Vol.

74, 1968] 55

CONTINUOUS FERMENTATION OF BREWER'S WORT

By A. D. Portno
{Brewing Industry Research Foundation, Nutficld, Surrey)

Received 26th June, 1967

Results from a wide range of systems for the continuous fermentation of brewer's
wort demonstrate that two criteria determine the operational aspects of such
systems. These are:
1I) The degree of mixing, and
(2) The degree of restraint over the escape of yeast.
A reduction in the ability of yeast to ferment occurs when high concentrations of
yeast are maintained and is evident to a greater extent in homogeneous systems than
in those which maintain a fermentation gradient.
Results suggest that satisfactory systems must allow partial escape of yeast and
that a minimum degree of heterogeneity should be maintained. These requirements
necessitate a system which either maintains a concentration gradient in an elongated
vessel or comprises a series of homogeneous vessels with a feed-back of cells so that,
in each instance, yeast in the system is regularly exposed to a high concentration of
nutrients.

Introduction and Discussion system of continuous fermentation. It is

Continuous fermentation methods were given by the expression
classified by Herbert7 into closed systems
from which no cells are allowed to escape and
open systems when the concentration of cells -OHO 100

where C = the closure index expressed as %

in the outflowing liquid is equal to that in
the fermenting medium. Both these types
of system can be further classified into
homogeneous systems, the contents of which
are thoroughly mixed in a single vessel thus
ensuring that the composition of the culture
is spatially constant and heterogeneous sys~
terns, in which exists a fermentation gradient.
Such a gradient can be brought about in
either a tubular reactor such as a packed
tower or a series of stirred vessels, thus
ensuring spatial variation in the culture.
Heterogeneous-closed systems can be sub
divided into those embodying a single-phase
when cells and substrate move together
through a fermentation gradient, a condition
which in practice is almost unattainable, and
two-phase systems such as a packed tower, in
which one phase, ordinarily consisting of the
micro-organisms, moves more slowly through
the system allowing a difference in the
residence times of liquid and yeast.
In this communication the term closure-
index is used to denote the degree to which
yeast is prevented from escaping from a
y = the concentration of yeast in the
outflowing liquid
and X = the concentration of yeast within
the system
both expressed as g. per litre.
Homogeneous systems for the continuous
fermentation of brewer's wort which accumu
late large concentrations of yeast by oper
ating under conditions when the closure
index is high and therefore nearly all the
yeast is retained, are inherently unstable7:
they are therefore unable to maintain steady
internal conditions and thus give rise to an
inconsistent product even though in some
circumstances the degree of fluctuation may
be small. It was shown that this effect is
caused by an overall loss of fermentative
power which is reflected by a reduction in
the ability of the cells to reduce methylene
blue.19
Most continuous fermentation systems
which function under partially-closed con
ditions rely on the sedimentation of yeast
56 PORTNO: CONTINUOUS FERMENTATION
under gravity in order to build up high
concentrations.6'10-11 •28 Under such condi
tions the fermentative capacity of the yeast
decreases and this is reflected by an increase
in the specific gravity of the liquid within
the system; this may induce the yeast to
deflocculate and therefore lead to it being
washed out of the apparatus. Such an
effect has been observed both in these
laboratories and elsewhere.10'11
In order to allow such a sequence of events
to proceed beyond this point and thus to
facilitate examination of the process in
greater detail, a new system of continuous
fermentation was constructed and is illus
trated in Fig. 1. The apparatus employs a
Arphon
Syphoning Foaming
break hamber
.eve. ,—-&fi«L
1
Level
fluctuation
Wort
Yeast
circulation
Fig. 1.—Closed system of continuous fermentation
with filtration barrier to prevent escape of
yeast.
filtration barrier in order to prevent the
escape of yeast in the outflowing liquid and is
capable of operation under fully closed
conditions or, when yeast is deliberately
removed from the system, at any closure-
index which might be required.
The fermenting vessel consists of a porous-
walled tube constructed of fibre glass and
impregnated with resin. The wall of the
tube has an average pore size of 2 microns;
[J. Inst. Brew.
it has an internal diameter of f in. and is 35 in.
long. The vessel broadens at its upper end
to form a glass bulb which serves as a cham
ber within which foam is confined. Fer
menting liquid which contains yeast is
withdrawn from the lower end of the tube
and recirculated to the foaming chamber
thus ensuring complete mixing throughout
the system. The tubular portion of the
fermenting vessel is enclosed within a glass
tube into which wort is introduced.
Equilibration occurs across the filter-tube
