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BIOTECHNOLOGY - Vol. V - Bioplastic and Biopolymer Production - Ian W. Sutherland
Encyclopedia of Life Support Systems (EOLSS)

BIOPLASTIC AND BIOPOLYMER PRODUCTION

Ian W. Sutherland,
Institute of Cell and Molecular Biology, University of Edinburgh, Edinburgh, United
Kingdom

Keywords: ABC-transporters, Archae, Bacterial alginate, Bioplastics, Biopolymers,
Biosynthesis, Carbon and energy reserves,Cellulose, Curdlan, Dextrans,
Dextransucrase, -D-Glucans, -D-Glucans, Exopolysaccharides (EPS), Fructans,
Gellan, Heparinm, Heteropolysaccharides , Hyaluronic Acid, Levan, levansucrase,
Negative regulation, Polyhydroxyalkanoic acids (PHAs), -hydroxybutyric acid (PHB),
Polysaccharides, Positive regulation, Regulation of synthesis, Scleroglucan, lucan,
Xanthan

Contents

1. Bioplastics
1.1. Introduction
1.2. Occurrence & Composition
1.3. Biosynthesis
1.4. Products
1.5. Production and Recovery
2. Biopolymers (Polysaccharides)
2.1. Introduction
2.2. Occurrence and Composition
2.3. Polysaccharide biosynthesis
2.3.1. Biosynthetic mechanisms
2.3.2. Genetics and Regulation of Exopolysaccharides Synthesis
2.4. Commercial Products
2.4.1. -D-Glucans
2.4.2. -D-Glucans
2.4.3. Bacterial Alginates
2.4.4. Gellan and Related Polymers
2.4.5. Hyaluronic Acid and Heparin
2.4.6. Succinoglycan and Galactoglucans
2.4.7. Xanthan
2.5. Production of exopolysaccharides
3. Future Developments
3.1. Bioplastics
3.2. Biopolymers
Glossary
Bibliography
Biographical Sketch

Summary

Micro-organisms provide a source of bioplastics and biopolymers (polysaccharides)
from renewable sources. Although currently considerably more expensive than plastic
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derived from petrochemicals, bacteria have proved capable of yielding bioplastics with
comparable properties. They have the additional advantage of being biodegradable. The
processes, which have been developed for their production, have been optimised and a
range of products obtained. Research is currently determining the possibility of
transferring the genes for synthesis of these bioplastics to plants in order to test the
commercial viability of such sources. Considerable effort is also being applied to the
discovery of new bacterial sources of polymers with different properties.

A very wide range of microbial polysaccharides has been studied and structure/function
relationships for a number of these macromolecules have been determined. Several
biopolymers have gained acceptability in a wide range of applications in manufactured
food technology, in other industrial applications and as useful adjuncts in oil exploration
and production. Some of the biopolymers possess unique properties and are of high
value. These products are generally competing against a range of established plant and
algal gums, including starch. However, they are also prepared from renewable resources
and are biodegradable.

1. Bioplastics

1.1. Introduction

When grown under conditions where growth becomes limited through exhaustion of a
key nutrient such as nitrogen or phosphorus and carbon substrate remains available,
many prokaryotes can synthesise intracellular storage compounds. These act as carbon
and energy reserves which can be utilised when balanced growth is resumed. One such
is glycogen, while another type was initially identified as poly--hydroxybutyric acid
(PHB). These storage compounds can represent over 50 per cent of cell dry weight
under some growth conditions and are normally recognised through use of iodine and
sudanophilic dyes respectively.

