Appl Microbiol Biotechnol (1986) 23:228--233
Microbiology
The fermentation of hexose and pentose sugars
by Candida shehatae and Pichia stipitis
J. C. du Preez, M. Bosch, and B. A. Prior
Department of Microbiology, University of the Orange Free State,
P.O. Box 339, 9300 Bloemfontein, Republic of South Africa
Summary. The fermentation by Candida shehatae
and Pichia stipitis of xylitol and the various sugars
which are liberated upon hydrolysis of lignocellu-
losic biomass was investigated. Both yeasts pro-
duced ethanol from D-glucose, D-mannose, D-gal-
actose and D-xylose. Only P. stipitis fermented D-
cellobiose, producing 6.5 g-1-1 ethanol from 20
g. 1-1 cellobiose within 48 h. No ethanol was pro-
duced from L-arabinose, L-rhamnose or xylitol.
Diauxie was evident during the fermentation of a
sugar mixture. Following the depletion of glu-
cose, P. stipitis fermented galactose, mannose, xy-
lose and cellobiose simultaneously with no no-
ticeable preceding lag period. A similar fermenta-
tion pattern was observed with C. shehatae, ex-
cept that it failed to utilize cellobiose even though
it grew on cellobiose when supplied as the sole
sugar. P. stipitis produced considerably more
ethanol from the sugar mixture than C. shehatae,
primarily due to its ability to ferment cellobiose.
In general P. stipitis exhibited a higher volumetric
rate and yield of ethanol production. This yeast
fermented glucose 30--50% more rapidly than xy-
lose, whereas the rates of ethanol production
from these two sugars by C. shehatae were simi-
lar. P. stipitis had no absolute vitamin requirement
for xylose fermentation, but biotin and thiamine
enhanced the rate and yield of ethanol production
significantly.
Offprint requests to: J. C. du Preez
Nomenclature: Iz . . . . Maximum specific growth rate, h - t ; Qp,
Maximum volumetric rate of ethanol production, calculated
from the slope of the ethanol vs. time curve, g.(l.h)-~; qp,
Maximum specific rate of ethanol production, g-(g
cells.h)-~; Yp/s, Ethanol yield coefficient, g ethanol.(g sub-
strate utilized)-~; Y,-/s, Cell yield coefficient, g biornass-(g
substrate utilized)- J; E, Efficiency of substrate utilization, g
substrate consumed. (g initial substrate)-~. 100
Applied
Biotechnology
© Springer-Verlag 1986
Introduction
In earlier reports it was shown that a strain of
Candida shehatae fermented D-xylose rapidly and
with a high ethanol yield in comparison with the
few other yeasts capable of xylose fermentation
(Du Preez and Van der Walt 1983; Du Preez et al.
1984). More recently the perfect stage of this spe-
cies, Pichia stipitis (Barnett et al. 1983), was also
shown to ferment xylose to ethanol efficiently
(Slapack et al. 1983; Bruinenberg et al. 1984;
Dellweg et al. 1984; Lindman and Bj6rling 1984;
Toivola et al. 1984; Du Preez and Prior 1985).
Apart from D-xylose, the hydrolysis of the
hemicellulose component of lignocellulosic bio-
mass also yields varying amounts of D-mannose,
D-galactose, L-arabinose, L-rhamnose, D-glucose
and cellobiose, depending on the nature of the
plant material and hydrolysis procedure (Suihko
1983). These sugars also occur in spent sulphite li-
quor with D-xylose and D-mannose as the princi-
pal sugars (Neirinck et al. 1982). Dilute acid hy-
drolysis of sugar cane bagasse, which is regarded
as one of the more economically viable lignocellu-
losic waste feedstocks for ethanol production,
yields xylose, glucose, arabinose and galactose as
the main sugars, in that order of concentration
(Du Toit et al. 1984). The extent to which these
sugars can be fermented to ethanol by the above-
mentioned yeasts is therefore of interest to the
commercial exploitation of lignocellulosic materi-
als.
This article describes the utilization and fer-
mentation of the above-mentioned seven sugars
singly and in a sugar mixture by a strain of C. she-
hatae and P. stipitis. Xylitol was also included in
the study because xylitol accumulation in the fer-
mentation broth is typical of xylose-fermenting
yeasts (Gong et al. 1983), and a yeast which can
J. C, du Preez et al.: The fermentation of hexose and pentose sugars 229