as the liquid is induced to flow alongside it,
causing wort to enter the fermentation
circuit and the products of fermentation to
leave: the nitration properties of the tube
are such as to prevent completely the escape
of j'east cells. Trials showed that free
transfer of liquid across the filter-tube was
insufficient to prevent the deposition of a
layer of yeast on the internal surface and
therefore, in order to prevent this, an
automatic back-flushing system was attached
to the liquid outlet. This, by an intermittent
syphoning action, induced a reciprocating
transfer of liquid across the filter-tube and
thus ensured freedom from obstruction.
Fig. 2 shows the changes in specific gravity
and yeast viability, as estimated from
methylene-blue staining, which occurred
during a fermentation when a powdery yeast
(N.C.Y.C. No. 240) was used. During this
trial no yeast was allowed to escape and the
system was thus operating under conditions
when the closure index was 100%; it can be
seen that under these conditions the specific
gravity of the liquid in the system fluctuated
considerably during a distinct overall in
crease. These altered levels of specific
Time (days)
Fig. 2.—Changes in specific gravity
and the ability of yeast to reduce methylene
blue (O—O—O) which occurred during the
operation of a closed system of continuous
fermentation.
Vol. 74, 1968] PORTNO: CONTINUOUS FERMENTATION
gravity were reflected by changes in the
ability of the yeast to reduce methylene
blue.
Examination of the sugars present in the
medium during the course of the trial
showed that the fluctuations in specific
gravity were largely caused by altered levels
of maltose in the fermenting medium; this is
plainly a reflection of changes in the ability
of the yeast to remove it. Table I shows
TABLE I
Changes in Specific Gravity and Concentra
tions of Maltose which Occurred in a Closed
System of Continuous Fermentation
Concentration
Time Specific of maltose
Sample* (days) 1 gravity (g. per litre)
A 4 10063 10
B 0 10122 120
C 12 10070 20
D 18 10135 17-0
E 30 1-0176 21-0
Wort — 10400 430
* Letters A-E refer to points labelled in Fig. 2.
alterations in concentrations of maltose
which accompanied the changes in specific
gravity illustrated in Fig. 2. It thus
appears that the failure of homogeneous-
closed systems of fermentation to maintain
stability in the manner previously described
may, at least in part, be caused by a decline
in the ability of the yeast to remove and
ferment sugars; the consequent build-up in
their concentrations leads to deflocculation
and thus the loss of a major proportion of the
yeast. The results obtained from the filtra
tion system of fermentation and shown in
Fig. 2 suggest that the fermentative capacity
of the yeast initially recovers so as almost to
reach its former level, even if its escape from
the system is prevented. Further studies
were therefore designed to examine the
implications of these findings for the opera
tion and design of continuous fermentation
systems.
Early work on the continuous fermentation
of brewer's wort was concerned predomin
antly with open-homogeneous systems and
the results reported by most authors were
obtained during the maintenance of steady
conditions at specific gravities which repre
sent only partial attenuation.8'21 This may
be attributed to the fact that when such
57
systems are employed, steady-state condi
tions become increasingly difficult to achieve
as the attenuation limit of the wort is
approached.
Changes which occurred in the specific
gravity of the contents of an open-homo
geneous continuous fermentation system
which was operated for four weeks at an
efficiency of 0-8 and under constant condi
tions are shown in Fig. 3. It may be seen
that a rather regular pattern of changes
occurred similar to those which were observed
in the closed fermentor which was described
earlier, although in the present example the
magnitude of the fluctuations was reduced
and no overall decrease in the fermentative
capacity of the yeast was observed.
16
Time (days)
Fig. 3.—Measurements of specific gravity during the
operation of an open-homogeneous continuous
tormentor.
Consideration of these two sets of obser
vations with due regard to the features of the
systems from which they were derived
suggests the existence of two effects:—
(a) a rhythmic fluctuation in specific
gravity caused by changes in the
fermentative capacity of the yeast
and which appears to be primarily
related to the homogeneity of the
system, and
(b) an overall slow rise in specific gravity
apparently due to a gradual loss of
fermentative capacity as a function
of the closure index.
In a fully heterogeneous system of con
tinuous fermentation such as that described
in an earlier publication and in which the
essential features of a batch fermentation are
reproduced, fluctuations such as those ob
served in homogeneous systems do not
occur.19 However, the operation of hetero
geneous systems under excessive degrees of
closure induces a slow loss of fermentative
capacity (effect (b) above) and which, in
closed-homogeneous systems such as that
described above, may occur superimposed
 PORTNO: CONTINUOUS FERMENTATION [J. Inst. Brew.

