1 s2.0 S0141022999001556 Main

Enzyme and Microbial Technology 26 (2000) 87107
Review
Factors affecting the fermentative lactic acid production from

renewable resources1
Karin Hofvendahl, Barbel HahnHagerdal*
Department of Applied Microbiology, Lund Institute of Technology/Lund University, P.O. Box 124, SE-221 00 Lund, Sweden
Received 15 February 1999; received in revised form 11 August 1999; accepted 20 August 1999
Abstract
Parameters affecting the fermentative lactic acid (LA) production are summarized and discussed: microorganism, carbon- and nitrogensource, fermentation mode, pH, and temperature. LA production is compared in terms of LA concentration, LA yield and LA productivity.
Also by-product formation and LA isomery are discussed. 2000 Elsevier Science Inc. All rights reserved.
Keywords: Lactic acid; Lactic acid bacteria; Lactococcus lactis; Lactobacillus; Starch hydrolysate; Lignocellulose; Whey; Nutrient; Fermentation mode;
Simultaneous saccharification and fermentation process; Cell recirculation; pH; Temperature; Productivity; Yield; By-product formation; Optical isomers;
Process optimization
1. Introduction
Lactic acid (LA) is a versatile chemical, used 1) as an
acidulant, flavor and preservative in the food, pharmaceutical, leather and textile industries, 2) for the production of
base chemicals, 3) and for polymerization to biodegradable
poly LA (PLA) [1,2]. LA exists as two optical isomers, Dand L-LA. Both isomeric forms of LA can be polymerized
and polymers with different properties can be produced
depending on the composition. Of the 80 000 tonnes of LA
produced worldwide every year about 90% are made by LA
bacterial fermentation and the rest is produced synthetically
by the hydrolysis of lactonitrile. Fermentative production
has the advantage that by choosing a strain of LA bacteria
(LAB) producing only one of the isomers, an optically pure
product can be obtained, whereas synthetic production always results in a racemic mixture of LA. It is also possible
to use renewable resources as substrates, such as starch and
cellulose in fermentative production. Renewable resources
do not give any net contribution of carbon dioxide to the
atmosphere, as do the limited oil- and fossil-fuel-based
sources. Cellulose, hemicellulose and starch are the most abundant compounds in the world, and when hydrolyzed to mainly
glucose they are fermentable by a number of microorganisms.
1
This paper is based on the Ph.D thesis of Karin Hofvendahl (1998)

* Corresponding author. Tel.: 46-46-222-8428; fax: 46-46-222-4203.
E-mail address: Barbel.Hahn-Hagerdal@tmb.lth.se (B. HahnHagerdal)
Hemicellulose, in contrast to starch and cellulose, contains

pentoses, which give rise to by-products such as acetate and
ethanol, decreasing the LA yield. Fermentative LA production
from renewable resources comprises the following steps: pretreatment of substrate including hydrolysis to sugars, fermentation of sugars to LA, separation of bacteria and solid particles
from the broth, and purification of LA.
The names of genera and strains used in this review were
updated to concur with our knowledge on LAB today, and
might therefore differ from those used by the authors cited.
The changes made are listed in Table 1. Today LAB consist
of the Gram-positive genera: Carnobacterium, Enterococcus (Ent), Lactobacillus (Lb), Lactococcus (Lc), Leuconostoc (Leu), Oenococcus, Pediococcus (Ped), Streptococcus
(Str), Tetragenococcus, Vagococcus, and Weissella [4].
LAB are cocci, with the exception of lactobacilli and carnobacteria which are rods, unable to synthesize ATP by
respiration, and that have LA as the major end product from
energy-conserving fermentation of sugars. Most LAB are
facultatively anaerobic, catalase negative, nonmotile and
nonspore forming. They have high acid tolerance and survive pH 5 and lower. Their acid tolerance gives them a
competitive advantage over other bacteria. The optimal
temperature for growth varies between the genera from 20
to 45C [5,6]. Most of them are considered GRAS (generally regarded as safe), but some strains of e.g. streptococci
are pathogenic. All LAB genera belong to the Clostridium
0141-0229/00/$ see front matter 2000 Elsevier Science Inc. All rights reserved.
PII: S 0 1 4 1 - 0 2 2 9 ( 0 0 ) 0 0 1 5 5 - 6
88
K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107
Nomenclature
ATP
Ent
LA
LA/tot
LAB
Lb
Lc
LDH
Leu
NADH
PDH
Ped
PFL
PLA
QV
SSF
Str
WWF
YLA/tot
adenosine triphosphate
Enterococcus
lactic acid
La per total products (g/g)
lactic acid bacteria
Lactobacillus
Lactococcus
lactate dehydrogenase
Leuconostoc
nicotinamide adenine dinucleotide, reduced
form
pyruvate dehydrogenase
Pediococcus
pyruvate formate lyase
poly-lactic acid
maximum columetric productivity (g/lh)
simultaneous saccharification and fermentation
Streptococcus
whole wheat flour
yield of LA per substrate provided (g/g)
subdivision of the Gram-positive bacteria; i.e. the GC