ferment xylitol or does not produce xylitol would
thus be desirable. Because limited aeration en-
hances ethanol production from xylose considera-
bly (Du Preez et al. 1984), presumably by promot-
ing the reoxidation of excessive reducing equival-
ents (Bruinenberg et al. 1984), all fermentations
were conducted under aerated conditions. The ef-
fect of biotin and thiamine on xylose fermenta-
tion by P. stipitis is also reported.
Cultivation conditions. Erlenmeyer flasks with cotton wool
plugs were incubated at 30°C on magnetic stirrers as de-
scribed previously (Du Preez et al. 1984). The fermentation of
individual carbon sources were conducted in 250 ml Eden-
meyer flasks containing 250 ml medium at a stirring speed of
700 rev. m i n - ~. For the fermentation of sugar mixtures 500 ml
Erlenmeyer flasks containing 500 ml medium and stirred at
900 r e v . m i n - I were used. The same conditions were used to
evaluate the effect of biotin and thiamine on the fermentation
of xylose by P. stipitis, and in these experiments all glassware,
etc., were rinsed with 1N HC1 or autoclaved with distilled wa-
ter prior to use. The flasks were inoculated to an initial dry cell
concentration of ca. 0.2 g. 1- t.

lnoculum. The inoculum was prepared in a similar fashion us-

Materials and methods
Microorganisms. The Candida shehatae CSIR-Y492 (CBS
2779) strain used previously (Du Preez et al. 1984) and Pichia
stipitis CSIR-Y633 (Du Preez and Prior 1985) were obtained
from J. P. van der Walt of the Council for Scientific and In-
dustrial Research, Pretoria.
Culture media. Medium CA containing casamino acids (Difco,
vitamin free), added vitamins and mineral salts as described
elsewhere (Du Preez et al. 1984) and adjusted to pH 5 was
used. The respective carbon sources were D-glucose, D-man-
nose, D-galactose, L-rhamnose, D-cellobiose, D-xylose, L-ara-
binose and xylitol at a concentration of 20 g . l - ~ or as indi-
cated in the text. Multiple substrate fermentations were con-
ducted with D-glucose, D-mannose, D-galactose, D-cellobiose
and o-xylose using 10 g.l -~ of each. The sugars were auto-
claved separately from the rest of the medium constituents.
ing a 150 ml and a 300 ml Erlenmeyer shake flask sequentially
as described elsewhere (Du Preez and Van der Walt 1983) with
20 g. 1- ~ D-xylose as carbon source, which was increased to 40
g. 1-~ in the vitamin requirement study. In evaluating the ef-
fect of biotin and thiamine, the second shake flask contained
no vitamins and a washed cell suspension was used as inocu-
lure as described elsewhere (Du Preez et al. 1985b).
Analyticalprocedures. Ethanol was determined by gas chroma-
tography and glucose and xylitol by high performance liquid
chromatography (Du Preez and Van der Walt 1983). Because
of the unsatisfactory resolution of mannose, galactose and cel-
lobiose during HPLC analysis of sugar mixtures, these assays
were supplemented with gas-liquid chromatography of trime-
thylsilyl ether derivatives of the sugars as outlined elsewhere
(Du Preez et al. 1985a) with multi-step temperature program-
ming from 160°C to 260°C. The internal standard was 10
g.1 t D-ribose. Growth was monitored with a Klett-Summer-
son colorimeter at 640 nm as well as gravimetrically by drying
centrifuged and washed samples to constant mass at 105 ° C.

