The Handbook of Polyhydroxyalkanoates Microbial
Biosynthesis and Feedstocks 1st Edition
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The Handbook of
Polyhydroxyalkanoates
The Handbook of
Polyhydroxyalkanoates
Microbial Biosynthesis and Feedstocks
Edited by
Martin Koller
First edition published 2020
by CRC Press
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and by CRC Press
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© 2021 Taylor & Francis Group, LLC
First edition published by CRC Press 2021
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ISBN: 9780367275594 (hbk)
ISBN: 9780429296611 (ebk)
Typeset in Times
by Deanta Global Publishing Services, Chennai, India
Dedicated to the fond memory of our father, Josef Koller
(1949–2019), who passed away during the creation of this book.
Contents
Foreword I.................................................................................................................xi
Foreword II ............................................................................................................ xiii
About the Editor.....................................................................................................xvii
Contributors ............................................................................................................xix
The Handbook of Polyhydroxyalkanoates, Volume 1: Introduction
by the Editor.........................................................................................................xxvii
PART I Enzymology/Metabolism/Genome
Aspects for Microbial PHA Biosynthesis
Chapter 1 Monomer-Supplying Enzymes for Polyhydroxyalkanoate
Biosynthesis..........................................................................................3
Maierwufu Mierzati and Takeharu Tsuge
Chapter 2 PHA Granule-Associated Proteins and Their Diverse Functions...... 35
Mariela P. Mezzina, Daniela S. Alvarez, and M. Julia Pettinari
Chapter 3 Genomics of PHA Synthesizing Bacteria .......................................... 51
Parveen K. Sharma, Jilagamazhi Fu, Nisha Mohanan, and
David B. Levin
Chapter 4 Molecular Basis of Medium-Chain Length-PHA Metabolism of
Pseudomonas putida .......................................................................... 89
Maria-T. Manoli, Natalia Tarazona, Aranzazu Mato, Beatriz
Maestro, Jesús M. Sanz, Juan Nogales, and M. Auxiliadora Prieto
Chapter 5 Production of Polyhydroxyalkanoates by Paraburkholderia
and Burkholderia Species: A Journey from the Genes through
Metabolic Routes to Their Biotechnological Applications .............. 115
Natalia Alvarez-Santullano, Pamela Villegas, Mario
Sepúlveda, Ariel Vilchez, Raúl Donoso, Danilo Pérez-Pantoja,
Rodrigo Navia, Francisca Acevedo, and Michael Seeger
vii
viii Contents
Chapter 6 Genetic Engineering as a Tool for Enhanced PHA Biosynthesis
from Inexpensive Substrates............................................................. 145
Lorenzo Favaro, Tiziano Cazzorla, Marina Basaglia, and
Sergio Casella
Chapter 7 Biosynthesis and Sequence Control of scl-PHA and mcl-PHA ....... 167
Camila Utsunomia, Nils Hanik, and Manfred Zinn
PART II Feedstocks
Chapter 8 Inexpensive and Waste Raw Materials for PHA Production ........... 203
Sebastian L. Riedel and Christopher J. Brigham
Chapter 9 The Sustainable Production of Polyhydroxyalkanoates from
Crude Glycerol ................................................................................. 221
Neha Rani Bhagat, Preeti Kumari, Arup Giri, and Geeta Gahlawat
Chapter 10 Biosynthesis of Polyhydroxyalkanoates (PHA) from Vegetable
Oils and Their By-Products by Wild-Type and Recombinant
Microbes........................................................................................... 257
Manoj Lakshmanan, Idris Zainab-L, Jiun Yee Chee, and
Kumar Sudesh
Chapter 11 Production and Modifcation of PHA Polymers Produced from
Long-Chain Fatty Acids................................................................... 291
Christopher Dartiailh, Nazim Cicek, John L. Sorensen, and
David B. Levin
Chapter 12 Converting Petrochemical Plastic to Biodegradable Plastic ............ 315
Tanja Narancic, Nick Wierckx, Si Liu, and Kevin E. O’Connor
Chapter 13 Comparing Heterotrophic with Phototrophic PHA Production:
Concurring or Complementing Strategies? ...................................... 331
Ines Fritz, Katharina Meixner, Markus Neureiter, and
Bernhard Drosg
Contents ix
Chapter 14 Coupling Biogas with PHA Biosynthesis......................................... 357
Yadira Rodríguez, Victor Pérez, Juan Carlos López, Sergio
Bordel, Paulo Igor Firmino, Raquel Lebrero, and Raúl Muñoz
Chapter 15 Syngas as a Sustainable Carbon Source for PHA Production ......... 377
Véronique Amstutz and Manfred Zinn
Index...................................................................................................................... 417
Foreword I
Even in technologically advanced countries, relatively little of the massive amount
of plastic produced each day is recycled. The vast majority of plastic placed in
recycling bins is sent to landfll. We know about the harm to aquatic life and other
detrimental effects of plastics that do not degrade in the natural environment.