content of the DNA is 55% [4].
LAB have complex nutrient requirements, due to their
limited ability to synthesize B-vitamins and amino acids [7].
Therefore they are naturally found in nutrient-rich environments such as in plants, milk, and inside the human and
animal bodies. A complex natural environment renders a
microorganism able to metabolize many different carbohydrates [8]. LAB have been used by humans for the fermentation of food and feed products since ancient days, and
today their major applications are still in the food and feed
industry, e.g. in the production of dairy products, pickles,
meat and wine. The present technical applications of LAB
include the production of dextran from sucrose by Leu.
mesenteroides [9], the production of nisin by Lc. lactis spp.
lactis and the production of LA for different applications,
see above. It has also been suggested that LAB could be
used as oral vaccine vectors [10].
LAB ferment sugars via different pathways resulting in
homo-, hetero-, or mixed acid fermentation (Fig. 1). Homofermentation gives only LA as the end product of glucose
metabolism, and the EmbdenMeyerhofParnas pathway is
used (Fig. 1A) [11,12]. In heterofermentation equimolar
amounts of LA, carbon dioxide and ethanol or acetate are
formed from glucose via the phosphoketolase pathway (Fig.
1B) [13,14]. The ratio of ethanol and acetate formed is dependent on the redox potential of the system [15]. This pathway is
used by facultative heterofermenters, such as Lb. casei, for the
fermentation of pentoses, and for the fermentation of hexoses
and pentoses by obligate heterofermenters, organisms such as
Leu. [15]. According to Kandler, all LAB except lactobacilli of
type I (e.g. Lb. delbrueckii) are able to ferment pentoses, i.e.
they are facultative heterofermenters [15].
Mixed acids are formed by homofermenters such as
lactococci during glucose limitation [16], and during growth
on other sugars, e.g. Lc. lactis on maltose [1722], lactose
[23,24] and galactose [23,24], or at increased pH and decreased temperature [25]. Ethanol, acetate and formate are
formed in addition to LA. The homofermentative pathway
is used, but the difference is in the metabolism of pyruvate,
Table 1
Names of organisms changed in the material
Before change
After change
Reference
Lb. arabinosus
Lb. bulgaricus
Lb. casei sp. rhamnosus
Lb. cellobiosus
Lb. xylosus
Str. diacetylactis
Str. cremoris
Str. faecalis
Str. faecium
Str. lactis
Str. salivarius sp. thermophilus
Lb. delbrueckii NRRL B-445
Lb. casei sp. casei ATCC 393/DSM 20011
Lb. delbrueckii ATCC 9649
Lb. salivarius NRRL B-1950
Lb. casei DSM 20244
Lb. casei ATCC 7469/IFO 3425
Lb. casei ATCC 11443
Lb. plantarum
Lb. delbrueckii sp. bulgaricus
Lb. rhamnosus
Lb. fermentum
Lc. lactis sp. lactis
Lc. lactis sp. lactis biovar diacetylactis
Lc. lactis sp. cremoris
Ent. faecalis
Ent. faecium
Str. thermophilus
Lb. rhamnosus ATCC 10863
Lb. zeae ATCC 393
Lb. delbrueckii sp. delbrueckii ATCC 9649
Lb. salivarius sp. salivarius ATCC 11742
Lb. paracasei sp. paracasei DSM 20244
DSM, ATCC
166
167, 168
169
170
170
170
171
171
170
172
ATCC
168
ATCC
ATCC, NRRL
167, DSM
ATCC
ATCC
When different names of the same strain were used in different studies, they have been harmonized to one, where the ATCC No. is the highest in hierarchy,
followed by the DSM No.
89
Fig. 1. Catabolic pathways in lactic acid bacteria. Homofermentation (A), heterofermentation (B) and mixed acid fermentation (C). P phosphate, BP
bisphosphate, LDH lactate dehydrogenase, PFL pyruvate formate lyase, and PDH pyruvate dehydrogenase.
which in addition to give LA is also metabolized into