1.2. Occurrence & Composition

The occurrence of polyhydroxyalkanoic acids as storage polymers in prokaryotic cells is
now known to be very widespread indeed. They are water-insoluble compounds with
the general structure shown in Figure 1. Many bacteria produce an intracellular carbon
and energy storage compound - poly--hydroxybutyric acid (PHB) - in relatively large
quantities . While this property is absent from enteric species, it is widely found in
Pseudomonads and related species including the plant symbiont Rhizobium and also in
nitrogen-fixing Azotobacter spp. Accumulation is normally a response to unbalanced
growth in the presence of excess carbon and energy source. Under appropriate
conditions the polymer can amount to more than 50-80 per cent of cell dry weight. The
storage product is found as granular inclusion bodies within the cytoplasm. However,
many of these compounds represent in relatively small amounts or because of their short
chain lengths or other properties are unsuitable as potential bioplastics. Among species
synthesising PHB and PHV are some Archae including Haloferax mediterranea. These
halophilic bacteria might present advantages for production as their culture
requirements of salinity and relatively high temperature provide little opportunity for
growth of contaminants. In species such as Azotobacter vinelandii, simultaneous
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production of large amounts of exopolysaccharide diverts substrate to alternative
products and makes recovery of PHB difficult. Development of high-yielding mutant
strains resulted in conversion rates of 65 per cent for PHB and eventual PHA yields of
71 per cent dry weight



Figure 1: The typical structure of a polyhydroxyalkanoate copolymer

1.3. Biosynthesis



Figure 2: The synthesis of PHB

Biosynthesis of PHB is much simpler than the formation of most polysaccharides, as
only 3 enzymes are normally involved - -thioketolase, acetoacetylCoA reductase and
PHB synthase (Figure 2). The genes involved have been studied in Ralstonia eutropha
(formerly designated Alcaligenes eutrophus) and Pseudomonas oleovorans as well as
other species. Genetic control of the process is also relatively simple; the 3 genes
involved are organised in an operon to form a sequence of 3 open reading frames and
these can be transferred relatively easily to other bacterial species and to yield
transgenic plant species. A slightly modified alternative pathway found in
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Rhodospirillum rubrum was outlined by Steinbchel and Fchtenbusch. The PHA
synthase gene products from different bacteria have been grouped into 3 classes varying
in their constitutive proteins and their substrate specificity.

Recent studies using recombinant strains of E.coli into which the PHA biosynthesis
genes from R. eutropha had been inserted, yielded PHB with mass of 3-11 x 10
6

daltons. The mechanical properties of films from this product were improved by
stretching over 400 per cent. The enzymes for PHB synthesis are normally constitutive
and regulation of synthesis appears to occur at the enzyme level.

1.4. Products

Bioplastics are thermoplastic compounds which, unlike products of the petrochemical
industry, are biodegradable. They have the further advantage that they can be produced
from renewable resources. They are normally highly crystalline, optically active and
possess piezoelecetric properties. ICI (Zeneca) put considerable effort into development
of bioplastics and soon found that PHB did not have all the properties they wanted.
However they discovered that it was possible to synthesise linear co-polymers
containing poly--hydroxybutyric acid and hydroxyvaleric acid. Such random co-
polymers were formed by R. eutropha with glucose and propionic acid as substrates.
Typically polymers consisted of 12-20 per cent hydroxyvalerate and approximately
300000 Da. The melting point of the co-polymers decreases with increasing
hydroxyvaleric acid content. The melting point of PHB is close to 175
o
C whereas those
of PHAs are lower. Unlike PHB which is brittle, the copolymers are elastic. PHA are
thermoplastic polymers and they become highly viscous at temperatures above their
melting point, thus rendering them mouldable. The melting point (T
m
), crystallinity and
glass transition temperature (T
g
) depend on the composition of the product. The
commercialised product 'Biopol' has been used to form biodegradable plastic bottles,
together with golf tees, disposable razors and other products. However the high cost of
the product in relation to chemically synthesised plastics has led to closure of
production. Alternative applications which have been proposed, are as water-
impermeable coatings for biodegradable packaging, as temporary plates and pegs in
repair of bone injuries etc. , but it is unclear whether a market exists under the current
economic climate.

