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uel ethanol is currently made by large-scale yeast

fermentation of sugars that are extracted or prepared
from crops followed by separation of the ethanol
by distillation
. The first major fuel-ethanol programme
(ProAlcool) started in Brazil in 1975
, followed by pro-
grammes in the USA in 1978 and, more recently, in
Canada (Table 1). Although many other countries pro-
duce ethanol for fuel and other purposes (http://www., major production has only occurred
in those countries with especially favourable agricul-
tural and economic conditions. This article will examine
current production, put the work into the context of
changing environmental and economic realities and
describe the latest technological developments.
Sugar extraction from sugar cane
Sugar cane (Saccharum officinarum) is a drought-tolerant,
tropical and subtropical crop that contains 1217% total
sugars on a wet-weight basis with 6872% moisture
(90% sucrose and 10% glucose or fructose). The aver-
age extraction efficiency to produce cane juice by
crushing is approximately 95% and the remaining solid
residue is cane fibre (bagasse).
In factories that only produce ethanol, the cane juice
is heated up to 110C to reduce microbial contami-
nation, decanted, sometimes concentrated by evaporation
and then fermented. In combined sugarethanol plants
(annexed distilleries), sucrose crystals that are formed
after cane-juice concentration are removed by centrifu-
gation, leaving a syrup (molasses) that contains up to
65% w/w sugars. Both sugar-cane juice and molasses
(after adjusting the sugar concentration) normally
contain sufficient minerals and organic nutrients to
be immediately suitable for ethanol production by
fermentation with Saccharomyces cerevisiae
Starch extraction and saccharification from grains
Maize (Zea mays) and other less commonly used
grains can be processed by wet or dry milling. Wet
milling was originally devised for the starch industry
and adapted for fuel-ethanol production. The grain is
soaked (steeped) in water with sulphur dioxide for up
to 40 h, followed by grinding and separation of starch
and co-products (Table 2). When wheat is used, the
valuable bran and germ are usually removed first by dry
processing in a flour mill before steeping in water. The
starch fraction is gelatinized by cooking at low and high
temperatures, followed by the addition of -amylase,
which yields dextrin oligosaccharides. In the final sac-
charification process, glucoamylase converts the starch
to glucose, which can eventually be fermented to ethanol.
The main difference with dry milling is that the
entire grain is milled to a median diameter of approxi-
mately 1 mm and the different components of the
cereal grain are not fractionated before the water and
enzyme are added; the slurry is processed as in wet
milling. In both systems, the sugar-containing juice
that leaves the processor (mash or wort) is essentially
sterile, and this is a crucial point in the subsequent
successful downstream processing.
Fermentation and distillation
In Brazil, 70% of the sugar-cane distilleries use a
batch process with a fermentation capacity of up to
1.5 million (l ethanol) d
; a continuous version of
the same process is also being used in approximately
350 distilleries, the latest of which was built in 1995.
In both processes, yeasts are separated from the fer-
mented medium by centrifugation and reused in sub-
sequent fermentations after dilute-sulphuric-acid
washing to reduce bacterial contamination. Very high
cell densities (817% v/v) and temperatures of 3335C
482 0167-7799/99/$ see front matter 1999 Elsevier Science Ltd. All rights reserved. PII: S0167-7799(99)01384-0 TIBTECH DECEMBER 1999 (VOL 17)
Fuel ethanol after 25 years
Alan E. Wheals, Luiz C. Basso, Denise M. G. Alves and Henrique V. Amorim
After 25 years, Brazil and North America are still the only two regions that produce large quantities of fuel ethanol, from sugar
cane and maize, respectively. The efficiency of ethanol production has steadily increased and valuable co-products are
produced, but only tax credits make fuel ethanol commercially viable because oil prices are at an all-time low. The original
motivation for fuel-ethanol production was to become more independent of oil imports; now, the emphasis is on its use as an
oxygenated gasoline additive. There will only be sufficient, low-cost ethanol if lignocellulose feedstock is also used.
A. E. Wheals ( is at the Department of Biology
and Biochemistry, University of Bath, Bath, UK BA2 7AY and also
at the Department of Biology, Federal University of Lavras, 37200-000,
Lavras, MG, Brazil. L. C. Basso, D. M. G. Alves and H. V. Amorim
are at the Department of Biological Sciences, ESALQ/University of
So Paulo, 13418-900 Piracicaba, SP, Brazil. D. M. G. Alves
and H.V. Amorim are also at Fermentec, Rua Treze de Maio, 768,
13400-900 Piracicaba, SP, Brazil.
