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Biochem. J. (2009) 418, 219232 (Printed in Great Britain)
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doi:10.1042/BJ20081769
REVIEW ARTICLE
plastic production. Bacterial bioreactors fed with renewable resources from plants so-called white biotechnology have also
been successful in producing biodegradable polymers. In addition
to these methods of exploiting plant materials for biodegradable
polymer production, the present review also addresses the
advances in synthesizing novel polymers within transgenic plants,
especially those in the polyhydroxyalkanoate class. Although
there is a stigma associated with transgenic plants, especially
food crops, plant-based biodegradable polymers, produced as
value-added co-products, or, from marginal land (non-food), crops
such as switchgrass (Panicum virgatum L.), have the potential to
become viable alternatives to petroleum-based plastics and an
environmentally benign and carbon-neutral source of polymers.
INTRODUCTION
Abbreviations used: ADM, Archer Daniels Midland; ASTM, American Society for Testing and Materials; CaMV, cauliflower mosaic virus; CAP,
cellulose acetate phthalate; CEN, European Committee for Standardization; ER, endoplasmic reticulum; EU, European Union; GMO, genetically
modified organism; HB, hydroxybutyrate; HV, hydroxyvalerate; IPP, isopentenyl diphosphate; PA, protective antigen; PCLH, polycaprolactone
(polyhexanolactone)/hexamethylene di-isocyanate; PDC, pyruvate dehydrogenase complex; PLA, poly(lactic acid); PHA, polyhydroxyalkanoate; PHB,
polyhydroxybutyrate; PHBV, PHB-co-polyhydroxyvalerate; SPI, soy protein isolate; UTR, untranslated region.
1
email mooneyb@missouri.edu
c The Authors Journal compilation
c 2009 Biochemical Society
Biochemical Journal
University of Missouri, Interdisciplinary Plant Group, Division of Biochemistry, and Charles W. Gehrke Proteomics Center, 214 Christopher S. Bond Life Sciences Center,
1201 Rollins St, Columbia, MO 65211, U.S.A.
220
B. P. Mooney
WASTE MANAGEMENT
Figure 1
The United States, the European Union and countries in Asia have specific requirements for
biodegradable polymer testing and certification. Companies producing biodegradable polymers
can submit the products for independent testing and certification through the US Composting
Council or the Biodegradable Products Institute in New York, Vincotte in Brussels, DIN
(Deutsches Institut fur Normung) Certco in Berlin, the European Bioplastics Association and
the Biodegradable Plastic Society in Japan. Their labels are based on the ASTM 6400-99
and EN 13432 standards for biodegradable and compostable plastics. As examples, the Figure
shows logos issued by Vincotte in Brussels, the European Bioplastics Association and the
US Composting Council/the Biodegradable Products Institute, and are reproduced with their
permission.
Figure 2
(A) Natural rubber is a polymer of isoprene units arranged in cis -1,4 linkages. Natural rubber
(unlike synthetic) also has minor protein, lipid, carbohydrate and mineral components that
are responsible for its properties of resistance to abrasion and impact, elasticity, efficient heat
dispersal, and resilience and malleability at cold temperatures. (B) Gluten is a polymer of
glutenin that has inter- and intra-disulfide bonds that promote elasticity of the gluten polymer.
The gliadin proteins are thought to intercalate within the glutenin matrix and are held in place
by hydrophobic interactions and hydrogen bonding.
Plants naturally produce a number of structural and carbonreserve polymers. Polysaccharides are estimated to make up
approx. 70 % of all organic matter. Cellulose accounts for about
40 % of total organic matter and is the most abundant macromolecule on earth. Lignin comprises 1525 % of a typical woody
plant. Starch is also a major component of global biomass, and
the ability to digest starchy foods is thought to have played a role
in human evolution and its success over other species [17]. A
number of natural polymers have been exploited for commodity
manufacturing, and most of these products retain the inherent
biodegradability of their carbohydrate building blocks.
Rubber
221
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B. P. Mooney
for its Yulex Natural Rubber product based on guayule latex and
they plan to couple latex production with cellulosic biomass for
biofuels and power generation.
Russian dandelion produces a higher-molecular-mass polymer
than either Hevea or guayule (about 2 MDa), which accumulates
in lactifers in the roots. It was discovered in Kazakhstan in the
1930s and was used as a source of motor-tyre rubber during World
War II by a number of countries, including the United States and
the U.K. Although production yields are poor and large-scale
cultivation is hampered by cross-breeding with weedy species
and labour-intensive crop maintenance, it can be useful as a model
species for rubber biosynthesis because, in part, of its relatively
short life cycle (altered rubber phenotypes can be screened within
6 months) compared with guayule and Hevea. A further advantage
is the double-cropping potential of Russian dandelion, both as
a source of rubber and its fructose-based storage sugar, inulin,
which accumulates at 2540 % of root dry weight and could be
used in bioethanol production [26].