Table 1. Parameters for the individual fermentation and utilization of 20 g . l - 1 each of different hexoses, pentoses and xylitol by
C. shehatae and P. stipitis

Substrate Parameter

]/ . . . . Qp, qp, Yp/s Y,-/s E, % Fermentation

h- i g- (1- h ) - 1 h- i time, h a

C. shehatae
D-Glucose 0.14 0.45 0.34 0.32 0.13 100 18
D-Mannose 0.20 0.48 0.45 0.32 0.12 100 21
D-Galactose 0.14 0.42 0.23 0.30 0.13 100 28
D-Xylose 0.14 0.47 0,33 0.37 0.10 100 28
L-Arabinose 0.04 0 0 0 0.36 > 99 211
L-Rhamnose 0 0 0 0 0 0 --
D-Cellobiose 0.04 0 0 0 0.49 75 196
Xylitol 0.11 0 0 0 0.09 6 120
P. stipitis
D-Glucose 0.23 0.70 0.39 0.40 0.18 100 17
D-Mannose 0.18 0.46 0.29 0.36 0.13 100 24
D-Galactose 0.20 0.44 0.18 0.32 0.19 100 24
D-Xylose 0.22 0.53 0.26 0.39 0.16 100 24
L-Arabinose 0.14 0 0 0 0.32 > 99 121
L-Rhamnose 0.16 0 0 0 0.40 100 143
D-Cellobiose 0.21 0.26 0.05 0.32 0.34 100 48
Xylit01 0.12 0 0 0 0.47 88 120

Time required for the maximum ethanol concentration to be reached, or until termination of the cultivation
230
Results
Fermentation o f individual carbon sources
As shown in Table 1, both yeasts fermented D-
glucose, D-mannose, D-galactose and D-xylose to
ethanol. D-cellobiose was only fermented by P.
stipitis, whereas C. shehatae utilized this disac-
charide for growth with no concomitant ethanol
production. Apart from similar volumetric etha-
nol productivities on mannose and galactose, the
ethanol yield and volumetric productivity ob-
tained with P. stipitis on the above sugars were
markedly higher than the values obtained with C.
shehatae. Whereas C. shehatae fermented glucose
and xylose at comparable rates, the volumetric
and specific rates of ethanol production from glu-
cose by P. stipitis were, respectively, about 30%
and 50% higher than the corresponding values on
xylose. Even though the specific rate of ethanol
production by C. shehatae was considerably
higher on D-mannose, the maximum volumetric
productivities on the four fermentable sugars
were of similar magnitude. No ethanol was pro-
duced from xylitol or from the two L-sugars by
either yeast, although they were utilized for
growth. However, C. shehatae lacked the ability
~ 20
Z o
~2c
<8
mO
~5 '
FERMENTATION TIME , h
Fig. 1. Fermentation of a sugar mixture by P. stipitis (A) and
C. shehatae (B). Symbols: [3, Ethanol; ©, D-Glucose; 0 ,
D-Mannose plus D-galactose; A, D-Xylose; A D-Cellobiose
Table 2. Parameters for a multiple substrate fermentation (10 g.1-~ each of D-glucose, D-mannose, D-galactose, D-xylose and
o-cellobiose) by C. shehatae and P. stipitis
Yeast Parameter
species
].z. . . . Qp,
h -1 g.(1.h) -1
C. shehatae 0.20 0.50
P. stipitis 0,22 0.73
J . C . du Preez et al.: The fermentation of hexose and pentose sugars
to utilize L-rhamnose as the sole carbon source.
Furthermore, P. stipitis utilized 88% of the availa-
ble xylitol within 120 h for growth, whereas C.
shehatae utilized less than 6% within the same
time period. Only C. shehatae produced xylitol
( < 0.5 g. 1-1) from D-xylose.
Fermentation o f sugar mixtures
The five sugars which proved to be fermentable
by P. stipitis, namely D-glucose, D-mannose, D-
galactose, D-xylose and D-cellobiose, were used in
a mixture in order to determine the utilization
pattern that could be expected during the fermen-
tation of a lignocellulosic hydrolysate. The analy-
tical procedures used failed to distinguish be-
tween galactose and mannose, therefore the joint
total concentration of these two hexoses are given
in Figs. 1A and B. As shown in these two figures,
both yeasts utilized glucose preferentially, and the
presence of glucose appeared to retard the fer-
mentation of mannos~ and galactose more se-
verely in the case of C.'shehatae than with P. stipi-
tis. With P. stipitis xy[0se and cellobiose utiliza-
tion commenced only after the glucose was de-
pleted, thereafter the simultaneous fermentation
of galactose, mannose~ xylose and cellobiose oc-
curred (Figl 1A). Ethanol production from cello-
biose continued after exhaustion of the xylose. A
similar pattern was observed with C. shehatae
(Fig. 1B), except that even after a fermentation
time of 120 h no cellobiose utilization was evident
(the linear regression correlation coefficient of the
cellobiose concentration vs. time curve was
0.085).
The maximum volumetric rate of ethanol pro-
duction by P. stipitis was considerably higher than
that of C. shehatae (Table 2), and was similar to
the value obtained with glucose (Table 1). The
ethanol yield coefficient (based on sugars con-
sumed) was similar for both yeasts (Table 2), but
the failure of C. shehatae to ferment cellobiose is
reflected by the 18.6 g. 1-1 ethanol produced by P.
qp, Yv/~ Y,/.~ E, % Ethanol,
h -1 g.1-1
0.40 0.35 0.08 79 13.4
0.49 0.37 0.13 100 18.6
 J. C. du Preez et al.: The fermentation of hexose and pentose sugars 231

