181
Production of new polymeric compounds in plants
Yves Poirier
Plant metabolic engineering has recently enabled the
synthesis of a range of polyhydroxyalkanoates as well as a
protein-based polymer. These novel compounds can be
harvested from plants as a renewable source of
environmentally friendly polymers or can be used to change
the physical properties of plant products, such as fibres.
Addresses
Institut dcologie-Biologie et Physiologie Vgtales, Universit de
Lausanne, CH-1015 Lausanne, Switzerland;
e-mail: yves.poirier@ie-bpv.unil.ch
Current Opinion in Biotechnology 1999, 10:181185
http://biomednet.com/elecref/0958166901000181
Elsevier Science Ltd ISSN 0958-1669
Abbreviations
dwt dry weight
MCL medium-chain length
PHA polyhydroxyalkanoate
PHB polyhydroxybutyrate
P(HB-HV) poly(hydroxybutyrate-hydroxyvalerate)
Introduction
Over the past 50 years, the development of synthetic
petroleum-based polymers has considerably reduced mans
dependence on the use of certain plant polymers, such as
wood, cotton and rubber. Advances in plant genetic engi-
neering, combined with the growing concerns about our
environment and the decreasing petroleum reserves, have
created new opportunities to use plants for the large-scale
production of novel renewable and environmentally
friendly polymers. This article will focus on the two novel
classes of polymers that have been produced in transgenic
plants, namely polyhydroxyalkanoates (PHAs) and pro-
tein-based polymers made from artificial genes.
Polyhydroxyalkanoate homopolymer
PHAs are polyesters of 3-hydroxyacids which have proper-
ties of biodegradable plastics and elastic polymers
(elastomers) [13]. PHAs are naturally synthesised by a large
variety of bacteria as a source of carbon and an energy
reserve. Numerous bacteria and some fungi have been
found to degrade extracellular PHAs and metabolise the
breakdown products to carbon dioxide and water [4,5].
Thus, products made out of PHAs can be degraded com-
pletely and rapidly by composting (typically 39 months).
Over 100 different hydroxyacids have been found to be
included in PHAs [6]. This large diversity in structure trans-
lates into a spectrum of physical properties, ranging from a
stiff and brittle plastic to soft plastics, elastomers, rubbers
and glues. The main limitation in using bacterial PHAs as a
source of biodegradable polymer for low-value commodity
products is their production cost, making them 510 times
more expensive than petroleum-based polymers such as
polypropylene [7]. It is in this context that agricultural crops
have been regarded as a promising alternative for the pro-
duction of PHAs on a large scale and at low cost [1,8].
Production of PHAs by bacterial fermentation will probably
remain the method of choice for low-volume high-value
applications, such as in the medical field.
The first PHA produced in plants was the homopolymer
polyhydroxybutyrate (PHB). PHB is synthesised from
acetyl-CoA, which is found in several subcellular com-
partments in plant cells, including cytoplasm, plastids
and peroxisomes.
Expression of the acetoacetyl-CoA reductase and PHA
synthase from the bacterium Ralstonia eutropha (formerly
Alcaligenes eutrophus) in the cytoplasm of Arabidopsis
thaliana plant cells lead to the production of PHB at levels
up to 0.10.2% of the shoot dry weight (dwt) [9,10]
(Figure 1). Growth of these transgenic plants was severely
reduced, presumably due to the depletion of the acetyl-
CoA pool available in the cytoplasm for isoprenoid and
flavonoid biosynthesis [11].
Expression of the PHB biosynthetic pathway in the plastids
of transgenic A. thaliana increased PHB production in shoots
up to 14% dwt with minimal effects on plant growth [12]
(Figure 1) This experiment demonstrated that sufficient
acetyl-CoA could be diverted from the plastid fatty acid
biosynthetic pathway for high-level PHB production. In an
effort to bring this technology to the field, Monsanto has
recently shown that transgenic rape expressing the PHB
pathway in leucoplast of developing embryos accumulated
PHB at levels up to 8% dwt in mature seeds of heterozygous
plants (K Gruys, personal communication). The effect of
PHB synthesis on triacylglycerides accumulation has not
been reported. It is possible that a diversion of acetyl-CoA
towards PHB in seeds may result in a decrease in
triacylglyceride accumulation. Although it is unlikely that a
moderate decrease in triacylglycerides would be problematic
for the viability or vigour of the seedling, it would have an
impact on the value of the crop as an oil producer.
PHB accumulation up to 2 mg/g dwt has recently been
demonstrated in transgenic Black Mexican sweet corn
suspension cell cultures expressing the PHB biosynthet-
ic pathway in the peroxisomes (Figure 1; F Srienc,
personal communication). In this system, -oxidation of
fatty acids is the source of acetyl-CoA and 3-hydroxybu-
tyryl-CoA used for PHB biosynthesis.
PHB synthesis in plants has thus been demonstrated in
three distinct subcellular compartments through metabol-
ic engineering of three distinct pathways, namely
isoprenoid biosynthesis in the cytoplasm, fatty acid
182 Plant biotechnology