Polyhydroxyalkanoates (PHA) are not only biodegradable in most natural environ-
ments but are derived from renewable resources. Yet most people have never seen
any products containing PHA because there are so few. The reason is the high cost of
production. Without some sort of government subsidy, factors like substrate, fermen-
tation, and separation costs prevent PHA from competing with conventional plastics.
We need to produce better and cheaper PHA.
This collection of works is targeted primarily at providing researchers with state-
of-the-art PHA technology required to address these challenges. Progress occurs
more rapidly when the necessary information is available. Without adequate knowl-
edge, we may stray down futile paths or draw incorrect conclusions. Years ago, we
were given a bottle of toxic black smelly liquid called “nonvolatile residue” (NVR),
the major waste stream from the manufacture of nylon 6’6’ containing a mixture of
~100 different mono- and dicarboxylic acids. The objective was to use microorgan-
isms to convert this waste material into something useful. After months of research,
it was determined that NVR could be used to produce poly(3-hydroxybutyrate)
(PHB) under chemostat and fed-batch conditions. This was wonderful, the conver-
sion of a plastics manufacturing waste material into another plastic, PHB. Perfect,
and who else could possibly think of such a weird idea? It turned out that researchers
in Graz, Austria (the birthplace of this current collection of works) had arrived at
exactly the same conclusion and that we were all wrong. At this point in time, the
standard method for PHB analysis, also developed in Graz, was gas chromatography
fame-ionization detection (GC-FID), but this method depends on the use of stan-
dards to establish the retention times of the target compounds. We ignored all of the
other peaks since, to our knowledge, PHB was the only PHA in existence. Had mod-
ern rapid publication methods summarized the information available at that time,
we would have been well aware that the use of NVR would lead to the formation of
PHA copolymers.
Why should anyone care about the conversion of NVR into PHA? The frst sec-
tion in the current treatise deals with such issues. While it may be diffcult for us in
the Americas to comprehend, there is a shortage of farmland in much of the world.
If one of the great attributes of PHA is that their production substrates are typi-
cally renewable, we must consider the effect of their large-scale production on food
availability and avoid competition with agricultural materials destined for human
nutrition. Speaking of competition with foodstuffs, there has been some effort to
produce PHA in the plastids of various plants. Researchers trying to clone the genes
for medium-chain-length PHA (mcl-PHA) synthesis into plants were stymied by a
missing link. When the plant produces mcl-carboxylic acids by de novo synthesis,
they are covalently linked to the acyl carrier protein (ACP), while they must be
xi
xii Foreword I
linked to coenzyme A (CoA) for PHA synthesis. Since some organisms can naturally
produce mcl-PHA through de novo fatty acid synthesis, there must be an enzyme
that can transfer the carboxylic acids to coenzyme A while still medium chain
length. The discovery of phaG coding for the PhaG transacylase fnally allowed
progress to continue and demonstrates the importance of mapping key enzymes in
a pathway and knowing the genes that code for them. Before developing a fermenta-
tion process for a specifc PHA, it is imperative that one has some knowledge of key
enzymes, their specifcity, and how their activity is controlled. This knowledge is
usually required when modeling the process as well. The frst half of this Handbook
deals with “Enzymology, Microbiology and Genetics,” information that is, in com-
bination with kinetic aspects dealt with in volume 2, required to model and or even
plan a PHA production process.