stipitis as opposed to the 13.4 g-1-1 by C. sheha-
tae. The ethanol yields were as anticipated from
the yield values obtained during the fermentation
of individual sugars. At the end of the 48 h fer-
mentation C. shehatae had produced 1.9 g-1-1 xy-
litol. With P. stipitis a small amount of xylitol
(1 g.1-1) accumulated after a 31 h fermentation
period, but this was completely assimilated at
48 h.
much slower in the total absence of vitamins (Fig.
2 and Table 3). Although all the xylose was uti-
lized in the latter case, the required fermentation
time was more than double that obtained with vi-
tamins, and the ethanol yield was also signifi-
cantly lower.
Discussion

Galactose is poorly fermented by Pachysolen tan-

Vitamin requirements of P. stipitis
During the course of this investigation it was dis-
covered that biotin and thiamine were the only
two vitamins required by C. shehatae CSIR-Y492
for the optimal fermentation of xylose (Du Preez
et al. 1985b). Thus the other vitamins (pyridoxine,
myo-inositol, nicotinic acid, pantothenate and p-
aminobenzoic acid) were omitted from the culture
medium to establish whether the vitamin require-
ments of P. stipitis were similar. Whereas little dif-
ference could be discerned in the fermentations
where the medium contained all seven vitamins or
only biotin and thiamine, the fermentation was
'7 15
=.
b~lC
Oi 30 60 90 J
FERMENTATION TIME , h
Fig. 2. Effect of vitamin supplementation on ethanol produc-
tion from 40 g. 1- ' D-xylose by P. stipitis. Symbols : Q, No vi-
tamins; O, Biotin and thiamine; A, Biotin, thiamine, pyridox-
ine, myo-inositol, nicotinic acid, pantothenate and p-amino-
benzoic acid
nophilus (Maleszka et al. 1982a; Neirinck et al.
1982; Debus et al. 1983; Deverell 1983). In this re-
spect both C. shehatae and P. stipitis have the ad-
vantage that their ethanol yield and maximum
volumetric productivity on galactose are compar-
able with the values for the other fermentable su-
gars (Table 1), even though their maximum spe-
cific rate of ethanol production is lower on galac-
tose.
Yeasts, in general, are unable to ferment L-
arabinose to ethanol (Barnett 1976). This was con-
firmed for our two strains (Table 1) as well as for
P. tannophilus (Neirinck et al. 1982; Schneider et
al. 1983), but Dekker (1982) claimed ethanol for-
mation from this pentose using a different strain
of P. tannophilus.
P. stipitis CSIR-Y633 possesses the seemingly
rare ability of fermenting both xylose and cello-
biose efficiently in comparison with other yeasts.
Other investigators have shown that yeasts which
can ferment cellobiose to ethanol have a poor ca-
pacity for xylose fermentation (Maleszka et al.
1982b), and of the type strains of 200 yeast species
tested by Toivola et al. (1984) only Candida tenuis
was capable of xylose fermentation as well as a
weak ethanol production from cellobiose. Al-
though Dellweg et al. (1985) reported good etha-
nol yields from xylose, mannose and galactose
with their strain of P. stipitis, it produced only
0.4 g.1-1 ethanol from 1% cellobiose. Our strain
of P. stipitis even compares favourably with Can-

Table 3. Effect of vitamin supplementation on the fermentation of 40 g - l - ' D-xylose by P. stipitis

