000)

Gonda et al. (2000)
Gonda et al. (2000)
PCL
Deuteromycota
Fields et al. (1974)
PEA, PPA, PBA
Deuteromycota
PCL, PU
Deuteromycota
PEA, PPA, PBA
Deuteromycota
Fields et al. (1974); Howard

(2002)
PHB
Deuteromycota
Gonda et al. (2000)
PHB
PCL
Deuteromycota
Ascomycota
Matavulj and Molitoris (1992)

PEA, PPA, PBA
Ascomycota
PHB
PE
Deuteromycota
Deuteromycota

Howard (2002)
PCL
Basidiomycota
Sky-Green
Deuteromycota
PHB
Ascomycota
Gonda et al. (2000)
Ascomycota
PHB
Ascomycota
McLellan and Halling (1988)
PCL
PCL, cutin
Poly[3HB-co-(12 mol%) 3HV]
PHB
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota

Murphy et al. (1996)
Sang et al. (2002)
A. fumigatus
A. fumigatus LAR 9
A. fumigatus ST-01
A. fumigatus Pdf1
A. niger ATCC 9642
A. niger QM386
A. penicilloides
A. ustus T-221
A. ustus M-224
A. ustus LAR 25
A. versicolor
QM432
Aureobasidium
pullulans
A. pullulans
QM279c
Aureobasidium
pullulans
A. pullulans
QM279c
Candida
guilliermondii
Cephalosporium sp.
Chaetomium
globosum ATCC
6205
C. globosum
QM459
Cladosporium sp.
Curvularia
senegalensis
Cryptococcus
laurentii
Curvularia
protuberate LAR
12
Debaryomyces
hansenii
Emericellopsis
minima W2
Eupenicillium sp.
IMI 300465
Fusarium sp.
F. moniliforme
F. oxysporum F1-3
F. solani LAR 11
4346
Table 2 (continued)
Fungal strain
Group
Reference
F. solani strain 77-23

F. solani ATCC
38136
Mucor sp.
Paecilomyces
farinosus F4-7
P. farinosus LAR 10
P. lilacinus D218
P. lilacinus F4-5
P. marquandii
PCL, cutin
Deuteromycota
PCL, cutin
Deuteromycota
PHB
Zygomycota
Deuteromycota

Sang et al. (2002)
PHB, Sky-Green
PHB, PCL
PHB
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota

Oda et al. (1995)
Sang et al. (2002)
P. simplicissimum
YK
Penicillium
adametzii
P. argillaceum IFO
31071
P. chermisinum
P. chrysosporium
P. daleae
P. dupontii IFO
31798
P. funiculosum
ATCC 9644
P. funiculosum IFO
6345
P. funiculosum
QM301
P. funiculosum
ATCC 11797
P. funiculosum LAR
18
P. janthinellum
P. minioluteum LAR
14
P. orchrochloron
P. pinophilium
ATCC 9644
P. pinophilium LAR
15
P. restricum
P. roqueforti
PE
Ascomycota
Yamada-Onodera et al. (2001)
PHB
Deuteromycota
PCL
Deuteromycota
PHB
PHB
PCL
Deuteromycota
Deuteromycota
Deuteromycota
Deuteromycota
Mergaert et al., 1995

Renstad et al. (1999)
PHB, PCL
Deuteromycota
PHB, PHV, poly[3HB-co-(7, 14 %) 4HB], poly[3HB-co-(7, 27,

45, 71 %) 3HV]
PEA, PPA, PBA
Deuteromycota
Brucato and Wong (1991); Oda et

al. (1995)
Miyazaki et al. (2000)
Deuteromycota
PCL
Deuteromycota
PHB
Deuteromycota
PHB
PHB
Deuteromycota
Deuteromycota

PHB
PHB
Deuteromycota
Deuteromycota

Han et al. (1998)
PHB
Deuteromycota
PHB
PLA
Deuteromycota
Deuteromycota
PHB
Deuteromycota

Torres et al. (1996); Pranamuda
et al. (1997)
McLellan and Halling (1988)
PHB, Sky-Green
Deuteromycota
PHB
PEA, PCL, polyalkylene dicarboxylic acids
Deuteromycota
Deuteromycota
Deuteromycota

Renstad et al. (1999)
Tokiwa and Suzuki (1977a, b)
PHB, PCL, polyalkylene dicarboxylic acids
Deuteromycota
Tokiwa et al. (1976)
Mater-Bi
Deuteromycota
Basidiomycota
Renstad et al. (1999); Lee et al.

(1991)
P. simplicissimum
IMI 300465
P. simplicissimum
LAR 13
P. simplicissimum
P. simplicissimum
Penicillium sp. strain
14-3
Penicillium sp. strain
26-1
P. verruculosum
LAR 17
Phanerochaete
chrysosporium
4347
Table 2 (continued)
Fungal strain
Group
Reference
Physarum
polycephalum
Polyporus circinatus
Pullularia pullulans
QM279c
Rhizopus delemar
PMA
Myxomycota
Korherr et al. (1995)
PHB
PEA, PPA, PBA
Basidiomycota
Deuteromycota

PPA, PET copolymers with dicarboxylic acids
Zygomycota
PCL, polyalkylene dicarboxylic acids

PHB
Zygomycota
Basidiomycota
Walter et al. (1995); Nagata et al.