Figure 1

Plastid

Threonine Propionyl-CoA Acetyl-CoA

PDC
(ilvA)
(bktBRe ) (phaARe)
2-ketobutyrate
3-ketovaleryl-CoA Acetoacetyl-CoA Fatty acid
biosynthesis
(phaBRe) (phaBRe)

R-3-hydroxyvaleryl-CoA R-3-hydroxybutyryl-CoA
(phaCRe) (phaCRe) (FatB3)

Isoleucine P(HB-HV) PHB

Long-chain Medium-chain
fatty acids fatty acids

Cytoplasm Peroxisome

Acetyl-CoA Acyl-CoA
Acetyl-CoA

(phaARe) 3-ketoacyl-CoA 2-trans-enoyl-CoA

Acetoacetyl-CoA Malonyl-CoA
acetoacetyl-CoA
(phaBRe) S-3OH-acyl-CoA
(phaBRe)
R-3-hydroxybutyryl-CoA
R-3-hydroxy- R-3OH-acyl-CoA
butyryl-CoA (phaCRe)
(phaCRe) (phaC1Pa )
(phaCRe) PHB
MCL-PHA
Isoprenoids PHB Flavonoids

Current Opinion in Biotechnology

Modification of plant metabolic pathways for the synthesis of PHAs.
The pathways created or enhanced by the expression of transgenes
are highlighted in bold letters and solid arrows, whereas endogenous
plant pathways are in plain letters and open arrows. The various
transgenes expressed in plants are indicated in parenthesis. The ilvA
gene encodes a threonine deaminase from E. coli. The phaARe,
phaBRe and phaCRe genes encode a 3-ketothiolase, an acetoacetyl-
CoA reductase and a PHA synthase from R. eutropha, respectively.
The PHA synthase from R. eutropha can polymerise 3-hydroxyacyl-
CoAs ranging from 35 carbons. The bktBRe gene encodes a second
3-ketothiolase isolated from R. eutropha which shows high affinity for
both propionyl-CoA and acetyl-CoA [17]. The phaC1Pa gene encodes
a PHA synthase from P. aeruginosa which can polymerise 3-
hydroxyacyl-CoAs ranging from 616 carbons. The FatB3 gene
encodes a capryl-ACP thioesterase from C. lanceolata. PDC refers to
the endogenous plant pyruvate dehydrogenase complex.

biosynthesis in the plastid and -oxidation in the peroxi-
some. PHB biosynthesis has also been demonstrated in
the cytoplasm of yeast [13] and insect cells [14].
Polyhydroxyalkanoate copolymers
PHB is a polymer with relatively poor physical properties,
being too stiff and brittle for use in most commodity prod-
ucts. Thus, success in PHA production in crops depends
on the synthesis of polymers that have better physical
properties. Incorporation into the polymer of a low amount
of longer chain monomers decreases the cristallinity of the
polymer resulting in increased toughness and flexibility
[15]. A bacterial PHA with such properties is the copoly-
mer poly(hydroxybutyrate-hydroxyvalerate) (P[HB-HV]),
which is commercially produced by Monsanto under the
trade name Biopol.
A significant advance has been accomplished recently by
the synthesis of P(HB-HV) copolymer in A. thaliana and
rapeseed (K Gruys, personal communication). In the com-
mercial production of P(HB-HV) from R. eutropha,
propionate is added to the growth media in order to create
an intracellular pool of propionyl-CoA which can be con-
densed with acetyl-CoA to form 3-ketovaleryl-CoA. For
the synthesis of P(HB-HV) in plants, the propionyl-CoA
has been generated by modifying the branched amino acid
biosynthetic pathway (Figure 1). An Escherichia coli threo-
nine deaminase modified for plastid targeting was