upon the fluctuations in fermentative Wort Beer + Gas Sample removal

capacity (effect (a)). The former effect
therefore appears to be a function of the
closure index and the consequent inadequate
growth of yeast when this becomes excessive.
In a batch fermentation, or in a hetero
geneous continuous system, such as a tubular
reactor with piston-flow, yeast is introduced
into a medium which contains a high con
centration of nutrients. In contrast to this,
yeast in a homogeneous system is maintained
in a medium of the composition of the out
flowing liquid and which thus contains low
Still zone
concentrations of nutrients.
It has been shown that changes occur in
the ability of yeast moving through a con
centration gradient in both a tubular reactor
and a multi-vessel system to remove and
ferment glucose, maltose and maltotriose
and that the ability to ferment such sugars is
related to their concentrations in the sur
rounding medium.17-20 It thus appeared
likely that yeast cells which were maintained
within a homogeneous system of continuous
fermentation which contained very low Magnetic
concentrations of fermentable sugars might Stlrrer
exhibit a decreasing ability to absorb and Fig. 4.—Simple homogencous-partially closed con
ferment them. Such an effect would ob tinuous fermentation vessel.
viously result in a rise in the specific gravity
of the medium until the concentration of between 1-014 and 1-016. When the former
sugars was sufficient to induce the enzyme level had been reached, agitation of the
systems, whereupon active fermentation fermentor was interrupted and the yeast
would recommence. Repetition of this effect allowed to sediment. Adjustment of the
with time would give rise to a periodic rate at which wort was introduced then
instability such as that observed in homo resulted in a swift rise in the specific gravity
geneous systems. Conversely, the exposure of the contents of the system to 1-032. The
of yeast to high concentrations of sugars at introduction of wort was then interrupted
the commencement of each cycle in a hetero and stirring recommenced, the system being
geneous system would stimulate induction allowed to operate as a stirred-batch fermen
of the necessary enzyme systems and thus tation until the specific gravity had been
prevent the effect. It was with this possi reduced to 1-008; at this time continuous
bility in mind that further investigations operation was resumed and it was found that
were devised. a significant increase in the ability of the
Three trials were conducted in order to yeast to ferment had occurred as a result of
test the validity of the concept. In the first, its exposure to a high concentration of
a simple homogeneous-closed system of nutrients. This treatment arrested the ten
fermentation similar to that illustrated in dency for the specific gravity to rise and
Fig. 4 and which operated at a closure allowed continued production of fully attenu
index of between 94 and 98% was allowed to ated beer at the original efficiency for a
ferment wort of specific gravity 1-040 under further period. In a control experiment,
constant conditions and at 20° C. until the which had been operated alongside that
specific gravity was observed to rise. Pre already described, but which was allowed to
vious work had shown that, with the strain run without interruption, the specific gravity
of yeast involved (N.C.Y.C. No. 1026) continued to rise until the yeast deflocculated
deflocculation and thus wash-out would and wash-out occurred. The system then
occur when the specific gravity had risen to operated as an open continuous fermentor
Vol. 74, 1968] PORTNO: CONTINUOUS FERMENTATION
producing partially attenuated liquid of
specific gravity 1-035 from brewer's wort of
specific gravity 1-040.
In the second investigation, the same
simple continuous fermentors were employed
and were operated under identical conditions,
using the same wort and yeast as in the
former trial. The two fermentation systems
differed only in that an additional stirred
vessel was installed in series with one of
them, yeast from the main fermenting vessel
being recirculated to meet the wort which
entered through the small pre-treatment
vessel. This arrangement was intended to
ensure that yeast which was recirculated was
exposed to a high concentration of nutrients
in order to stimulate induction of the neces
sary enzyme systems. The specific gravity in
the main fermenting vessels was 1-011 and
that in the pre-treatment vessel 1-026. The
volumes of liquid in the vessels was 1200 ml.
in the main fermenting vessel and 600 ml. in
the pre-treatment vessel. Both fermentation
systems were operated at an efficiency of 3
which corresponds to a dilution rate of
0-125 h.-i
Fig. 5 shows the changes in specific gravity
which were observed when the systems were
operated for a period of 20 days. In the
case of the single-vessel fermentation system,
the specific gravity rose after 10 days
resulting in wash-out and the subsequent
functioning of the system as an open con
tinuous fermentor attenuating through 4
degrees only. A parallel rise in specific