formate and acetyl-CoA by pyruvate formate lyase (PFL)
(Fig. 1C). In the presence of oxygen PFL is inactivated [26],
and an alternative pathway of pyruvate metabolism becomes active via pyruvate dehydrogenase (PDH), resulting
in the production of carbon dioxide, acetyl-CoA and NADH
[27]. LAB are also capable of forming other products, e.g.
flavors such as diacetyl and acetoin or bacteriocins, which is
not further discussed in this review.
Numerous processes of fermentative LA production have
been patented. They include the fermentation of industrial
starch waste, e.g. potato, with a mixture of five Lb. strains
in a continuous simultaneous saccharification and fermentation process (SSF) with electrodialytic purification [28],
the use of whey permeate in a continuous process with cell
recirculation and electrodialysis [29], the use of lignocellulosic material and a recombinant strain of Lb. able to ferment xylose [30], and the recovery of municipal solid waste
to solids and LA by fermentation [31]. Several patents claim
the production of optically pure D- [3234] or L-LA [32,35,
36]. In one patent, the development of a LA-tolerant strain
of Lb. delbrueckii is described [37]. Methods for the purification of LA from the fermentation broth with electrodialysis [38,39], membrane separation [40], and esterification
[41] have also been patented.
2. Effectiveness of various processes

The selection criteria for the material presented in this
review were fermentative LA production by LAB for the
purpose of industrial processes. Therefore, papers with a
metabolic approach have not been included. Many parameters influence the efficiency of a fermentation process,
several of which are discussed below. Due to the oxygen
tolerance of LAB most fermentations were performed without aeration control. However, anaerobic conditions were
preferred to aerobic or microaerophilic conditions.
The effectiveness of a process can be measured as the
concentration of LA produced, as the yield of LA based on
substrate and as the productivity (LA production rate). The
yields presented in Tables 210 were calculated as g LA per
g substrate provided (YLA/tot), and the productivities are
given as the maximum volumetric productivity (QV) in g
LA per liter per hour. The overall most effective way to
produce LA in terms of concentration was to continuously
remove the acid by extraction, resulting in at most 771 g/l
[42]. The highest yields, 3.1 g/g, were achieved from whey
using electrodialysis [43] or a semicontinuous fermentation
mode [44]. The highest values of QV, 52-144 g/(lh), were
obtained by recirculating the cells, thus increasing the cell
density [45 48].
90
Table 2
Comparison of different strains for lactic acid production
Organism
Lb. casei NRRL B-441
Lb. delbrueckii sp. lactis ATCC 8000
Lb. delbrueckii sp. lactis DSM 20073
Lb. delbrueckii mutant DP3
Lb. delbrueckii mutant DP3, 19
Lb. zeae ATCC 393
Lb. rhamnosus DSM 20024
Lb. zeae ATCC 393
Lb. coryniformis sp. torquens ATCC 25600
Lb. amylovorus ATCC 33622
Lb. delbrueckii sp. bulgaricus ATCC 11842
Lb. plantarum ATCC 14917
Lb. delbrueckii sp. lactis 447
Lb. rhamnosus CCM 1753
Lb. delbrueckii sp. bulgaricus AU
Lb. kefir
Lb. acidophilus R
Lb. casei SU No 22
Le. lactis WS 1042
Str. thermophilus
Lb. delbrueckii sp. bulgaricus 5085
Lc. lactis sp. lactis 2432
Str. thermophilus
Lc. lactis sp. cremoris 2487
Lb. casei SU No 22
Lc. lactis WS 1042
Str. thermophilus
Lb. acidophilus CRL 640
Lb. delbrueckii sp. bulgaricus CRL 870
Str. thermophilus CRL 807
Lc. lactis sp. cremoris SBT 1306
Lc. lactis sp. lactis ATCC 19435
Lc. lactis sp. lactis AS211
Lb. plantarum NRRL B-787
Lb. pentosus NRRL B-227
Lb. plantarum USDA 422
Lc. lactis sp. lactis NRRL B-4449
Ent faecium
Lb. plantarum
Ped. acidilacti
Substrate
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
hydr barley flour
hydr barley flour
sorghum
sorghum
lignocellulose hydr
lignocellulose hydr
molasses
molasses
molasses
paneer whey
paneer whey
whey
whey
whey permeate
whey permeate
whey permeate
whey permeate
whey permeate
whey permeate
whey permeate
whey permeate
whey permeate
deproteinised whey
deproteinised whey
whey permeate
whey permeate
whey permeate
skim milk
skim milk
skim milk
lactose
lactose
hydr wheat flour
hydr wheat flour
hydr wheat flour
hydr wheat flour
solid waste
solid waste
solid waste
solid waste
solid waste
solid waste
solid waste
solid waste
hydr cod corn syrup
hydr cod corn syrup
hydr cod corn syrup
Temp
C
LA
g/l
g/g
YLA/tot
41
41
45
45
45
45
45
45
45
36
36
36
36
36
37
37
42
42
37
37
47
47
47
43
43
32
32
42
42
30
42
42
37
40
30
30
32
32
42
42
30
37
37
37
37
30
30
30
37
45
32
32
32
32
32
32
32
32
37
37
37
82
68
83
82
58
77
68
28
28
21
22
24
37
39
93
120
0.91
0.76
0.83
0.82
0.48
0.64
0.57
0.93
0.92
0.71
0.74
0.80
0.98
0.98
0.52
0.67
55
37
20
26
18
9.8
8.6
16
11
18
15
8.3
7.9
15
30
30
37
37
20
15
19
16
9.0
14
12
8.5
21
80
96
95
106
18
17
19
18
18
21
18
18
6.6
11
17
13
0.85
0.74
0.45
0.58
0.40
0.20
0.17
0.32
0.22
0.50
0.41
0.21
0.18
0.35
0.71
0.71
0.88
0.88
0.39
0.30
0.47
0.38
0.20
0.38
1.5
0.76
0.77
0.82
0.11
0.42
0.46
0.43
0.43
0.51
0.43
0.43
0.16
0.45
0.70
0.51
Qv
g/(lh)
Ref
5.6
3.5
146
146
32
32
49
49
49
51
51
173
173
173
50
50
92
92
100
100
113
113
89
89
89
174
174
129
129
137
137
137
140
140
52
52
52
52
175
175
139
139
139
133
133
133
53
53
18
18
18
18
111
111
111
111
111
111
111
111
176
176
176
0.72
1.7
1.2
8.0
11
4.0
2.6
2.0
1.5
4.5
2.0
5.9
4.0
2.1
1.9
1.5
4.6
2.4
2.0
1.5
6.0
4.4
2.3
5.6
4.1
3.0
1.7
1.6
0.56
0.10
0.10
0.11
Abbreviations: LA lactic acid; YLA/tot yield of g LA per g substrate provided; Qv maximum volumetric LA productivity in g LA per l per h; Ref
reference; hydr hydrolysed.
91
3. Parameters affecting efficiency