overexpressed, leading to the accumulation of -ketobu-
tyrate, which can be converted, albeit at low efficiency, to
propionyl-CoA by the plant pyruvate dehydrogenase com-
plex [16]. A new 3-ketothiolase, encoded by the bktB gene
isolated from R. eutropha and showing high affinity for both
propionyl-CoA and acetyl-CoA [17], was also targeted to
the plastids along with an acetoacetyl-CoA reductase and a
PHA synthase. Expression of all four genes in A. thaliana
under the control of the CaMV35S promoter resulted in
the accumulation of PHA between 0.20.8% dwt in shoots,
with a 417 mol% hydroxyvalerate content in the polymer.
Seed-specific expression of the same pathway in leucoplast
of rape embryos resulted in PHA accumulation up to 2.3%
dwt in seeds, with up to 6.5 mol% hydroxyvalerate content
(K Gruys, personal communication). Although the present
level of P(HB-HV) copolymer production is lower than for
PHB homopolymer, synthesis of Biopol in plants is like-
ly to be a key factor on the route to commercialisation of
PHA-producing crops, as P(HB-HV) has good physical
properties and a market for this product has already been
developed. The challenge in this particular system is to
appropriately control the flux of two independent path-
ways supplying different monomers for the synthesis of a
single copolymer and to take into account the fact that
variable environmental conditions encountered in the field
may differentially affect the supply of one monomer.
Medium-chain length (MCL) PHAs represent a broad
family of PHAs containing 3-hydroxyacids ranging from six
to 16 carbons, which may have functional groups, such as
unsaturated bonds, epoxy and hydroxy groups [6]. The
presence of such reactive groups offers the opportunity to
modify the structure and physical properties of the PHAs
after extraction [18]. MCL-PHAs are generally attractive
as a source of elastomers, rubbers and glues.
Synthesis of MCL-PHAs in plants has recently been
demonstrated in A. thaliana expressing a PHA synthase from
Pseudomonas aeruginosa modified for peroxisome targeting
[19,20]. In this system, MCL-PHAs are synthesised using
the 3-hydroxyacyl-CoA intermediates generated by fatty
acid -oxidation (Figure 1). MCL-PHA production in per-
oxisomes was low, reaching 0.4% dwt in seven-day-old
germinating seedlings, and the monomer composition was
complex, including saturated and unsaturated monomers
ranging from six to 16 carbons. Growth of these transgenic
plants in media containing various unusual fatty acids result-
ed in an increased accumulation of MCL-PHA containing
monomers derived from the -oxidation of these fatty acids
(Y Poirier and V Mittendorf, unpublished data). For exam-
ple, addition of either tridecanoic acid, tridecenoic acid
(C13:1, 12) or 8-methyl-nonanoic acid resulted in the pro-
duction of PHA containing mainly saturated odd-chain,
unsaturated odd-chain or branched-chain 3-hydroxyacid
monomers, respectively. These studies demonstrated that
the plant -oxidation pathway was capable of generating a
large spectrum of monomers from fatty acids that can be
included in MCL-PHAs. An endogenous control over the
Production of new polymeric compounds in plants Poirier 183
flux of substrates towards -oxidation and PHA synthesis
has been achieved through the combined expression of a
peroxisomal PHA synthase with an acyl-acyl carrier protein
thioesterase (Y Poirier and V Mittendorf, unpublished data).
Constitutive expression of a plastidial capryl-acyl carrier pro-
tein thioesterase from Cuphea lanceolata in A. thaliana was
used to channel decanoic acids towards peroxisomal -oxi-
dation, resulting in an increased synthesis of MCL-PHA
composed of approximately 40% 3-hydroxydecanoic acid,
32% hydroxyoctanoic acid and 4% hydroxyhexanoic acid. A
working model thus emerges where enzymes and genes
involved in the synthesis of unusual fatty acids in plants can
be used to modulate the quantity and quality of substrates
channelled towards MCL-PHAs. The absolute amount of
MCL-PHA accumulating in mature shoots of transgenic
plants expressing both thioesterase and PHA synthase
remains relatively low (0.1% dwt) compared to PHB syn-
thesis in peroxisomes or plastids. The reasons for this
difference could be multiple, including differential activity
of bacterial enzymes in various plant subcellular compart-
ments and the relative ability of the created and endogenous
metabolic pathways to compete for the same substrates
(competition for acetyl-CoA for lipid and PHB biosynthesis;
competition for 3-hydroxyacyl-CoAs for fatty acid -oxida-
tion and MCL-PHA biosynthesis).
Novel uses of polyhydroxyalkanoates in plants
A novel perspective was brought by the expression of the
PHB biosynthetic pathway in the cytoplasm of cotton fibre
cells [21,22]. In this system, the polymer is produced in
plants to change the physical properties of the fibre and not
as a source of extractable biopolymer for industrial uses.
Accumulation of PHB to only 0.3% dwt of the fibre was suf-
ficient to significantly decrease the rate of heat uptake and
cooling of the fibre, resulting in a higher heat capacity and
improving the fibres insulating properties. It is intriguing
to speculate what the changes in fibre properties would be
if higher amounts of PHB could be synthesised through
expression of the pathway in the plastid, or if different PHA
