J. Inst. Brew.. March-April. 1996, Vol. 102, pp.

97-102

MALTING AND BREWING SCIENCE: CHALLENGES AND OPPORTUNITIES'*

By A. W. MacGregor

(Grain Research Laboratory, Canadian Grain Commission, 1404-303 Main Street, Winnipeg, Manitoba R3C3G8, Canada)

Received 15 September 1995

Molecular technologies have been developed for the transformation of barley. These technologies
complement current methods of barley breeding. In addition, they offer the potential of altering
specific components in barley that affect malting quality and of introducing foreign genes, control
ling desirable traits, into barley. Application of genetic engineering to improving malt quality factors
such as cell wall degradation, protein modification, starch hydrolysis and flavour stability, is
discussed. Limitations to the use of this technology for improving malt-related functional properties
of barley components such as cell walls and starch granules are also evaluated. Some possible
constraints to the utilization of genetic engineering for malt quality improvements are identified.

Key Words: Barley, genetic engineering, malt quality, cell walls,
proteins, starch, hydrolytic enzymes.
Introduction
Horace Brown was a scientist with very wide interests who
made significant contributions to a range of scientific disci
plines31. These included studies on the effects of rock form
ations on brewing water quality, the microbiology of spoilage
organisms in beer, the use of hops in brewing, the role of
nitrogen in malting and brewing and the physiology of barley
germination. It is this latter work, some of which is described
in a classic publication10 of 1890, that is associated most
closely with Brown and that is still referred to today. The
publication contains a detailed description of the structure of
the barley kernel based on painstaking observations using
microscopy, carefully developed arguments to support the con
clusions drawn and thoughtful and insightful speculation.
The intent of this aspect of Brown's research was to build a
sound knowledge-base of the germination process for the
malting and brewing industries. This would not only provide
answers to problems encountered during processing of barley
and malt but would provide a spring board to improve
the efficiency of malting and brewing technologies, provide
impetus to the development of new technologies and improve
the quality of the final product. In other words, Brown was
intent on building a strong scientific foundation for the techno
logies (malting and brewing) with which he was working so
that advantage could be taken of advances in relevant scientific
knowledge as these become available. This is an approach that
is as relevant today as it was in Brown's time.
What are some of the challenges and opportunities facing
the malting and brewing industries today? Stable, if not
declining, beer and malt markets in many countries have led to
increased competition for market share and the development
of "new" beer types such as dry and ice beer with associated
technological problems. As governments and industries curtail
spending there is less funding for research and there has been a
repositioning in research emphasis leading to a decrease in
longer term fundamental research and more emphasis on
shorter term "problem solving" research. An exploration of
the long term negative impact of this de-emphasis of funda
mental research is beyond the scope of this discussion. Along
with these changes are the increasing demands of consumers
for purity in food systems., i.e., a demand for "natural" foods
or foods made with minimum additives. This applies also to
the preparation of malt and beer and so extra demands will be
made on the quality of raw materials such as barley and malt.
Advantage must be taken, therefore, of new technologies that
have potential for improving the malting quality of barley.
Such a technology is plant genetic engineering3456-70 or the
manipulation, addition or deletion of quality-related genes in
barley. This offers the possibility of manipulating individual
components within the grain without changing other com
ponents. Equally important, however, is the potential for trans
ferring desirable genes from any organism into barley. This
opens up exciting possibilities for quality improvement of
malting barley.
Prerequisites for Genetic Manipulation
Several technological problems had to be overcome before
the new technique of genetic manipulation could be applied
successfully to cereal grains, such as barley35. For example,
efficient methods had to be developed for the delivery of DNA
into single cells37, selection of the transformed cells, re
generation of fertile plants from the transformed cells, and
expression of the new or modified DNA in appropriate tissues
of the grain96. Most of these problems have been overcome so
that all the common cereals have been transformed with at
least some degree of success (Table I). Several groups have
reported the successful transformation of barley so the techno
logy is now available43"521771'00
From past experience with transformation of other crops it
is likely for economic reasons that most of the immediate
research effort on barley will concentrate on developing
cultivars with herbicide tolerance and increased resistance to
diseases22-69. Use of the technology to improve malting quality
should be driven by the malting and brewing industries. They
have the monetary and personnel resources to control and
guide this approach to barley improvement.
Several different strategics arc available for manipulating
genes in cereals96. Insertion of desirable genes from other
TABLE I Transformation of Cereals
Cereal Year Transformed