Table 1. Major fuel-ethanol producers
Country Crop Fuel-ethanol
Brazil Sugar-cane (juice and 10.5 hydrous
molasses) 6.5 anhydrous
(in 1999)
USA Maize (95%) plus 5.3 anhydrous
some wheat and (in 1998)
Canada Maize plus 15% 0.24 anhydrous
wheat (in 1998)
TTEC Dec 99 24/10/99 9:51 am Page 482
contribute to reduced yeast growth and high ethanol
concentrations (811% v/v) and a high ethanol yield
(9092%). The very short (610 h) fermentation times
allow yeast to be recycled up to three times a day for as
long as 200 days. Deleterious bacterial contamination
can occur but is controlled using antibiotics. Some
more advanced distilleries are completely automated
There is a wide variation in the technology used in
maize-fermentation plants in North America. Distil-
leries that use older batch-culture technology do not
recycle yeast cells. Fresh yeast cells can be added every
time and are either bought from a supplier or produced
on site from a propagation unit. More-modern plants
have extensive computerized control and clean-in-
place technology, which is low on labour costs and
reduces equipment down time. Continuous input and
output processes are used together with simultaneous
saccharification with fermentation (SSF). This can even
be combined with yeast propagation in a continuous
process, with antibiotics often added as a precaution
against bacterial contamination.
Finally, the fermented liquor is distilled, producing
a constant-boiling (azeotropic) mixture of hydrous
ethanol at 95.5% v/v. To produce anhydrous ethanol,
the water is removed with dehydrated carbohydrate or,
more recently, with an appropriately sized molecular
sieve to which the water adsorbs. Distillation leaves
behind an aqueous residue called stillage (or vinasse in
Brazil) 1015 times the volume of the ethanol pro-
duced. In sugar-cane distilleries, all vinasse that is pro-
duced is used as fertilizer and in irrigation. In maize
wet mills some of this, called backset, can be used to
dilute the maize wort; in maize dry mills, the residual
non-starch fraction of whole stillage can be separated
by centrifugation or screening into approximately equal
parts of distillers grains (containing protein, fibre and
fat) and thin stillage (distillers solubles) that contains
yeast cells, soluble nutrients and small maize particles.
When dried in evaporators, these produce distillers
dried grains and distillers dried solubles, respectively,
or a combined product called distillers dried grains
with solubles; all of these are used as animal feeds. Many
Brazilian distilleries remove part of the yeast from the
fermentation as dry yeast for animal feed. By using a
process in which the yeast carbohydrate reserves (glyco-
gen and trehalose) undergo endogenous fermentation
the yeast-protein content, and thus the market value,
can be increased if the protein content is higher than
40% w/w.
Motor-grade fuel ethanol
Fuel ethanol is used in a variety of ways (Table 3).
Ethanol has a higher octane rating than petrol
enabling combustion engines to be run at a higher
compression ratio and thus giving a net performance
gain of approximately 15% w/w
, even though a litre
of pure ethanol contains only about two-thirds of the
calorific value of a litre of conventional gasoline. In the
past, direct government intervention in Brazil ensured
that hydrous ethanol was made available at all 26 000
gasoline stations and there are still over 3.5 million
vehicles that run on hydrous ethanol and consume two-
thirds of production (Table 1). Sales of ethanol-only
vehicles virtually ceased in the 1990s after confidence
in supply waned, but production is now resuming; at
the same time, the government has approved an increase
in ethanol content in gasoline to 26% and in diesel to 3%.
Today, the major use of ethanol is as an oxygenated
fuel additive that reduces emissions of carbon mon-
oxide, nitrogen oxides and hydrocarbons. In the USA,
there is considerable growth in E10 fuels (Table 3) but
these still only have approximately 12% of the market.