3.1. Microorganism
Strains of lactobacilli were compared with regard to the
fermentation of various sugars (Table 2). Strains giving the
highest LA concentration and yield usually also showed the
highest productivity, but in some reports different strains
gave the highest yields and productivities [49 51]. Lb.
delbrueckii emerges as the most efficient strain.
On lactose, including whey and milk, Str. thermophilus
was in most studies superior to Lb. delbrueckii spp. bulgaricus and Lc. lactis, but not to Lb. acidophilus (Table 2). Lc.
lactis, on the other hand, gave higher LA concentrations and
yields than Lb. rhamnosus [52,53], and sometimes also a
higher productivity [53].
In wheat flour hydrolysate Lc. lactis showed the highest
productivity, whereas Lb. delbrueckii spp. delbrueckii resulted in the highest LA concentration and yield (Table 2)
[18]. Generally the temperature used was the same for all
strains, but in some studies it was adjusted to the optimum
for each organism.
Metabolic engineering and traditional strategies for mutation and selection can be used to alter the properties of
an organism. Lb. delbrueckii has been subjected to mutagenesis to enhance its tolerance to LA (Table 2) [49].
The mutants resulted in better LA production than the
wild type.
3.2. Carbon source
A number of different substrates have been used for the
fermentative production of LA by LAB. The purest product
is obtained when a pure sugar is fermented, resulting in
lower purification costs. However, this is economically unfavorable, because pure sugars are expensive and LA is a
cheap product. Instead waste products from agriculture and
forestry are utilized.
3.2.1. Whey
The most common substrate for fermentative LA production is whey [54 59], a waste product from cheese
production normally used as animal feed [60]. It contains
proteins, salts, and lactose [60]. Whey can be hydrolyzed to
glucose and galactose [61,62], deproteinised by ultrafiltration [63 66], and demineralised [67]. Whey has been
supplemented with yeast extract [68 71], peptone [72,
73], milk powder [33,74], soy flour [36], or corn steep
liquor [75]. The strain most used for LA production from
whey was Lb. delbrueckii spp. bulgaricus [76 78], but in
many studies Lb. helveticus [79 81] or Lb. casei [82,83]
were utilized.
3.2.2. Molasses
Molasses, a by-product of the sugar manufacturing
process, is used as an animal feed, and for ethanol and
Fig. 2. A process of lactic acid production from wheat flour with simultaneous (SSF) (A) and separate (B) saccharification and fermentation. Process parameters according to references [18,20,106].
yeast production [60]. In addition it could be used for LA