•Based on Horace Brown Lecture presented at the Royal Society, Rice 1988 "■"■"
,990U.23J4
London, October 1993. Maize
fPaper No. M201 of the Grain Research Laboratory, Canadian Grain Wheat 199297102
Commission. 1404—303 Main Street, Winnipeg, Manitoba R3C 3G8, Oats I99290
Canada Bariey I99443-52-76-'00

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98 OPPORTUNITIES IN MALTING AND BREWING SCIENCE
organisms into barley has the potential to exert the greatest
change in quality parameters. It is possible also to alter exist
ing genes to produce subtle but specific changes in a gene
product but little progress has been made to date, with this
approach. Yet another approach would be to alter the pro
moter part of genes that control hydrolytic enzymes21 so that
synthesis of the enzymes would be switched on or activated
more rapidly during germination. This should have obvious
advantages in decreasing malting times. The use of antisense
technology to block the action of existing genes is a potentially
powerful aspect of genetic manipulation103. This technique has
been used with considerable success in a number of plants and
is being used currently to change the starch characteristics in
wheat".
Potential for Improving Malt Quality through
Genetic Engineering
Desired improvements in malt quality through changes in
barley characteristics have been discussed in several published
reports25-45-55. Many of these improvements could be achieved
through conventional plant breeding but the specificity of
genetic manipulation and the access it offers to a range of gene
pools make it a powerful complement to breeding programs.
Some possible applications of the technology will now be
discussed.
Endosperm Cell Walls
These form a barrier to the movement of enzymes in the
endosperm during malting and cause severe technological
problems during brewing if not degraded adequately during
malting3. The major component of the walls is a mixed-linkage
P-glucan that has been well characterized107. Despite that, little
is known about the biosynthesis of the polysaccharide in
barley. It is desirable to lower the levels of barley P-glucan but
the most effective way of accomplishing this at the present
time is through traditional breeding methods. The growing
environment also affects P-glucan levels in barley but this is
difficult to counteract6. Arabinoxylan, the minor component
of endosperm cell walls in barley, may also cause technological
problems because this polysaccharide, also, has the potential to
form viscous solutions58. The composition, structure, proper
ties and biosynthesis of arabinoxylans are not well understood.
Therefore, genetic manipulation of this minor but techno
logically important barley component is unlikely in the near
future.
Cell Wall Degrading Enzymes
Barley contains low levels of P-glucanase but a high pro
portion of the enzyme found in malt is synthesized in scutellar
and aleurone cells during germination1. It is important that
P-glucanase diffuses through the endosperm during malting
and degrades p-glucan throughout the endosperm3. The
enzyme has poor heat stability and may show limited activity
during mashing42. Therefore, to ensure minimal processing
problems, P-glucan degradation should be completed by the
end of malting. There are two major p-glucanases in malt and
they have been well characterized20-28. Their functional proper
ties, including their detailed action on P-glucan, have been
documented, their sites of synthesis during barley germination
have been located and the genes controlling synthesis of the
enzymes have been identified.
It is not surprising that the p-glucanase system in malt has
been targeted for genetic manipulation21. One approach has
been to insert genes for a heat stable p-glucanase from Tricho-
derma reesei into barley52 but the effectiveness of manipulating
barley in this way has yet to be evaluated. Parameters that
confer heat stability on proteins are not well understood60 but
when these have been identified, it may be possible to re-design
the appropriate barley gene to produce a more heat stable
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[J. Inst. Brew.
P-glucanase. Research is underway to produce hybrid P-
glucanases that would combine desired heat stability with
specified hydrolytic activity8. Results of such research may
find application in fermentation industries. P-Glucanases have
limited time to complete P-glucan hydrolysis during malting
and so their synthesis must be initiated rapidly during germin
ation. Research aimed at increasing the effectiveness of the
promoter region of the P-glucanase gene in barley would be
a worthwhile approach to increasing the effectiveness of P-
glucanase during malting21.
Even low molecular weight products from P-glucan hydro
lysis can cause technological problems108. Complete hydrolysis
of these products to glucose would prevent such problems and
also provide extra fermentable extract to the brewer. Barley
does contain a P-glucosidase83 that hydrolyzes the tri- and
tetrasaccharides produced by p-glucanase*0 but the effective
ness of the enzyme during malting and mashing has not yet
been determined. The enzyme is encoded by a single gene and
the probable sequence of the gene has been reported. This is
another barley gene that could be manipulated to increase the
usefulness of an enzyme for malting and mashing purposes