1.5. Production and Recovery

Wild-type strains of R. eutropha utilise fructose but not glucose. However, glucose-
utilising strains have been developed for use in the commercial production of 'Biopol'.
PHA-producing bacteria can be grown on a large scale to high cell densities in stirred
tank fermenters using glucose, sucrose or molasses as carbon source. Fed-batch systems
have been preferred to continuous processes and yields of 70-80 per cent PHA have
been reported for R. eutropha grown in a mineral salts medium on glucose
supplemented with propionate as the sole carbon sources. Cell densities were of the
order of 100g. dry weight litre
-1
. Following cell harvesting by centrifugation or by
flocculation, the PHA can be recovered by the use of surfactants and hypochlorite to
lyse the cells and release the intracellular product. Although hypochlorite provided a
suitable laboratory procedure it can lead to some degradation of the product. Flotation
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processes have also been used to separate the PHA from soluble intracellular products.
Alternatively, solvent extraction has been proposed. This can only be effectively
achieved if the cell mass has first been dried by spray drying or by lyophilisation, thus
adding to the cost of the process. Large amounts of solvent are needed as concentrated
solutions of PHA are highly viscous. Chloroform or methylene chloride can dissolve
most PHA products but pretreatment of the cells may be required to achieve maximum
extraction efficiency. The actual yield and composition of the product depend on the
substrate regime chosen. Choi and Lee observed initial low polymer content of cells due
to the high residual content of propionic acid in the medium. By altering the feed
strategy, use of acetic acid induction and oleic acid addition, P(3HB-co-3HV)-polymer
yields of 78 per cent wt. and productivity of 2.88 g l
-1
h
-1
were obtained.

2. Biopolymers (Polysaccharides)

2.1. Introduction

A large number of micro-organisms produce exopolysaccharides. The structures of
many of these polymers have now been accurately determined. Some chemical
structures have also been correlated with the physical functions of EPS. In several
bacteria of medical significance, including Streptococcus pneumoniae, Escherichia coli
and Klebsiella aerogenes, systematic studies on a large number of different serotypes
have also determined the relationship between serological specificity and chemical
structure. Other bacterial groups have been studied because of either their pathogenicity
or their symbiotic interactions with plants and the roles which polysaccharides play in
such processes.

Although large numbers of microbial polysaccharides are potentially available,
relatively few have been commercially developed. There are many reasons for this: the
microbial source may be pathogenic; production costs may be very high; product quality
may be difficult to maintain and to guarantee; the product may not achieve regulatory
acceptability or (very commonly) there is no market niche. Despite such problems,
several EPS are now recognised products of biotechnology. Several more may be
developed in the next few years, especially as we look to renewable resources for
alternatives to several chemical products.

Substituent Occurrence Linkage Charge
conferrred
Organic Acids
Acetate
Very common - e.g. Klebsiella
spp.; colanic acid
Ester None
Glycerate Sphingomonas
elodea
Ester Negative
Hydroxybutanoate Rhizobium
trifolii; R. leguminosarum, etc.
Ester None
Propionate Rare - some
Ester None
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Escherichia coli
Pyruvate Very common - e.g.
Klebsiella spp.; colanic acid
Ketal Negative
Succinate Rhizobium spp.;
Agrobacterium spp.
Half ester Negative
Inorganic acids
Phosphate Common in some
genera, Gram positive spp.

Negative
Sulphate Cyanobacteria;
Haloferax mediterranea

Negative

Table 1: Non-Carbohydrate Substituents of Exopolysaccharides

Microbial homopolysaccharides are mainly neutral glucans, although levans (fructans)
may also have potential uses. The majority of heteropolysaccharides are polyanionic
due to the presence of uronic acids; alternatively, or additionally, charge can be
conferred by the presence of organic or inorganic substituents. These include pyruvate
ketals or succinyl half-esters, phosphate and (relatively rarely) sulphate groups. Further
intriguing properties may be conferred by the presence of O-acetyl groups which are of
widespread occurrence. Almost all microbial heteropolysaccharides are composed of
regular repeating units varying in size from disaccharides to octasaccharides. These
frequently contain one mole of a uronic acid. D-glucuronic acid is most common but
some heteropolysaccharides contain D-galacturonic acid. D-Mannuronic acid is found
in bacterial alginates and a few other polysaccharides, as is L-guluronic acid. Rare
aminouronic acids are also found. Very occasionally, two uronic acids are present in a
repeat unit. The proposed uniformity of the repeat units is mainly based on chemical
studies and some irregularities may possibly be found, especially in polymers composed
of larger and more complex repeat units. Short side-chains, varying from one to four
sugars in length may be present. Bacterial alginates, linear molecules composed only of
uronic acids, lack any regular repeat unit. They are heteropolysaccharides composed of
D-mannuronic and L-guluronic acids in an irregular linear structure.