Proteins
Figure 3
223
Cellulose and starch can be modified to produce biodegradable polymers with various R groups in place of hydrogen. Nitrocellulose is cast as films/paper, and cellulose acetate and CAP have
been used for both films and nanospheres. Derivatization is achieved by chemical means from the isolated plant material. Starch has been used directly for polymer production and also through
derivatization with caprolactone, cellulose and various vinyl alcohols. The degradation properties of starch polymers are dependent on the relative proportion of pure starch to secondary compounds.
The degradation of the Mater-BiTM starch polymers is shown in the bottom left-hand panel.
224
B. P. Mooney
Starch
Using plant carbon reserves for industrial biotechnology feedstocks typically sugar, starch,
or oils various polymers can be produced by bacterial fermentation. (A) PLA is produced
by bacteria of the genus Lactobacillus , and often D-glucose (maize dextrose) is the carbon
source. It is a high-molecular-mass polymer (> 1 MDa) that can be used for films and moulded.
(B) PHB and (C) PHBV are in the class of PHAs that naturally occur in many bacteria as carbon
reserves. Typically, highly refined sources of glucose derived from plants are used in batch-fed
bacterial cultures; however, use of industrial wastewater and plant oils have also shown promise
for PHA production through fermentation.
225
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B. P. Mooney
A number of hurdles need to be tackled before commercialscale production of value-added heterologous proteins (and other
polymers) can be achieved in plants. The DNA encoding the
foreign protein must be stable across a number of generations,
and synthesis of the protein from mRNA, through processing,
translation and, addition of post-translation modifications, must
be considered. Additionally, the type of crop used and the site of
accumulation must be carefully considered to allow high levels
of expression and ease of purification (reviewed in [87]).
Among the number of novel proteins being expressed in
transgenic plants, many of these confer traits on the plant itself, for
example the Bacillus thuringiensis cry toxin for insect resistance
(see [88]). However, in the context of polymer production,
proteins with an inherent commercial value following purification
from the plant are also being produced. Typically, these are highvalue proteins with specialized uses, for example as vaccines.
Production of heterologous proteins within plants has numerous
advantages over production in animal or bacterial systems. Plantbased therapeutic proteins, for example, are theoretically safer
from a disease point of view than those purified from animal
sources (whether native or recombinant). Also, incorporation
of post-translational modifications such as glycosylation can
be achieved by in planta production, in contrast with bacterial
systems. Although eukaryotic culture systems such as insect and
mammalian cell cultures can be used in many cases to produce
viable proteins, an additional advantage of plants is that they are
renewable and scaleable and require less energy and nutrient input
than bioreactors, which should make plants cost-effective for bulk
applications (reviewed in [89]).
Heterologous protein accumulation within plants can be
achieved through standard transgenic approaches (i.e. nuclear
c The Authors Journal compilation
c 2009 Biochemical Society
227
PHAs
228
Figure 5
B. P. Mooney
Accumulation of PHBV within plastids uses the endogenous PDC for generation of the two
precursors of the polymer acetyl-CoA and propionyl-CoA. Introduction of the bacterial PHA
synthase operon comprising three enzymes (blue italics) allows accumulation of the polymer
within the plastid. The plastidial -oxobutyrate (-ketobutyrate) is produced from threonine by
an introduced threonine deaminase (not shown).
Figure 6
229
The present review discusses three main themes for synthesis of biodegradable polymers. There are a number of naturally occurring plant polymers that have been exploited by direct harvesting
for biodegradable plastics rubber, starch, proteins and cellulose sometimes followed by derivatization. Industrial or white biotechnology using plant extracts as carbon sources for bioreactor
fermentative production of PLA and PHA polymers is an alternative. Through engineering of bacteria, a number of valuable polymers can be produced with various monomers based on the nutrient
input. Finally, using transgenic technologies, plants can be engineered to produce novel polymers, including value-added proteins and PHAs, and therefore function as solar-driven biofactories. In
all cases, the polymers produced are biodegradable, so the carbon cycle is maintained the transgenic route to plastics is considered the closest to a carbon-neutral method for polymer production.
CONCLUDING REMARKS
230
B. P. Mooney
FUNDING
This research received no specific grant from any funding agency in the public, commercial
or not-for-profit sectors.
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