and so lead to an improvement in the malting quality of the
barley.
Extensive hydrolysis of arabinoxylans is also desirable
during malting. Limited information is available on the malt
enzymes required for such hydrolysis7-86 and so it is premature
at this time to envisage using genetic engineering technology to
increase the levels of these enzymes in malt although this
would be beneficial for malt quality.
Non-Enzymic Barley Proteins
The influence of barley proteins on malt and beer quality is
complex because of the large number of proteins present
in barley, the diversity of their functional properties and
the changes that they undergo during malting53-71-88-104. Some
effects of these interactions are listed in Table II.
Hordeins are a complex group of proteins80 that must be
modified extensively during malting or they will cause numer
ous processing problems during brewing. They are composed
of three main groups (B, C, D), each of which is heterogenous
but group D contains only one major component. Attempts to
evaluate the effects of each of these groups on malting quality
have had limited success although there is some indication that
a low D- to B-hordein ratio would be desirable71. Progress has
been made on identifying the hordcin genes98 but practical
utilization of this knowledge must wait for more detailed
information on the functionality of individual hordein pro
teins. It would then be feasible to alter proportions of different
proteins present in barley81 without increasing the total protein
content, which normally would not be desirable. The hordein
proteins also form a matrix in the barley endosperm that
embeds the starch granules. This matrix must be extensively
hydrolyzed during malting to release the starch granules so
that they can be solubilized effectively during mashing.
Gel-forming proteins have been identified in barley63-85-89.
These proteins reduce malt extract and cause filtration prob
lems and so have been the subject of several studies. The gels
consist of mixtures of high and low molecular weight proteins,
including B- and D-hordeins, that are joined together through
intra- and inter-molecular disulphide bonds63. Some com
ponents of the gel that is formed during mashing appear to be
TABLE II Protein Interactions During
Malting and Brewing
Viscosity-associated proteins
Head retention
Beer stability
Wort fermcntability
Protein matrix and starch granules
Protein type/quantity
Vol. 102, 1996] OPPORTUNITIES IN MALTING AND BREWING SCIENCE
derived from barley proteins that have been partially hydro-
lyzed by proteolysis during malting85. Again, insufficient in
formation is available on proteins that are essential for gel
formation and so it is premature to develop a strategy for
limiting their levels in barley.
Proteins are also actively involved in the formation and
retention of beer foam110 along with other components derived
from malt and hops. Recent results suggest that hydrophobic
proteins derived from hordeins are major components of beer
foam2-36. Other proteins such as protein Z44 and a trypsin/
a-amylase inhibitor36 have also been identified in beer foam. It
is the hydrophobic nature of a range of polypeptides rather
than a few specific proteins that appear to be important
constituents of beer foam. It is difficult with current know
ledge to increase specifically the head retention potential of
barley and malt because the property does not reside in
specific proteins.
Proteins and polyphenols have been implicated in the forma
tion of hazes in beer during storage61. This problem has been
controlled by judicious use of proteases to degrade the pro
teins and selective absorbents to remove proanthocyanidins,
the active polyphenol component of haze16-51. Identification of
the proanthocyanidins99 in barley that are responsible, along
with proteins and other beer components, for haze formation
has led to the development of proanthocyanidin-free barley
that does reduce the haze-forming potential of the resulting
malt". Little information is available on the barley proteins,
either intact or modified by proteolysis during malting, that
participate in haze formation. It is possible that some protein
cross-linking is involved in haze formation and so some of the
same proteins may be responsible for foam stability and haze
formation. Until more information is available on the identity
of haze-related proteins, little can be done to lower their levels
in barley either through genetic manipulation or through more
traditional breeding methods.
Proteolytic Enzymes
During germination barley proteins, mainly insoluble, stor
age proteins in the endosperm, must be converted into soluble
proteins, peptides and amino acids to supply nutrients to the
developing embryo. From a technological standpoint, the same
changes are important during malting because they lead to
destruction of the protein matrix, release of starch granules,
and to formation of amino acids in the malt that are required
for yeast nutrition during brewing. These changes are brought
about by a complex array of proteolytic enzymes15-54'74'"3.
Some of these are present in barley, others are synthesized
during malting and are active in the endosperm, while some
may be active only in the aleurone29. There is some controversy
about the activity of these enzymes during mashing41 because
they arc heat labile and so may be inactivated rapidly. There