production [84 89]. The most abundant sugar is sucrose,
the high concentration of which makes the viscosity of
the liquid high [60]. Lb. delbrueckii has generally been
used.
3.2.3. Starch
Another common substrate for LA production is starch
from crops or wastes [18,90 92]. It has to be hydrolyzed to
glucose and maltose to be fermentable by LAB. Starch from
various origins has been used for LA production, including
wheat [20,93], corn [94,95], cassava [96], potato [97,98],
rice [93], rye [35], sorghum [99,100], and barley [95,101].
In some studies, starch either remained untreated or was
liquefied/gelatinized and then fermented by an amylaseproducing organism such as Lb. fermentum [98], Lb. amylovorus [102,103], or Lb. amylophilus [104,105]. Amylases
can also be added to hydrolyze the starch and LAB fermented the produced glucose to LA [99,106,107]. In one
study, amylase was produced by Aspergillus awamori and
Lc. lactis made LA from the glucose produced [108]. LAB
used for LA production from starch include: Lb. casei [92],
Lb. plantarum [109], Lb. delbrueckii [97,107], Lc. lactis
[18,108], and Lb. helveticus [28].
In the above mentioned reports hydrolysis was performed simultaneously with fermentation (SSF) (Fig. 2A).
Another solution is to use pre-hydrolyzed starch as a substrate [20,97]. This is an advantage if the saccharifying
enzymes and the bacterium have different temperature and
pH optima. This was the case in the fermentation by Lc.
lactis of wheat starch hydrolyzed by a commercial enzyme
preparation composed of - and gluco-amylase and a protease, with 30C, pH 6 and 55C, pH 5 as optima respectively (Fig. 2B) [18]. Starch was often supplemented with
nutrients, mostly in the form of yeast extract [18] or peptone
[102]. In a few studies, the starchy material was considered
to be sufficient as nutritional source [18,20,92,95,97,98,104,
106,109]. In SSF of whole wheat flour (WWF) with Lc.
92
Table 3
Influence of carbon source and substrate treatment on lactic acid production
Organism
Substrate and treatment
LA
g/l
YLA/tot
g/g
Qv
g/(lh)
Ref
Lb. amylophilus ATCC 49845
glucose
hydr corn starch
cassava starch
corn starch
potato starch
rice starch
wheat starch
raw corn starch
liq corn starch
liq barley starch glucoam
liq barley starch glucoam alpha
barley flour
barley flour glucoam
hydr newspaper
hydr pure cellulose
glucose
lactose
glucose
cellobiose
xylose
glucose
fructose glucose
sucrose
glucose
lactose
xylose
21
33
4.8
10
4.2
7.9
7.8
45
55
112
162
36
114
24
53
35
37
56
32
41
87
94
85
58
40
14
0.95
0.73
0.48
1.0
0.42
0.79
0.78
0.82
1.0
0.68
0.87
0.20
0.63
0.48
0.53
0.85
0.82
2.8
1.6
2.1
0.87
0.94
0.85
0.85
0.75
0.70
1.6
0.88
0.69
1.2
0.14
0.86
1.2
8.6
20
104
11
18
42
95
91
46
27
90
40
6.0
7.4
6.8
0.55
0.36
0.84
0.95
0.91
0.92
0.54
1.8
0.70
0.60
0.74
0.68
0.51
0.69
0.59
0.74
0.14
0.43
0.30
0.30
0.35
0.54
0.37
0.57
0.43
0.62
0.40
0.66
0.42
0.60
0.49
0.46

Lb. amylovorus NRRL B-4542
Lb. delbrueckii IFO 3534
Lb. delbrueckii sp. bulgaricus CBS 743.84
Lb. delbrueckii sp. bulgaricus CNRZ 369
Lb. delbrueckii sp. delbrueckii

