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Article history:
Received 3 March 2014
Received in revised form 13 May 2014
Accepted 3 June 2014
Available online 26 June 2014
Keywords:
Polyhydroxyalkanoate
Biodegradable plastic
Polyhydroxybutyrate
Burkholderia xenovorans
pha gene
a b s t r a c t
Petroleum-based plastics constitute a major environmental problem due to their low biodegradability and accumulation in various environments. Therefore, searching for novel biodegradable plastics is
of increasing interest. Microbial polyesters known as polyhydroxyalkanoates (PHAs) are biodegradable
plastics. Life cycle assessment indicates that PHB is more benecial than petroleum-based plastics. In this
report, bacterial production of PHAs and their industrial applications are reviewed and the synthesis of
PHAs in Burkholderia xenovorans LB400 is described. PHAs are synthesized by a large number of microorganisms during unbalanced nutritional conditions. These polymers are accumulated as carbon and energy
reserve in discrete granules in the bacterial cytoplasm. 3-hydroxybutyrate and 3-hydroxyvalerate are two
main PHA units among 150 monomers that have been reported. B. xenovorans LB400 is a model bacterium
for the degradation of polychlorobiphenyls and a wide range of aromatic compounds. A bioinformatic
analysis of LB400 genome indicated the presence of pha genes encoding enzymes of pathways for PHA
synthesis. This study showed that B. xenovorans LB400 synthesize PHAs under nutrient limitation. Staining with Sudan Black B indicated the production of PHAs by B. xenovorans LB400 colonies. The PHAs
produced were characterized by GCMS. Diverse substrates for the production of PHAs in strain LB400
were analyzed.
2014 Elsevier B.V. All rights reserved.
1. Introduction
The highly versatile petroleum-based plastics have made them
valuable materials for modern life with a wide range of applications. However, due to their low biodegradability and accumulation
in various environmental compartments they are major environmental pollutants. Searching for novel biodegradable polymers
that possess similar physicochemical properties as conventional
plastics is of increasing interest. PHAs are biodegradable plastics
synthesized by diverse bacteria from a range of substrates including
sugars and fatty acids. These biopolymers are stored as intracellular
carbon and energy reserve granules by Gram-negative and Grampositive bacteria under nutrient limitation. 3-hydroxybutyrate
(3HB) and 3-hydroxyvalerate (3HV) are two common and widely
studied PHA monomers. However, 150 different PHA monomers
have been reported. Depending on the microorganism, the carbon source and culture conditions different PHA homopolymers,
copolymers and combination thereof are synthesized. A number of
Fig. 1. Plastic consumption in Chile in different areas. Adapted from Ref. [1].
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Table 1
Production of polyhydroxyalkanoates by diverse bacteria.
Bacterial strain
Carbon source
PHA
PHA content
(%)
References
Glucosa,
fructose, acetic
acid, valeric
acid
Sucrose or
gluconate
Xylose:levulinic
acidb
Sucrose:propionic
acidc
Glucose
PHB
PHV
80
90
[14,18]
PHB
P(3HPEa )
P(3HB-co-3HV)
45
[19,20]
49
[21,22]
P(3HB-co-3HV)
30
[23,24]
PHB
ND
[25,26]
Fig. 2. Structural formulas of PHAs. General PHA structural formula (a). R derived from radical represents different substituents. PHB structural formula (b). Structural formulas
of short-chain-length PHA monomers (c).
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Fig. 3. Biosynthesis of 3-hydroxybutyrate and 3-hydroxyvalerate from different substrates. 3HB is synthesized from sugars via the Entner-Doudoroff pathway and from few
fatty acids. 3HV is synthesized from diverse fatty acids. Abbreviations: 3-hydroxyvalerate (3HV), 3-hydroxybutyrate (3HB), phosphate (P); Coenzyme A (CoA) and 2-keto-3deoxy-6-phosphogluconate (KDPG).
For example, the hmgABC gene cluster encoding the homogentisate central pathway and the hpaG1G2EDFHI gene cluster encoding
the homoprotocatechuate central pathway are located at chromosome 1 (C1) and chromosome 2 (C2), respectively. Additional
hmg gene copies were identied within the LB400 genome [43,44].
The functionality of diverse catabolic pathways such as biphenyl,
homogentisate, homoprotocatechuate, gentisate, protocatechuate
and 2-aminophenol pathways has been reported [42,4446]. B. xenovorans LB400 is able to grow on 3-hydroxyphenylacetate (3-HPA)
Fig. 4. The pha gene cluster organization in B. xenovorans strain LB400 and R. eutropha strain H16. The arrows indicate the PHA metabolic pathway genes wherein the black
arrows indicate the PHA polymerase genes.
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Fig. 5. Synthesis of polyhdroxyalkanoates by B. xenovorans strain LB400. LB400 colonies grown on M9 medium with mannitol (10 g L1 ) as sole carbon source and 10% N (NH4 Cl
0.1 g L1 ) were stained with Sudan Black B dye (a). Magnication of the image (b).
[26]
[27]
[28]
[29]
[30]
213