gravity was observed in the two-vessel
system although its magnitude was con
siderably reduced and the fermentative
capacity of the yeast recovered before the
point was reached where deflocculation
would have occurred; this allowed recovery
and a consequent reduction in the specific
gravity to the original level. Plainly this
degree of heterogeneity is sufficient to
20
Time (days)
Fig. 6.—Changes in specific gravity in two systems of
continuous fermentation:—
(a) A single-vessel system (0—0—0) and
(6) A two-vessel system (■—■—■).
69
minimize the effect and therefore to prevent
wash-out, but is insufficient to suppress all
instability.
In the third trial the two fermentation
systems were again operated side by side;
on this occasion the pre-treatment vessel
was absent and the systems thus were
identical in all respects. The specific gravity
of the contents of both vessels rose to
between 1-014 and 1-017 and at this point
the inflow of wort was stopped. Into one
fermentor 100 ml. of a solution of maltose
was added; this contained 51-6 g. of maltose
and resulted in the total concentration of this
sugar in the vessel being elevated to that of
unfermented wort of specific gravity 1*040
(43 g. per litre). 100 ml. of sterile water was
introduced into the other fermentor as a
control.
Fig. 6 shows the results obtained when,
after allowing the fermentors to stand for
24 hr., the introduction of wort was recom
menced at the former rate. In the control,
to which water had been added, the specific
gravity continued to rise at an increasing
rate and ultimately induced deflocculation
of the yeast and wash-out. On the other
hand, exposure of the yeast to the high
concentration of maltose delayed the sub
sequent rise in specific gravity. However,
by reference to the first trial and to Fig. 5 it
is clear that maltose alone was less effective
than wort in so doing and therefore, although
concentrations of maltose which influence the
ability of yeast to remove and ferment it are
of obvious importance in this respect, other
i-ok
Time (days)
Fig. 6.—The effect on specific gravity of the addition
of maltose to a closed-homogeneous system as
the fermentative ability of the yeast com
menced to decline (O—O—Q) and results
obtained from a control to which water was
added (©—#—#).
60 PORTNO: CONTINUOUS FERMENTATION
sugars, and possibly additional materials of
the type which are required for the synthesis
of enzymes, are probably involved.
It is possible that, as a yeast cell ages, its
ability to bring about fennentation declines
and that, at a point dependent upon fermen
tation conditions and the individual strain
of yeast, its fermentative capacity becomes
seriously impaired. Clearly, therefore, it
seems desirable to adjust the conditions
under which a continuous fermentor operates
so that the time during which yeast cells
remain in the system is regulated. This
would enable the turnover to be maintained
at such a level that a large proportion of the
ageing cells would leave the system before
the time at which their fermentative
capacities commenced to decline.
Fig. 7 shows the rate of reduction in the
ability of cells to reduce methylene blue with
time in the case of two closed systems of
continuous fermentation, one of which was
homogeneous whilst the other was hetero
geneous. It can be seen by reference to the
Figure how the fluctuations which occur in a
homogeneous system confuse the secondary
effect which is predominantly due to an
excessive closure-index. Such fluctuations
do not occur in heterogeneous continuous
fermentors and although the mean slope of
the curve which depicts the reduction in
viability is the same in both types of system
this becomes confused in the case of homo
geneous fermentors by the fluctuations
which occur around the mean. In this
instance a mean reduction of 40% in the
ability of yeast cells to reduce methylene
blue occurred over a period of 40 days and
at closure indices of 94% and 100%.
20 30 40 SO
Time (days)
Fig. 7.—Alterations in the ability of cells to reduce
methyleno blue which occurred in
(a) a closed-homogeneous continuous fer
mentor (O—D—D)
and (6) a closed-heterogeneous continuous fer
mentor (O—O—O).
[J. Inst. Brew.
The growth of yeast in continuous fer
mentation systems is frequently considered
as specific growth rate (p). This expression
is a measure of the rate at which new cells
are produced and is related to the generation
or doubling time.
However, the specific growth rate is
essentially a measure of the rate at which
new cells are produced and, as such, was
devised in connection with studies on the
continuous culture of micro-organisms when
the main objective was to produce such
organisms at or near maximum rates.
In the continuous fermentation of wort the
end product is the fermented liquor and
growth of yeast is clearly a secondary con
sideration. In these circumstances such
growth is required for two purposes:—
(a) to produce new cells at a rate at
least equal to the rate at which
(i) cells escape from the fennentation
system,
(ii) old cells lose the ability to
ferment; and
(b) to regulate the flavour of the finished