The European Union has recommended a 5% admix-
ture of ethanol in all gasoline for internal combustion
engines before 2010. As an additive, ethanol must com-
pete with methanol and two petroleum derivatives,
methyl and ethyl tertiary-butyl ether (MTBE and
ETBE, respectively). The latter is made from ethanol
and both are preferred by the petroleum industry,
although MTBE pollution is causing concern because
of its toxic and alleged carcinogenic properties
Biological aspects
A great deal of effort has gone into improving ethanol
yield during the past 25 years
, but there is still hope
for increasing throughput and reducing costs. During
the ethanolic fermentation of sugar-cane juice, up to
10% of the metabolized sugar can be diverted to form
glycerol and succinic acid, thus reducing the ethanol
yield (D. M. G. Alves, MSc thesis, University of So
Paulo, 1994). However, attempts to reduce succinic-
acid formation have resulted in reduced yeast competi-
tiveness in the industrial environment because of the
antibacterial action of this acid
. Increased glycerol
production is correlated with the acidic, ethanolic and
temperature-stressing conditions that are encountered
by the cells in the fermenter
. Because glycerol is
Table 2. Representative yields of products from
a maize wet-milling process
Product Per bushel
% w/w
Ethanol 9.84 litres 30.7
Crude maize oil 1.5 lb 2.7
Corn gluten feed (21% protein) 12 lb 21.4
Corn gluten meal (60% protein) 3 lb 5.4
Carbon dioxide 17 lb 30.4
A bushel is the commercial measure used for maize and is a
weight of 56 lb.
Table 3. Common ethanolic motor-fuel
Ethanol content
(% v/v)
Hydrous ethanol (lcool) (Brazil) 95.5
E85 (North America) 85
Gasoline (Brazil) 24
E10 (gasohol) (North America) 10
Oxygenated fuel (USA) 7.6
Reformulated gasoline (USA) 5.7
Biodiesel (Sweden) 15
Hydrous ethanol contains 4.5% (v/v) water. The other formu-
lations contain hydrocarbons and a trace of water.
TTEC Dec 99 24/10/99 9:51 am Page 483
synthesized specifically in response to osmotic stress, it
probably has an essential role in cell viability
. Using
very high sugar concentrations can reduce the fer-
mentation time but additional assimilable nitrogen is
often needed
and there can be rheological problems
with such a viscous fluid. Yeasts with improved proper-
ties such as ethanol and temperature tolerance have
been created in the laboratory and can be used in single-
batch cultures, but none have succeeded in the industrial
fermentation process with cell re-use.
It was only possible to demonstrate that the tradi-
tional inocula (bakers yeast and other selected strains)
were unable to compete with wild yeasts that contami-
nated the industrial process after strain identification
by karyotyping
. The probable reason for this is that
such strains have been selected to survive in laboratory
conditions rather than in the less-well-understood con-
ditions of commercial fermenters. Wild strains that
contaminated, dominated and persisted in the fermenter
and also had good fermentation abilities are now used
as inocula, dominating the fermentation in 21 out of 30
distilleries in Brazil (L. C. Basso, pers. commun.).
Environmental aspects
The main environmental advantages of ethanol over
gasoline are that it is a renewable and, in principle, a
fully sustainable resource that is less polluting than gaso-
line. The most important aspect is that the CO
released by ethanol combustion has been fixed recently
by growing plants and therefore this greenhouse gas
makes no net contribution to global warming, an
important fact for the USA, which is responsible for
23% of total global emissions. However, energy is still
required for all other aspects of the process, including
the production and distribution of fertilizers, planting
and cropping, transport of the ethanol, and manufac-
ture of distilleries; most of this energy comes from
fossil fuels. In Brazil in 1988, ethanol production rep-
resented 19% of the total liquid fuel that was consumed
(P. N. Nastari, pers. commun.). With bagasse used as
fuel, the energy output from ethanol exceeds energy
input by a factor of two
. With maize, the associated
production and distribution costs that require fossil fuels
give a negative energy balance
but this could still be
environmentally worthwhile if it replaces the use of
coal in power stations
Many of the early criticisms of fuel-ethanol produc-
tion concerned its low efficiency and level of pollution.
Total ethanol production in Brazil has increased at a rate
of approximately 4% per annum since 1990, thus pro-
viding 13% of total national energy requirements
(P. N. Nastari, pers. commun.). Concomitantly, costs
have reduced by 3% per annum since the introduction
of the ProAlcool programme, owing to a combination
of new sugar-cane varieties, improved cultivation tech-
niques and improvements in the industrial process
(extraction, fermentation and distillation). Similar gains
have been seen in the USA, largely as a result of imple-
menting newer technologies at more-modern fermen-
tation plants and because the cost of producing ethanol
has reduced by two-thirds over the past 15 years
Improvements in all aspects of the productivity of the
process have been successful and sustained (Table 4),
and new developments are continually being explored
(Table 5), although many remain to be proved in an
industrial setting.