Lb. DSM 20605 MONT4, plasmid
X1 XK reg
Lb. helveticus sp. milano
Lb. paracasei No 8
Lb. pentosus
Lb. plantarum
xylose glucose
glucose
maltose
glucose
sweet sorghum
glucose
xylose
glucose xylose
hydr wood
glucose
galactose
mannose
hydr cellulose: glu,
glucose
galactose
mannose
xylose
hydr soluble starch
hydr tapioca starch
hydr tapioca flour
glucose
galactose
mannose
glucose
galactose
mannose
glucose
galactose
man, xyl, gal

6.9
5.9
7.4
1.4
man, xyl, gal
15
15
17
5.4
3.7
5.7
man, xyl, gal
6.2
4.0
6.6
man, xyl, gal
6.0
4.9
man, xyl, gal
93
102
101
1.1
2.0
0.85
0.60
92
114
33
159
5.5
5.5
6.2
118
177
30
4.2
5.0
5.6
10
2.4
0.59
4.0
1.3
120
99
67
111
111
96
111
111
111
93
Table 3 (continued)
Organism
glucose
galactose
mannose
hydr cellulose: glu, man, xyl, gal
glucose
galactose
mannose
xylose
glucose
fructose
glucose fructose
sucrose
alpha-cellulose
switch grass cellulose
glucose
starch
xylose
xylose glucose
glucose
xylose
glucose
maltose
hydr wheat flour, 3 l enz/g starch
glucose
galactose
mannose
xylose
Lb. plantarum USDA 422

LBM5 mix of 5 Lb strains
Lc. lactis IO-l JCM 7638
Lc. lactis sp. lactis NRRL B-4449
LA
g/l
7.3
4.7
8.3
5.2
3.1
6.2
1.3
17
14
16
15
45
28
99
90
23
28
36
13
4.9
3.2
75
75
90
87
6.6
2.8
5.8
1.8
YLA/tot
g/g
Qv
g/(lh)
0.73
0.47
0.83
0.43
0.52
0.31
0.62
0.13
0.43
0.86
0.71
0.81
0.73
111
111
88
0.96
0.50
0.90
0.82
0.45
0.70
1.0
0.42
0.86
0.70
0.78
0.83
0.98
0.93
0.66
0.28
0.58
0.18
0.16
Ref
112
28
0.30
2.2
3.6
0.37
2.5
1.0
1.2
0.85
1.5
1.7
160
161
25
196
111
Abbreviations as in Table 2 and: liq liquefied, glucoam glucoamylase; alpha alpha-amylase; Xl gene encoding xylose isomerase; XK gene
encoding xylulokinase; reg regulating gene xy/R; glu glucose; man mannose; xyl xylose; gal galactose; enz enzyme.
lactis, nutrient limitation occurred [106]. This was overcome by adding proteasereleasing nutrients present in the
flour or by increasing the flour concentration.
3.2.4. Lignocellulosic materials
Lignocellulosic materials have also been used for the
production of LA in similar ways as starch. It consists
mainly of the hexoses glucose, galactose, and mannose and
the pentoses xylose and arabinose, and has to by hydrolyzed
to monomers to be fermentable [110]. The lignocellulosic
materials included waste paper [111], plant material [112,
113], and wood [67]. SSF of cellulose has been studied with
Lb. delbrueckii [114] and Lb. rhamnosus [115,116]. A
mixed culture with the cellulase-producing fungus Trichoderma reesei has also been reported [117].
3.2.5. Effectiveness
Comparing different carbon sources showed that glucose resulted in higher LA concentrations and yields
compared with other sugars (Table 3). Xylose, galactose,
arabinose, lactose, fructose, and hydrolyzed cellulose