Vitamins " Parameter

added
It .... Qp, qp, Yp/s Y,-/s E, % Fermentation
h- ' g-(l- h ) - t h-1 time, h a

None O. 16 O. 17 O. 10 0.33 0.14 100 99

Biotin & thiamine 0.16 0.65 0.35 0.41 0.14 100 36
Sevenb 0.17 0.64 0.34 0.41 0.15 100 36

a Time required for the maximum ethanol concentration to be reached

b Biotin, thiamine, pyridoxine, myo-inositol, nicotinic acid, pantothenate and p-aminobenzoic acid
232
dida wickerhamii, one of the more efficient cello-
biose-fermenting yeasts (Freer and Detroy 1983;
Kilian et al. 1983b). Though the ethanol yield
coefficient of 0.34 obtained with P. stipitis on cel-
lobiose was lower than the values of 0.35 to 0.41
reported for C. wickerhamii (Kilian et al. 1983b),
its rate of ethanol production from cellobiose was
considerably more rapid (Table 1) than that of C.
wickerhamii, which required a fermentation time
of 125 h for the fermentation of 20 g . l - I cello-
biose with a maximum volumetric ethanol pro-
ductivity of only 0.12g (l-h) -1 (Kilian et al.
1983b). Furthermore, C. wickerhamii produced
ethanol from cellobiose only under non-aerated
conditions (Kilian et al. 1983b).
None of the xylose-fermenting yeasts tested so
far has the ability to produce significant amounts
of ethanol from xylitol. This was the case with our
two strains (Table 1), the P. stipitis strains of Dell-
weg et al. (1985), as well as with 15 yeast taxa sur-
veyed by Maleszka and Schneider (1982).
Thus P. stipitis can ferment all the major su-
gars produced by the hydrolysis of plant biomass,
namely D-glucose, D-mannose, D-galactose, D-xy-
lose and D-cellobiose. The remaining sugars, L-
arabinose and L-rhamnose, usually occur at very
low levels (Neirinck et al. 1982), although L-arabi-
nose is more abundant in certain hemicellulose
sources (Finn et al. 1984).
Particular problems can be encountered when
attempting to ferment sugar mixtures such as
those found in lignocellulosic hydrolysates. For
example, Slapack et al. (1983) reported that after
the depletion of glucose and mannose, Candida
tropicalis consumed the produced ethanol before
the onset of galactose and xylose utilization, fol-
lowed by the concurrent uptake of these three
substrates. Diauxie can also cause a considerable
lag period during the fermentation of sugar mix-
tures by P. tannophilus (Detroy et al. 1982) and
appears to be a common phenomenon during the
fermentation of xylose or cellobiose in the pres-
ence of glucose (Gond6 et al. 1982; Kilian et al.
1983a; Slapack et al. 1983). In some cases a total
diauxie can occur during the fermentation of a
glucose-cellobiose mixture (Gond6 et al. 1982)
such as that observed with C. shehatae (Fig. 1B),
but P. stipitis does not suffer from such a degree
of diauxie so that the fermentation of xylose and
cellobiose commenced immediately after glucose
depletion (Fig. 1A).
As was the case with C. shehatae (Du Preez et
al. 1985b), biotin and thiamine enhanced xylose
fermentation by P. stipitis. However, the fermen-
tation performance of P. stipitis was much less de-
J. C. du Preez et al.: The fermentation of hexose and pentose sugars
pendent on these two vitamins, because in their
absence this yeast still exhibited a maximum volu-
metric rate of ethanol production of
0.17 g.(1.h) -1 and an ethanol yield coefficient of
0.33 (Table 3), in contrast with the corresponding
values of 0.03 g.(1-h) -1 and 0.015 with only
23.9% xylose uptake obtained with C. shehatae
(Du Preez et al. 1985b). This is in agreement with
the results of Dellweg et al. (1984), who reported
that P. stipitis has no requirement for growth fac-
tors, although yeast extract stimulated growth.
Acknowledgements. The authors thank P. J. Botes and Marieta
Cawood for their competent technical assistance. The finan-
cial support of the Council for Scientific and Industrial Re-
search (Foundation for Research Development) and of the
Central Research Fund, University of the O. F. S., is gratefully
acknowledged.
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