Bagasse is used mainly for steam generation and sup-
plies the energy for crushing, heating, juice concen-
tration and ethanol distillation. There is still a 20% sur-
plus that is used in the cogeneration of electricity,
which can be sold to electricity companies during the
dry season in Brazil, when the electricity demand is
high and the hydroelectric supply is at its lowest. Par-
tially hydrolysed bagasse is also being used as an animal
feed and in the paper industry, and is even being
explored as a building material
. Sugar-cane and maize
stillage was originally disposed of in water courses or
in surface ponds and was a serious pollutant. Today,
however, almost all of it is returned to the land as fer-
tilizer or used in the controlled production of methane
(biogas). Traditionally, before harvesting sugar cane
manually, the leaves and tops are burned in the field.
However, mechanical harvesters can harvest the whole
plant so, if the tops are burned in the distilleries in
Table 4. Comparison of ethanol production from
sugar cane and maize
Production stage Sugar cane Maize
Yield of crop (tonnes hectare
) 80100 12
Yield of ethanol best 9000 4400
(litres hectare
Yield of ethanol average 7750 2750
(litres hectare
Average fermentation 92 89
efficiency (%)
Typical fermentation time (hours) 610 40
Based on a theoretical maximal production of 511 g ethanol (kg glucose)
Table 5. Recent developments in fuel-ethanol production
Area Development Refs
Fermentation Highly concentrated wort (very high gravity) 19
Continuous CO
stripping of ethanol 20
Continuous membrane-bioreactor separator to retain yeasts and remove ethanol 21
Fluidized beds 22
Selection of suitable yeast strains 12,13
Continuous fermentation unit 23
Immobilized yeast 24
Distillation Pressure-swing adsorption 25
Fuel ethanol Exploration of direct use in a fuel cell 26
New coproduct Corn-fibre oil and gum 27
TTEC Dec 99 24/10/99 9:51 am Page 484
high-efficiency boilers (using gasification
), there is an
increase of up to 40% in the total energy return.
One serious environmental problem has been the soil
degradation that occurs in all intensive agriculture: ero-
sion is 1030more rapid than for undisturbed soil
Today, however, low- or no-till practices are reducing
this problem. Most agricultural crops need a large input
of nitrogen to maximize production, which leads to sig-
nificant energy costs for the synthesis, transport and
application of fertilizer, as well as pollution because of
excess nitrogen run-off and nitrification of watercourses.
As nitrogen inputs become limiting, farmers who mon-
itor nitrogen content can reduce usage substantially.
Nitrogen-fixing endophytic bacteria (mainly Acetobacter
diazotrophicus, Azospirillum spp. and Herbaspirillum spp.)
have been discovered in both sugar cane and in maize
Their biological nitrogen-fixing role needs to be con-
firmed directly, but it has been suggested that up to 60%
of the plants nitrogen needs are supplied endogenously
when sugar-cane is grown in a low-nitrogen soil
In general, as a fuel, ethanol produces less CO, NO
and photochemical pollutants than gasoline. The amount
of gasoline pollutants is reduced both by replacing
gasoline with ethanol and by the use of ethanol as an
. Ethanol combustion does produce acetalde-
hyde and, because it has a low vapour pressure
ethanol releases volatile organic compounds (VOCs).
As a result of the USA Clean Air Act Amendments
(1990), oxygenated gasoline additives (MTBE, ETBE
or ethanol) were required to provide at least 2% by
weight of oxygen to reduce the ozone-forming poten-
tial of gasoline by reducing CO, NO
and VOCs. This
led to the Reformulated Federal Gasoline (RFG) Pro-
gram, which regulated the content and volatility of
gasoline during the summer months in specified areas,
and the Federal Oxygenated Fuels Program, which
required gasoline to contain 2.7% oxygen in other
designated areas of high risk of CO pollution in win-
ter. The RFG programme has been successful at reduc-
ing the ozone-forming potential of gasoline both
directly and indirectly, although probably by less than
Both maize and sugar cane are part of the global agri-
cultural industry and their prices vary depending on
supply and demand. The distillery-gate price of fuel
ethanol is higher than that of gasoline but it is harder
to estimate the full and true cost of both fuels. There
are certain principal components that need to be
known if a fair and accurate assessment is to be made
(Table 6). Many of these are indirect costs (externali-
ties) that have not until recently even been considered.