copolymers, including MCL-PHAs, could be accumulated.
It is also tempting to speculate whether the properties of
wood, rubber or starch could also be positively modified
through the co-accumulation of PHAs.
Synthesis of PHAs in plants can also be used as a unique
tool in basic studies of plant biochemistry. PHA synthe-
sised in plants acts as a terminal carbon sink, as plants do
not have enzymes, such as PHA depolymerases, required
for degradation of the polymer. Thus, any modifications in
plant metabolism that can result in either changing the
quantity or the quality of substrates for the PHA synthase
could be detected by monitoring the quantity and compo-
sition of PHA. For example, PHB synthesis could be used
to study how carbon flux to endogenous compounds also
utilising acetyl-CoA, such as lipids and isoprenoids, can be
modulated to impact acetyl-CoA availability for PHB
biosynthesis. In the peroxisomes, MCL-PHAs are
synthesised from monomers derived from -oxidation
184 Plant biotechnology
intermediates of fatty acid degradation. The nature and
proportion of the monomers found in MCL-PHA can,
therefore, be used to elucidate the pathways involved in
the degradation of unsaturated and unusual fatty acids,
while the quantity of PHA can be used to study flux of
fatty acids towards peroxisomal -oxidation.
Protein-based polymers
Nature produces a range of proteins with unique properties.
For example, spiders produce a variety of silks that range
from lycra-like elastic fibres to kevlar-like superfibres [23],
whereas mussels can produce strong protein-based bioadhe-
sives that are effective in aqueous environments [24]. These
proteins, as well as elastin, collagen and keratin, are distinc-
tive because of their highly repetitive primary structure,
being composed of repeated blocks of amino acids.
Expression in E. coli of a spider dragline silk-like protein
based on a synthetic gene has led to production of approxi-
mately 300 mg/L of protein [25]. The protein produced was,
however, heterogeneous in size, due to premature termina-
tion of translation. Expression of a similar gene construct in
Pichia pastoris resulted in a higher level of accumulation of
protein (900 mg/L) without evidence of truncated protein
synthesis [26]. These studies opened the door to the pro-
duction of a range of protein-based polymers where primary
structure and molecular weight can be genetically controlled.
In this context, could plants be used as efficient production
vectors of interesting protein-based polymers?
Expression of a synthetic protein made of 121 repeats of
the peptide GlyValGlyValPro has been reported in
both E. coli [27], Aspergillus nidulans [28] and tobacco [29].
This peptide was based on the sequence of the bovine
elastin. Synthetic proteins made from multiple repeats of
this sequence display elastic properties, are biodegradable
as well as biocompatible, being non-toxic and naturally
resorbed by animal tissues [30]. The natural degradability
of these protein polymers in animals could be used for the
gradual release of drugs enclosed in a protein matrix.
While expression of (GlyValGlyValPro)121 in E. coli
leads to high polymer accumulation, expression of the
same gene in A. nidulans and tobacco led only to relatively
low protein accumulation despite high levels of mRNA. It
is probable that efficient expression of protein polymers
having a high proportion of a limited number of amino
acids will require modification in the amino acid biosyn-
thetic pathways and/or tRNA pools. The future of
producing protein-based polymers in plants as a source of
useful commodity products is at present uncertain in view
of the low level of protein accumulation in tobacco
(< 5 g per gram fresh weight) [29]. As with PHAs, howev-
er, it is perhaps possible to use protein-based polymers to
modify the properties of plant fibres, such as strength,
water absorption or dye-binding properties of cotton fibres.
Conclusions
A broad range of PHAs covering a spectrum of physical prop-
erties have now been synthesised in plants. The challenge
for the future is to succeed in high level production (i.e.
15% dwt) of a limited number of useful PHAs and in
developing efficient extraction procedures. Initial experi-
ments with expression of protein-based polymers in plants
indicate that, similar to PHAs, complex engineering of
multiple pathways is also likely to be necessary before
high-level accumulation of polymer is accomplished. For
both PHAs and protein-based polymers, production of
polymers for the modification of the properties of plant
products, such as fibres, are attractive alternatives to
extraction of polymers for industrial uses. PHAs and pro-
tein-based polymers are only two out of many groups of
polymers produced in Nature which have interesting prop-
erties. Plants alone produce several kinds of
carbohydrate-based polymers, such as cellulose and starch,
which can be modified and used for the synthesis of
biodegradable plastics [8]. Thus, novel industrially useful
polymers could also be developed in plants through modi-
fications of existing plant compounds.
Acknowledgements
I thank Daniela Caldelari, Philippe Reymond and Christiane Nawrath for
critical evaluation of the manuscript, and Marie-Madeleine Defago for
editorial assistance.
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