were less effective. However, mannose gave a higher LA
concentration and higher or the same yield as glucose
[111]. The simultaneous utilization of glucose and fructose was more effective than glucose alone with Lb.
delbrueckii [118], whereas the opposite was true for Lb.
rhamnosus [88]. When comparing glucose and maltose
fermentation by Lc. lactis, glucose resulted in higher LA
concentration, yield and productivity than maltose (Table
3) [21,25,119]. The main reason is that on maltose Lc.
lactis gives mixed acids, whereas on glucose the fermentation is almost homofermentative [25]. Lb. helveticus,
on the other hand, showed higher values on maltose than
on glucose (Table 3) [120]. Amylase addition to starch
also resulted in enhanced LA production (Table 3) [20,
92,101], by increasing the concentration of sugar and
nutrients [25]. Also, increased amylase concentrations
increased the LA concentration, yield and productivity
(Table 3) [106].
The concentration of the substrate had no large effect on
yield and productivity [25].
94
Table 4
Influence of nutrient type and concentration on lactic acid production
Organism
Nutrients
LA
g/l
YLA/tot
g/g
Lactobacterium delbrueckii sowjeskij
sucrose
sucrose
whey
whey
glucose
glucose
hydr wheat flour, SSF
glucose
glucose
hydr potato
hydr potato waste
lactose
whey
whey permeate
whey permeate
whey permeate
hydr whey
hydr whey, clarified
whey, UF
glucose
hydr rye
whey
whey
sweet sorghum
sweet sorghum
glucose
glucose
sorghum
sorghum
sorghum
hydr molasses, SSF
hydr molasses, SSF
glucose
glucose
hydr wood
hydr wood, SSF
glucose
glucose
glucose
glucose
glucose
soy
soy molasses
potato starch
potato starch
glucose, cont substr feed
glucose, pH dep substr feed
glucose
glucose
glucose
unhydr wheat flour glucose
unhydr wheat flour
hydr wheat flour
20% YE
1% YE 0.5% pep
21
18
13
20
93
96
18
26
58
67
106
109
100
93
17
8.9
36
36
40
44
41
37
93
92
9.8
14
106
91
45
46
1.1
0.90
0.22
0.34
0.93
0.96
0.11
0.18
0.48
0.56
0.82
0.91
1.0
0.78
0.38
0.20
0.75
0.75
0.83
Lb. acidophilus R
Lb. delbrueckii sp. lactis ATCC 12315
Lb. helveticus ATCC 15009
Lb. helveticus Milano
Lb. helveticus sp. milano
Lb. IMET 11466

Lb. kefir
Lb. paracasei No 8
Lb. pentosus


Lc. lactis IFO 12007 Aspergillus awamori IFO 4033
Lc. lactis JO-l JCM 7638

YE
malt sprouts
YE
YE
MRS 1% YE
MRS 3% YE
YE
CSL
MRS
YE
YE higher conc
YE pep
YE
CSL
CSL
MRS
5% MRS
YE
YE pep
YE
MRS
5% vetch juice
15% vetch juice
25% vetch juice
YE pep
0.25% YE 0.5% trp
0.5% YE 1% g/trp
YE pep
YE pep
0.4% YE
0.8% YE
0.2% YE
1% YE
YE
YE trp
0.3% YE
0.5% YE
1% YE
YE
YE
YE
80
16
14
57
58
27
29
53
53
25
34
26
4.2
5.5
25
10
28
34
24
37
43
95
107
96
106
90
87
75
1.0
0.95
0.92
0.20
0.28
0.79
0.91
0.90
0.92
0.81
0.70
0.81
0.95
0.96
1.0
0.66
0.66
0.83
1.1
0.81
0.76
0.85
0.50
0.20
0.56
0.68
0.96
0.74
0.86
0.77
0.91
0.76
0.88
0.98
0.96
1.0
4.0
Qv
g/(lh)
Ref
1.0
0.83
122
174
2.3
3.9
0.56
0.9
0.72
1.4
1.6
3.6
146
18
49
18
28
178
5.8
9.4
12
5.5
4.4
2.7
123
124
35
174
10
10
2.3
2.4
2.2
2.0
2.8
99
67
100
88
156
2.3
1.5
2.8
3.7
0.2
0.5
2.6
115
179
180
87
0.72
0.43
2.0
2.1
1.2
2.1
2.3
1.7
2.4
3.0
3.3
1.5
3.3
2.1
108
143
181
18
18
106
20