Why do microorganisms accumulate PHA? Typical answers include that they
serve as carbon and energy storage and/or that their production and degradation can
be used by microbes to control their internal redox potential. Since there are dif-
ferent classes of PHA and many diverse organisms that synthesize them, there are
likely specifc reasons for different materials and microbes. Nevertheless, “stress”
conditions are often cited as stimulating synthesis and stress-synthesis relationships
are explored in several chapters of volume 2 dealing with “Environmental and Stress
Factors.” Stress conditions may also be used to prevent the growth of undesired
microbes. Growing PHA-accumulating halophiles under osmotic conditions where
few other microorganisms survive may eliminate the need for costly sterilization
(one of the challenges facing the use of waste substrates that may resist sterilization)
or perhaps fnally allowing PHA to be produced in continuous culture over extended
periods of time, greatly decreasing capital cost. Cost is presently limiting the wide-
spread adoption of PHA as commodity plastics. The topics covered in the Handbook
allow us to examine the latest approaches to better understand the processes that
may allow economic PHA production, and the world should be a better place for it.
Bruce Ramsay and Juliana Ramsay
Foreword II
The Handbook of Polyhydroxyalkanoates comprises, in total, 42 chapters, which
have been divided into three volumes. Volume 1 at hand focuses on various aspects
of the production of polyhydroxyalkanoates (PHA) regarding suitable technical
substrates, production organisms, and the cellular PHA biosynthesis machinery,
whereas volumes 2 and 3 focus on the properties and downstream processes of PHA
production although also substrates and mixed cultures for PHA production are sub-
ject of the second volume. A clear thematic cut between the topics of all volumes was
not reached but is also diffcult to reach. The three volumes are edited by Dr. Martin
Koller from the University of Graz, who is very well known for his contributions to
the feld.
There are several chapters contributed to volume 1 by renowned scientists, which
focus on the feedstocks that can be used for PHA production. A good overview
and introduction are provided in Chapter 8, pledging for inexpensive and waste
raw materials instead of the more expensive refned carbon sources. Other chapters
review the use of long-chain fatty acids, vegetable oils, and byproducts, glycerol,
CO2, biogas, and syngas. Even the use of conventional plastics as a carbon source
for PHA production is described. This means that conventional petrochemical plas-
tics are converted into biodegradable plastics! One chapter describes how bacteria
can be genetically engineered to produce PHA from inexpensive substrates. Another
focus is the different organisms that can be used for PHA production. Phototrophic
organisms like cyanobacteria will allow the production of PHA from CO2 and sun-
light. Haloarchaea and halophilic eubacteria will allow PHA production at high
salt concentrations in the medium, which is highlighted in volume 2. Furthermore,
PHA production by hydrogen-oxidizing bacteria, Pseudomonas putida, as well as
Burkholderia and related bacteria is described in this Handbook. In addition, an in
silico analysis of the genomes of PHA-synthesizing bacteria is provided in one chap-
ter. Thermodynamic aspects and the mathematical modeling of PHA biosynthesis
were also analyzed in two chapters at the beginning of volume 2. The relation of
PHA metabolism and stress robustness, the analysis of the composition and structure
of PHA granules, aspects of PHA degradation, and the control of PHA composition
are subjects of other intriguing chapters.
We have to consider that intensive research on PHA has been conducted for
more than 40 years. This research aimed at unraveling all biochemical factors that
infuence the biosynthesis and production of PHA. This aim has more or less been
achieved. If one considers that about 40 years ago, not even the genes encoding the
enzymes involved in PHA biosynthesis were known and only vague imaginations on
the enzymes catalyzing the polymerization were present, we now know many details
and much more than was expected. These details include the elements constituting
the PHA granules and the regulation of PHA biosynthesis in the bacterial cells. This
is fne.
The research on PHA was also aiming at fnding conditions at which PHA could
be commercially produced under economically reliable conditions. This has not at
xiii