Various acyl groups and other substituents may be found in exopolysaccharides in
addition to a wide range of monosaccharides (Table 1). The most common substituents,
ester-linked O-acetyl groups, do not confer any charge on the macromolecule but may
greatly affect the physical characteristics of aqueous solutions. Pyruvate ketals are also
common and they confer a negative charge due to the free carboxylic acid group. Acetyl
and pyruvate groups are more common in heteropolysaccharide structures than
homopolysaccharides. Succinyl half esters are found in polymers from Rhizobium
species. Indeed, the succinoglycan EPS from Rhizobium and related spp. usually contain
all three acyl groups. Two types of inorganic substituents may be present. Phosphate
groups are relatively common in polysaccharides from Gram positive bacteria though
rare in Gram negative products; sulphate groups have been found in EPS of
Cyanobacteria and of halophiles including Haloferax mediterranea.

In electron micrographs, some EPS appear to form network-like structures; in others,
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macromolecular strands have been demonstrated. The EPS are usually highly hydrated.
EPS apparently serve to protect the bacterial cells against desiccation, phage and other
agents. Cellulomonas flavigena provides a rare example in which the EPS (curdlan) has
been shown to function as a carbon and energy reserve.

2.2. Occurrence and Composition

Three types of microbial homopolysaccharide structure have been characterised:

1. linear neutral polymers composed of a single linkage type (the "mixed linkage" type
of glucan found in cereal plants such as oats and barley have not been detected in
micro-organisms);
2. other homopolysaccharides are exemplified by the fungal polymer scleroglucan.
These are linear polysaccharides with regular, short side-chains composed on one or
two residues of the same monosaccharide as the main chain. Scleroglucan possesses
tetrasaccharide repeating units due to the 1,6--D-glucosyl side-chains present on
every third main chain residue.
3. The third type, including dextrans, are branched homopolysaccharides, as are levans
(polyfructans). A few polyanionic homopolymers are also known. They include
poly-D-glucuronic acid formed by a Rhizobium mutant, the 'Vi' antigen found in
some species of Enterobacteriaceae and some bacterial sialic acids. Also in this
category are the acetylated, poly-D-mannuronic acid polymers produced by some
Pseudomonas aeruginosa mutants .
-
-
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Encyclopedia of Life Support Systems (EOLSS)

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Biographical Sketch

Ian W. Sutherland is Professor of Microbial Physiology in the Division of Biological Sciences at
Edinburgh University.
He holds the degrees of B.Sc., Ph.D. and D.Sc. from Edinburgh University.
His major fields of interest are - the cell surface, its composition, structure and biosynthesis; the use of
enzyme methods in structural analysis of polysaccharides; biotechnological aspects of polysaccharides;
the physical properties of polysaccharides in aqueous solution. He has spent study leave at the Max
Planck Institut fr Immunbiologie, Freiburg, West Germany, and at J ohns Hopkins University (Baltimore,
U.S.A.).
He has served as - Member of Council - Society of General Microbiology 1984-93,
Chair of British Coordinating Committee for Biotechnology 1993-1996, an Editor of Journal of General
Microbiology 1985-1990 and MEB- Journal of Bacteriology 1980-1986; Journal of Microbiological
Methods 1983-1987, Carbohydrate Polymers 1980 onwards, Journal of Industrial Microbiology 1988-
1994, Journal of Marine Biotechnology 1999 onwards.
He is the author of 38 Book Chapters and Reviews and 118 Published scientific papers in refereed
J ournals.