Usually, the retail price of ethanol is kept lower than
that of gasoline by highly visible tax advantages. The
USA has recently renewed (until 2007) a 5.4 cents per
gallon excise-tax exemption for E10 fuels (at the
pump), which corresponds to a 54 cents per gallon tax
credit for the producer or blender. However, the pump
price of gasoline also conceals huge, hidden indirect
costs. For example, the nominal world oil price is
approximately US$13 per barrel but, on the basis of the
existing data, one US Government agency estimates that
the indirect cost of gasoline (with respect to regulated
air pollutants alone) is US$45 per barrel, and the costs
of defending oil supplies in the Middle East could be
as high as US$9 per barrel (
Approximately 50% of oil used in the USA is
imported, thus accounting for 45% of the US trade
deficit, because domestic production of oil is decreasing
while the demand is increasing. Current US ethanol
production contributes US$1.5 billion per annum to
the trade balance and, up to 1997, Brazil had saved
US$35.6 billion in foreign exchange in reduced oil
imports. As Brazil is a net debtor to the financial market,
this saving has been worth another US$35 billion in
interest payments (P. N. Nastari, pers. commun.). For
both countries, an increase in domestic fuel production
could be very valuable.
Because approximately 50% of the revenue from
maize processing comes from the coproduct stream, it
is arguable whether coproducts have been developed as
solutions to a waste problem in ethanol manufacture or
whether ethanol production is a solution to an excess-
starch problem. What is clear is that coproducts have
not been utilized to their maximum potential; a switch
from animal feed to products for the food (including
nutraceuticals), cosmetic and pharmaceutical industries
could increase their value greatly (
Lignocellulose feedstock
With oil prices at an all-time low and political reluc-
tance to implement carbon taxes, fuel-ethanol produc-
tion will remain uncompetitive unless some other form
of cost reduction can be made. For instance, lignocel-
lulose has long been acknowledged to be an abundant
and potentially cheap feedstock for ethanol produc-
. It has been estimated that using agricultural,
forest and municipal wastes alone could replace the equiva-
lent of 40% of the US gasoline market, and growing
crops specifically for energy production could provide
the remainder
. Extensive research over the past 20 years
has been directed towards making an efficient conver-
sion of lignocellulose into sugars for fermentation
there are four major aspects that should be considered.
Table 6. Costs and benefits of fuel-ethanol production
Direct costs Direct benefits
Grain or sugar-cane feedstock Sale of ethanol
Electricity and water use Major national agricultural industry
Transport, processing, Cogeneration of electricity
fermentation and distillation Sale of coproducts
Distribution and supply Technological development
Tax subsidy or exemption
Indirect costs Indirect benefits
Land degradation Reduced emissions of green
Increased acetaldehyde pollution house gases
Foreign currency payments and Reduced pollution by CO and NO
interest Rural employment and social
Military protection of gasoline stability
supplies Renewable resource
Reduction in fossil-fuel use
Reduced dependency on
imported oil
TTEC Dec 99 24/10/99 9:51 am Page 485
Wood fibres consist of crystalline cellulose (~45% dry
weight) that is embedded tightly within hemicellulose
(~30% dry weight) and lignin (~25% dry weight). The
aim of pretreatment is to render the lignocellulose
structure suitable for subsequent enzymatic attack and
to solubilize some of the constituents, particularly
hemicellulose, which is usually broken down into its
constituent monomers. After the dry material is com-
minuted, several different treatments are possible,
including high-pressure hot water
, dilute acid
, cold
concentrated acid
, acid-catalysed steam hydrol-
, steam explosion
, ammonia fibre explosion
alkaline hydrolysis
and the use of nonionic surfactants
Each technique has its own associated problems, par-
ticularly in relation to the ability to recycle expensive
reagents and to avoid solubilization or the creation of
toxic compounds
A consortium of enzymes [endoglucanase, exoglu-
canase and -glucosidase (cellobiase), collectively known
as cellulase] are needed to break down cellulose into its
constituent glucose monomers
. Although they are
commercially available, usually derived from the fungus
Trichoderma reesei, their high cost is a major concern
Various approaches to reducing costs include improving
cellulase production by fungi
, recycling the cellu-
, binding the cellulase to inert supports
, cloning
the relevant genes into bacteria such as Escherichia coli
and the use of bacterial species such as Klebsiella oxytoca,
which naturally possess some of these genes