fore, it is important that effective protein degradation is com
pleted by the end of malting. Over 40 different proteolytic
activities have been identified in malt"3 but only a few pro
teases have been characterized in detail. Proteases, like other
enzymes, have bond specificities and some of these have been
determined through elegant and very necessary studies33 but
much remains to be done in this area. With current knowledge
it is not possible to contemplate changing the protease com
plement of malt in the near future. An additional complication
is the presence of endogenous protease inhibitors in barley32.
The extent to which they may modulate protein modification
during malting has yet to be determined.
Barley Starch
Rapid degradation of starch to fermentable carbohydrates
during mashing depends on the starch being fully gelatinized18
before any of the starch-degrading enzymes are inactivated.
Starch granules in barley, therefore, should have the lowest
gelatinization temperatures possible. Factors affecting the gela-
tinization temperature of barley starch granules include
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99
growing environment95, granule size48 and, possibly, starch
structure"2. Little can be done to offset the environmental
effect. Barley cultivars containing a high proportion of large
granules would be preferred for malting because small granules
have higher gelatinization temperatures48 and do have the
potential to cause problems during brewing. Although signi
ficant progress has been made in identifying the enzymes
responsible for starch synthesis in cereals5172-73'87, controlling
and changing the proportion of large granules does not yet
appear to be feasible. There is some evidence that reducing the
length of the outer chains in amylopectin may reduce gelatiniz
ation temperatures of the starch"2, but, again, the biochemical
pathways controlling this parameter are not yet understood.
High amylose and waxy barley starches tend to have higher
gelatinization temperatures than do normal, large granules46
and so they do not offer any advantage to the brewer. The
starch content of potatoes has been increased by the insertion
of a starch synthesis gene92 and a similar effect could be
achieved in barley.
Starch-degrading Enzymes
It is unlikely that a-amylase levels are quality limiting in
barley malts66. Dextrin profiles in wort and beer4' suggest that
a-amylolysis of starch is usually complete by the end of mash
ing. The two major a-amylase components in malt have similar
activities and action patterns on solubilized starch51 so there is
no incentive to change the proportion of these components in
malt. ct-Amylase gene families in malt have been identified and
characterized65 and so could be manipulated if required. There
is a potential inhibitor of malt a-amylase 2 in barley64105106.
Since the inhibitor does not appear to play a role in malting or
mashing there is no obvious advantage to lowering its level in
barley.
P-Amylase plays a crucial role during mashing because it is
responsible for the degradation of starch and products of
a-amylase hydrolysis of starch to maltose, the most abundant
fermentable carbohydrate in wort94. The two amylases working
together are more effective in degrading starch than are the
two enzymes acting independently, p-amylase is synthesized
during barley development but is rendered fully active during
malting26-91. The enzyme is heat labile and a significant pro
portion of the activity is lost during kilning4-9. Enzyme activity
is also lost rapidly during mashing84 when the temperature
approaches 65°C. It is important, therefore, that sufficiently
high levels of p-amylase are present during mashing to hydro-
lyze the starch completely in the temperature range 58-60°C
(starch gelatinization temperature) to 65°C (temperature of
rapid inactivation of P-amylase). The presence of linear malto-
dextrins such as maltotetraose or maltohexaose in wort or
beer indicates incomplete hydrolysis of starch dextrins by (3-
amylase47. Obviously, it would be beneficial to increase levels
of p-amylase in barley and/or increase the temperature stabi
lity of the enzyme. Genes coding for P-amylase have been
detected391" so it would be possible to increase barley P-
amylase levels. The parameters that control the heat stability of
enzymes are poorly understood but some success has been
achieved in improving the heat stability of barley p-amylase68.
This opens up the possibility of improving the thermal stabi
lities of other barley and malt enzymes.
The technological significance of a-glucosidase is not clear.
It is important during malting for the conversion of maltose to
glucose93, a sugar that can be assimilated and metabolized by
the active embryo. However, brewing yeasts rapidly metabolize
maltose and so do not require the prior hydrolysis of maltose
to glucose. a-Glucosidase may increase the effectiveness of
P-amylase during mashing by removing maltose, a possible
competitive inhibitor of P-amylase75. There is no obvious
reason at the present time to alter a-glucosidase levels in malt.
Beer contains significant levels of branched starch dextrins"
indicating that the starch debranching enzyme, limit dextrinase,
is largely ineffective during mashing. This is due, in large
100 OPPORTUNITIES IN MALTING AND BREWING SCIENCE [J. Inst. Brew.