beer which, in several respects, has
been shown to be influenced by the
amount of yeast growth which
occurs.18'18
In order that a continuous fermentation
system should function under steady con
ditions, it is plainly essential to maintain a
constant concentration of active yeast
within it and the population level which is
necessary will depend on both the efficiency
at which the system is required to operate
and the fermentative ability of the par
ticular yeast involved.
Fig. 8 shows the concentrations of yeast
required to ferment wort from a specific
gravity of 1-040 to 1-010 over a range of
efficiencies from 1 to 13 and is based on the
results reported by several authors8'8'9'8'10-13
all of whom claimed that the continuous
fennentation systems from which the results
were obtained operated under steady condi
tions. Clearly, if the fermentative capacity
of the yeast is maintained at a constant level
these proportions should stand in linear
relationship. However, in the presence of
very high concentrations of yeast, the volume
occupied by the yeast cells in the fermentation
vessels becomes a significant proportion of
the total capacity and extrapolation of the
graph is thus only possible if compensation
Vol. 74, 1968] PORTNO: CONTINUOUS FERMENTATION
ine corrected
Tor volume of /east
10
I 7
tfc
HI
0 20 « 60 80
Concentration of yeast (g. per litre, dry weight)
Fig. 8.—Concentrations of yeast required to ferment
wort from a specific gravity of 1-040 to 1-010
at a range of efficiencies.
is made for this effect. The deviation of the
curve from the corrected line is clearly a
function of decreased fermentative capacity
in systems which functioned at high effi
ciencies and may have been largely caused
by inadequate levels of growth in the presence
of high concentrations of yeast and by low
levels of enzymic activity in cells which
fermented in homogeneous systems.
Adjustment of the amount of yeast growth
which occurs within a continuous system
can be achieved by regulation of the supply
of oxygen to the culture, accompanied by
control of the rate at which yeast is removed
or allowed to escape. Such control can be
used to regulate the residence time of yeast
cells in a suitable continuous system at such
a figure that the viability and fermentative
capacity of the yeast is maintained at a
consistently high level.
A further aspect of the growth of yeast in
continuous fermentation systems stems from
its influence upon the production of com
pounds which are of importance in connec
tion with flavour and aroma, and it has been
suggested that a level of yeast growth equal
to that which occurs in a batch fermentation
is necessary in order to produce beer of
comparable quality from a conventional
brewety wort.1
61
The results in Table II partly confirm this
suggestion and show analyses obtained,
using beers produced at three levels of yeast
growth in a homogeneous pilot scale con
tinuous fermentation system. The inter
mediate level of growth, 3-0 g. per litre (dry
weight), is comparable to that generally
achieved in a batch fermentation using the
same wort and yeast and it can be seen that
the beer produced under these conditions
was very similar to that produced in the
control batch fermentation. However, the
results show that, whilst levels of growth in
excess of that which occurs during a batch
fermentation may result in the production of
undesirable flavour and aroma character
istics, cell yields below such a level appear to
have no such detrimental influence. Fur
thermore, previous studies, when the in
fluence of different levels of wort aeration on
the course of fermentation in a heterogeneous
system was investigated, show that little
difference could be detected in the quality
of the beer produced at levels of growth
varying from 2-36 to 2-98 g. per litre (dry
weight) although the viability and fer
mentative capacity of yeast cells in the
system remained greater at the higher
levels of growth.
It is concluded that decreasing fermenta
tive capacities due to ageing populations of
cells are not likely to present a serious
problem provided that:—
(a) a level of growth comparable to that
achieved in a batch fermentation is
sustained, and
(b) a concentration of yeast is maintained
which is related to the throughput
and the growth of yeast is not allowed
to result in the accumulation of
excessive concentrations; this results
in prolonged residence times of yeast
cells and thus a population the average
age of which may become excessive.
Fig. 9 illustrates the changes in specific
gravity with time which typify several types
of system for the continuous fermentation of
brewer's wort. Obviously, if a high efficiency
is required and it therefore becomes necessary
to employ a system which functions at a
high closure index, a significant degree of
heterogeneity will be required in order to
ensure adequate stability.
It has been suggested, as a result
of theoretical studies, that a two-stage
62 PORTNO: CONTINUOUS FERMENTATION [J. Inst. Brew.
TABLE II
Analysis of Beers Produced at Different Levels of Yeast Growth in a Homogeneous System
Compared with that from a Batch Fermentation