Pentose utilization and fermentation
Cellulose monomers are principally glucose and cel-
lobiose, whereas hemicellulose monomers are a mix-
ture of hexoses (principally glucose, with some mannose
and galactose) and substantial amounts of pentoses
(principally xylose, with some arabinose). The hexoses
can be fermented by S. cerevisiae but the pentoses can-
not. Three approaches have been attempted to enable
the xylose to be utilized: (1) to clone genes from pentose-
utilizing species into S. cerevisiae
; (2) to co-culture
two different strains of genetically modified Zymomonas
; and (3) to clone pentose-utilizing genes into
ethanol-resistant strains of E. coli
Simultaneous saccharification and fermentation
Because high concentrations of sugar monomers can
inhibit the enzymatic hydrolysis of cellulose, there are
advantages to continuously removing them from the
reaction vessel using SSF (Refs 61,62). One problem
has been that the optimal temperatures for saccharifi-
cation and fermentation are approximately 45C and
30C, respectively; however, this has been solved by
developing a non-isothermal system
Commercial plants
Although complex, in principle, all parts of the
problem have now largely been solved. However, the
greatest difficulty has been integrating these solutions
into an economic process. Several pilot plants have
existed for over 15 years
and the first commercial plant
is now being built. The feedstock will be sugar-cane
bagasse and rice hulls using a two-stage acid-hydrolysis
pretreatment, first to solubilize the hemicellulose and
then the cellulose. Following neutralization, the sugars
will be fermented by a hexose- and pentose-utilizing
E. coli strain (KO11)
. The solid-waste stream will be
burned in boilers, even though it will contain lignin,
which is a potentially valuable byproduct. The plan is
to produce 75 million (l ethanol) y
with a price simi-
lar to gasoline. When it comes on-line in the middle
of the year 2000, the plant will provide the first real
technical and economic test of such a production sys-
tem. If the technology is successful, sugar-cane produc-
ers in Brazil will be in a particularly favourable position,
as this will make virtually their entire sugar-cane crop
convertible into ethanol without significant additional
transport costs (most of the material is already being
brought to the distillery).
Future prospects
Justification for the fuel-ethanol industry has always
been contentious
but political priorities have deemed
it desirable. Initially, the aims were to become inde-
pendent of the petroleum market and to reduce the
costs of expensive oil imports. Now, the emphasis is on
reducing pollution and making a contribution towards
the Kyoto protocol by limiting global warming. Adding
ethanol as an oxygenate is successful, with E24 being
used in all Brazilian gasoline and the use of E10 and
RFG fuel increasing in the USA; however, the use of
pure ethanol is in decline
. Fuel-ethanol production
will only remain a significant industry and become a
potentially self-sustainable agriculture system for the
21st century if the utilization of lignocellulose becomes
a commercial reality.
We thank CAPES, FAPEMIG and the Leverhulme
Trust for financial support, PADCT/FINEP for grant
64.96.0118.00 and P. M. Nastari for advice on economic
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TIBTECH DECEMBER 1999 (VOL 17) 0167-7799/99/$ see front matter 1999 Elsevier Science Ltd. All rights reserved. PII: S0167-7799(98)01207-4 487
Large-scale applications of NAD(P)-dependent
oxidoreductases: recent developments
Werner Hummel
NAD(P)-dependent dehydrogenases are useful catalysts for the synthesis of chiral compounds. Many active and stable enzymes
are available that (with high enantioselectivity) reduce ketones or keto acids to chiral alcohols, hydroxy acids or amino acids.
For economic reasons, these reactions need coupling to the simultaneous regeneration of NAD(P)H. For preparative applications,
three components have to be combined: an appropriate enzyme, an efficient coenzyme-regenerating step and a suitable reaction-
engineering technique.
xidoreductases, which include the dehydrogen-
ases, are enzymes that catalyse redox reactions.
To do this, they require a coenzyme. Some of
these coenzymes, such as flavine adenine dinucleotide
and flavine adenine mononucleotide are bound to the
enzyme. By contrast, nicotinamide adenine dinucleotide
(NAD) and nicotinamide adenine dinucleotide phos-
phate (NADP) mostly exist as a soluble component of
the enzyme. A typical reaction scheme for the use of
dehydrogenases in the synthesis of chiral alcohols,
hydroxy acids or amino acids is shown in Fig. 1. Many
specific enzymes of this type are known to be available
W. Hummel ( is at the Institut fr
Enzymtechnologie, Heinrich-Heine-Universitt, Forschungszentrum
Jlich, PO Box 2050, D-52404 Jlich, Germany.
TTEC Dec 99 24/10/99 9:51 am Page 487