measure, to the presence of limit dextrinase inhibitors in the TABLE m Constraining Factors on
malt49. These low molecular weight proteins are present in Genetic Manipulation
barley30 and, although their levels are reduced during malting,
sufficient inhibitor remains in the malt to inhibit a high pro Effect on plant/grain
Specificity of desired change
portion of the malt limit dextrinase. The heat stability of limit
Environment
dextrinase is similar to that of (J-amylase84 and so the enzyme
Multiple genes/quality factor
should be reasonably effective during mashing in the absence Consumer resistance
of the inhibitors. It would be beneficial to develop barleys with
low levels of inhibitor, using antisense technology, and in
creased potential to produce limit dextrinase during malting by
increasing the gene dosage for the enzyme. Neither of these
approaches is feasible at the moment because the genes coding
this area will be made now that genetically engineered food
for limit dextrinase and the inhibitors have not been identified
stuffs are available30.
and characterized. Increased limit dextrinase activity during
Changes to plants through genetic engineering are now a
mashing would have to be controlled carefully so as not to
reality. The malting and brewing industries must take advan
remove all branched dextrins because they contribute to mouth
tage of this powerful, new technology. Brewers should decide
feel and body in the final beer79. Genetic manipulation of the
what types of malt they will require in the next 5-10 years,
limit dextrinase system in malt does have the potential to
keeping in mind that public pressure will reduce the use of
increase markedly the fertnentability of worts.
additives in food production so malt quality will become even
more important. Maltsters, in turn, must take the initiative in
identifying more clearly desired quality parameters in barley
Flavour Stability
and malt so that appropriate genes can be identified, located,
Numerous components contribute to the flavour of beer and
characterized and altered appropriately. A collaborative effort
several that produce undesirable off-flavours have been identi
involving many disciplines will be required if the full potential
fied59'67. Only one example will be discussed here to illustrate
of genetic engineering is to be exploited for the improvement
how genetic manipulation could be utilized to improve flavour
of barley malting quality.
stability in beer. The undesirable "cardboard" flavour some
times found in beer has been traced to trans-2-nonenal18. This
aldehyde originates, via a series of reactions, from the un-
saturated fatty acids linoleic and linolenic that are produced
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