(1997); Fukuzaki et al. (1989)
Tokiwa et al. (1986)
Gonda et al. (2000)
PHB, PCL, PBS
Ascomycota
PLA
Deuteromycota
Jarerat and Tokiwa (2001a, b)
PHB
Deuteromycota
R. arrhizus
Rhodosporidium
sphaerocarpum
Thermoascus
aurantiacus IFO
31910
Tritirachium album
ATCC 22563
Verticillium
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). Moreover, a novel PHB depolymerase from a thermophilic Streptomyces 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 polyesters with high melting points of 112114 and 103106 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 distributed in the environment, but their ratio to the total microorganisms 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 actinomycetes 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 polymerization of succinic anhydride with ethylene oxide or by polycondensation of succinic acid and ethylene glycol (Maeda et
al. 1993). In contrast with microbial polyesters which are

susceptible to degradation in various environments, the degradability of PES was found to be strongly dependent on
environmental factors (Kasuya et al. 1997). Moreover, PESdegrading 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 g/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 (4352 C) had the highest biodegradation rate followed by PES, owing to the lower crystallinity
of PPS compared to PES and PBS (Bikiaris et al. 2006).
Aliphaticaromatic copolyesters (AACs)
It has been reported that aliphaticaromatic copolyester
(AAC), which consisted of PCL and aromatic polyester such
4348
as polyethylene terephthalate (PET), polybutylene terephthalate, and polyethylene isophthalate (PEIP), was hydrolyzed 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 degradation 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 produced 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 renewable low-cost feedstocks and the polymerizations are
operated under mild process conditions with minimal environmental 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 Streptomyces 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 microorganisms in the environment was estimated to be 0.59.6 % of the
total colonies (Suyama et al. 1998a, b). Majority of the PHBdegrading microorganisms were isolated at ambient or
mesophilic temperatures, and very few of them were capable
of degrading PHB at higher temperature. Tokiwa et al.
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 carbonate)]. 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, Streptomyces, Thermoactinomyces, and Saccharomonospora
(Tseng et al. 2007).
Polylactic acid (PLA)
PLA ([O(CH3)CHCO]n) is a biodegradable and biocompatible thermoplastic which can be produced by fermentation 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 microbial attack compared to other microbial and synthetic aliphatic 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 microorganisms (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 enzymes, proteinase K from Tritirachium album was the most
effective for PLA degradation. Proteinase K and other serine
4349
proteases are capable of degrading L-PLA and DL-PLA, but

not D-PLA. Furthermore, proteinase K preferentially hydrolyzes 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, especially Rhizopus delemar lipase (Fukuzaki et al. 1989).
The purified PLA depolymerase from Amycolatopsis sp.
was also capable of degrading casein, silk fibroin,
Suc(Ala)3pNA, 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 -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 Pseudomonas 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).
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 miscibility, morphology, and biodegradability of PHB blends
with PCL, PBA, and polyvinyl acetate (PVAc) were investigated. PHB/PCL and PHB/PBA blends were immiscible in
the amorphous state while PHB/PVAc are miscible. Enzymatic 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. 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).
Polymer blends
Blends of synthetic polymers and starch offer costperformance 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 enzyme, -amylase. Their biodegradabilities rapidly decreased with an increase in PCL content (Takagi et al. 1994).
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 contrast, 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 conventional 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
Blends of polyester with starch
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 degrade 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 polyestertype polyurethane (ES-PU) and the secretion of an enzymelike 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, hydrolyzed 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 therefore, it is difficult to clarify the fate of residues after degradation 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 molecular weight PE oligomers (MW=600800) 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 proceeds by synergistic action of photo- and thermo-oxidative
degradation and biological activity (i.e., microorganisms). When PE is subjected to thermo- and photooxidization, 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, reduces 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 degraded 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 actinomycete 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 packing and insulation materials.
Table 3 revealed 15 bacterial genera which have the capacity 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 Gramnegative 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., Pseudomonas sp., and Moraxella sp. (Kathiresan 2003).
4351
Table 3 List of microbial strains and the types of plastic which they degrade
Plastic
Microorganism
Polyethylene
Brevibacillus borstelensis
Rhodococcus rubber
Pseudomonas chlororaphis
Comamonas acidovorans TB-35
Polyvinyl chloride
Pseudomonas putida AJ
Ochrobactrum TD
Pseudomonas fluorescens B 22
BTA copolyester
Thermomonospora fusca
Some biodegradable/natural plastics and their degrading microorganisms
Poly(3-hydroxybutyrate-co-3-mercaptopropionate)
Schlegelella thermodepolymerans
Poly(3-hydroxybutyrate)
Pseudomonas lemoignei
Poly(3-hydroxybutyrate-co-3-mercaptopropionate)
Pseudomonas indica K2
Poly(3-hydroxybutyrate), poly(3-hydroxybutyrateStreptomyces sp. SNG9
co-3-hydroxyvalerate)
Poly(3-hydroxybutyrate-co-3-hydroxypropionate)
Ralstonia pikettii T1
Acidovorax sp. TP4
Poly (3-hydroxybutyrate), poly(3Alcaligenes faecalis
hydroxypropionate), poly(4-hydroxybutyrate),
Pseudomonas stutzeri
polyethylene succinate, polyethylene adipate
Comamonas acidovorans
Poly(3-hydroxybutyrate)
Alcaligenes faecalis
Schlegelella thermodepolymerans
Caenibacterium thermophilum
Poly(3-hydroxybutyrate-co-3-hydroxyvalerate)
Clostridium botulinum
Clostridium acetobutylicum
Polycaprolactone
Clostridium botulinum
Clostridium acetobutylicum
Amycolatopsis sp.
Bacillus brevis
Polymer blends and its degrading microorganisms
Starch/polyester
Streptomyces
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 enzymatic 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
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Lee et al. (1991)
enzymes secreted by microbes for plastic biodegradation. 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, especially 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 components 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 usually 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 noxious gases like furans and dioxins, 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. 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, Rhizopus, 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 biodegradation of plastic wastes.
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