Homogeneous continuous system Batch

Yeast growth (y) g. per litre 2-36 3-04 3-80 3-10

Cone, of yeast (x) g. per litre . 36 42 41 —

Percentage non-staining cells . 62 87 97 84

pH 4-09 4-03 3-02 4-00
Specific gravity 10113 1-0102 1-0105 1-0100
Total nitrogen (mg./l.) ■ ■ 503 452 393 380
a-Arnino nitrogen (mg./l.) 481 310 29-0 24-0
Bitterness (mg./l.) 26-6 22-7 24-4 230
Diketones (mg./l.) 0-4 0-8 21 0-7
Ethyl acetate (mg./l.) .. 86 7-7 12-4 8-9
n-Propanol (mg./l.) 30-4 20-1 310 19-4
Isobutanol (mg./l.) 32-5 36-1 41-3 29-4
Iso-amyl acetate (mg./l.) 014 0-19 0-23 0-20
Iso-amyl alcohol (mg./l.) 53-2 38-8 59-2 42-4
Ethyl caproate (mg./l.) .. 008 0-02 0-01 0-02
Acetaldehyde (mg./l.) • ■ 7-4 12-0 370 8-6
Acetone (mg./l.) 053 4-0 3-0

HOMOGENOUS Present results agree well with this suggestion

1-040
and preliminary trials, when such a system
Partially-closed was operated on a pilot scale, showed that it
1-030
was capable of maintaining very stable
conditions for prolonged periods; during this
trial the viability of the yeast was main
tained at a high level. It appears that
further studies on this type of system might
Open result in the development of a satisfactory
Time
stable system for the continuous production
of beer.
Experimental
HETEROGENEOUS All wort used in these trials was of 1040
[■(MO
gravity, was derived entirely from pale ale
Partially- malt and, in the case of the laboratory-scale
5" 1-030 Wash-out due/ closed
trials, was brewed in a conventional pilot-
to deflocculation
scale plant as described by Hudson &
u 1020 Fully-closed
S Deflocculation Birtwistle,12 and was collected whilst boiling
into 20-litre reservoirs which had previously
1-010
Open
been sterilized in an autoclave. Wort used
in pilot-scale trials was of similar quality and
Time
was brewed continuously using the equip
ment described by Hudson & Button14; it
Fig. 9.—The changes in specific gravity with time
was subsequently stored at 0° C. and under
which characterize different types of system
for the continuous fermentation of brewer's sterile conditions. Typical analyses of such
wort. The figures are based on changes which worts are shown in Table III. Concentra
occur when brewer's wort of specific gravity tions of yeast were determined on the basis
1-040 is fermented to a level approaching the
of dry weight; other analyses were performed
attenuation limit.
as described by Maule et a/.16
fermentation system consisting of a homo The closed continuous fermentor which
geneous vessel in series with a heterogeneous retained yeast by a system of nitration was
tubular reactor might be the most suitable constructed primarily from standard sections
arrangement for the continuous fermentation of industrial glass tubing (Q.V.F. Co.).
of complex media such as brewer's wort.4 The nitration element was assembled from
Vol. 74, 1968] PORTNO: CONTINUOUS FERMENTATION

TABLE III 2. Cowland. T. W.. unpublished.

Analysis op Worts Produced from Kilned 3. Cowland, T. W., and Maule, D. R., this
Malt in a Pilot-scale Conventional Brewery, Journal, 1966, 480.
and in a Pilot-scale Continuous Brewbry 4. Fcncl, Z.. in A Theoretical and Methodological
Basis of the Continuous Culture of Micro
organisms, by Molek, I., and Fond, Z.,
Conventional Continuous Academic Press, 1066.
browery brewery 5. Hall. R, D., & Fricker, R.. Proc. Conv. Aust.
Sect. Inst. Brew., 1966, 45.
pH 6-2 6-OS 6. Harris, G., & Merritt, N. R., this Journal,
Specific gravity .. 1040 1040 1962, 33.
Total nitrogen (ing. 7. Herbert. D., S.C.I. Monographs, (12), 21.
per litre) 713 662 8. Hough. J. S., S.C.I. Monographs, (12), 219.
a-Amino nitrogen 9. Hough, J. S., Proc. Eur. Brew. Conv., Vienna,
(mg. per litre) .. 206 183-5 1961, 160.
Fermentable sugars 78% 76% 10. Hough, J. S., &. Ricketts, R. W., this Journal,
Bitterness (mg. per 1060. 301.
litre) 36-0 350 11. Hough, J. S., & Watson. T., this Journal.
1966, 647.
12. Hudson, J. R., & Birtwistle, S. E., this
Journal. 1966, 46.
fibre-glass filter-tubes (Whatman Co. Ltd.). 13. Hudson. J. R., & Button, A. H., Proc. Eur.
In all trials, samples were removed into a Brew. Com. Stockholm, 1986, 121.
14. Hudson, J. R., & Button, A. H., this Journal,
syringe through special sample points which in the press.
incorporated self-sealing rubber septa; these 16. Lawrence, W. C, Wallerstein Lab. Commun.,
have been described elsewhere.18 1964, 123.
16. Maule, D. R.. Pinnegar, M. A., Portno, A. D.,
Acknowledgement.—The author thanks Dr. & Whitear, A. L., this Journal, 1966, 488.
A. H. Cook, F.R.S., for his continued interest 17. Millin, D. J.. this Journal, 1966, 394.
and encouragement. 18. Portno, A. D., this Journal, 1966, 458.
19. Portno, A. D., this Journal, 1967, 43.
20. Portno, A. D.. this Journal, 1867, 478.
References
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