Table 4
95
(Continued)
Organism
Nutrients
hydrwheatflourprotease
hydrwheatflour
vitamins
amino acids
peptides
LA
g/l
YLA/tot
g/g
43
46
53
44
17
Qv
g/(lh)
Ref
1.5
2.4
2.8
2.2
0.23
106
Abbreviations as in Table 2 and: dep dependent; YE yeast extract; pep peptone; CSL corn steep liquor; trp tryptone; SSF simultaneous
saccharification and fermentation; cont continuous; Substr substrate; UF ultrafiltrated.
Table 5
Influence of fermentation mode on lactic acid production
Organism
Fermentation
mode
Substrate
LA
g/l
YLA/tot
g/g
Ent. faecium
batch
semicont
batch, imm
cont, imm
batch
fed-b
batch
cont
batch
cont
batch, imm
cont, imm
batch
cont
batch, imm
cont, imm
batch
cont
batch
cont
batch
cont
batch
cont, imm
batch, dial
cont, dial
batch
cont, extract
batch
cont, recirc,
extract
batch
cont
batch
cont
batch, imm
cont, imm
batch, extract
rep b, extract
batch
fed-b
alfalfa
alfalfa
corn starch
corn starch
whey
whey
glucose
glucose
hydr maize barley
hydr maize barley
glucose
glucose
whey
whey
whey
whey
whey
whey
lactose
lactose
whey
whey
whey
whey
whey
whey
sucrose
sucrose
glucose
glucose
27
30
50
40
22
45
83
55
85
71
90
75
44
13
50
9.5
115
117
45
39
49
48
0.91
0.99
0.83
0.67
0.44
0.45
0.83
0.55
0.87
0.73
0.9
0.75
0.95
0.28
1.0
0.19
0.86
0.87
0.90
0.78
1.1
1.2
Lb. casei L100

Lb. casei SU No 22
Lb. delbrueckii MIX several strains
Lb. delbrueckii NCIM-2365
Lb. delbrueckii sp. bulgaricus
Lb. delbrueckii sp. bulgaricus Ch H 2217
Lb. delbrueckii sp. bulgaricus NRRL B-548
Lb. helveticus ATCC 15009
Lb. helveticus L89
Lb. helveticus NCDO 1844

Lc. lactis 65.1
Lc. lactis IFO 12007
Str. thermophilus
glucose
glucose
glucose
glucose
potato starch
potato starch
glucose
glucose
lactose
lactose
47
125
77
80
80
47
38
10
39
28
25
10
0.29
0.50
40
39
1.2
3.1
0.73
0.74
0.89
0.48
0.76
0.20
0.75
0.56
0.50
0.20
0.3
0.4
Qv
g/(lh)
Ref
130
94
129
1.5
5.3
127
95
131
182
0.65
12
138
183
11
2.2
1.3
2.7
3.1
29
134
116
184
43
1.7
8.0
5.1
4.2
185
141
186
187
0.72
0.43
108
126
7.1
1.4
125
Abbreviations as in Table 2 and: dial electrodialysis; extract extraction, adsorption; imm immobilised cells; fed-b fed-batch; rep b repeated
batch; cont continuous culture; semicont semicontinuous culture.
96
Table 6
Influence of cell recirculation on lactic acid production
Organism
Fermentation
mode
Substrate
LA
g/l
YLA/tot
g/g
Lb. casei
cont,
cont,
cont
cont,
cont
cont,
cont
cont,
whey
whey
glucose
glucose
glucose
glucose
glucose
glucose
22
28
26
32
17
47
27
45
0.44
0.57
0.65
0.80
0.68
0.48
0.54
0.90

imm
imm, recirc
recirc
recirc
recirc
Qv
g/(lh)
Ref
7.3
9.4
72
142
5.1
4.2
3.4
10
141
143
Abbreviations as in Table 5 and: recirc recirculation of cells.
3.3. Nitrogen source

The medium composition has been investigated from
many aspects, including the addition of various concentrations of nutrients in the form of e.g. yeast extract, peptone
or corn steep liquor [28,69]. The addition of nutrients and
higher nutrient concentrations generally had a positive effect on the LA production (Table 4). MRS medium, which
contains yeast extract, peptone and meat extract, was superior to yeast extract, which in turn was better than malt
extract. This reflects the complex nutrient demands of LAB,
being fastidious because of limited biosynthesis capacity
[121]. Yeast extract alone at high concentration gave higher
LA production than yeast extract and peptone in low
amounts [122], but the opposite resulted when the concentration of yeast extract was kept constant and peptone was
added [123]. Whey treatment also affects the outcome of
fermentation (Table 4) [124]. WWF did not supply enough
nutrients for Lc. lactis [20]. This was overcome by adding
hydrolyzing enzymes, - and gluco-amylase and a protease,
releasing nutrients in forms of amino acids from the flour
[106].
3.4. Fermentation mode
LA is most commonly produced in the batch mode [111],
but numerous examples of continuous culture exist [90], as
well as some fed-batch [125] and semicontinuous/repeated
batch fermentations [126] (Table 5). When comparing batch
and continuous fermentation modes, the former gave higher
LA concentrations and yields in most of the studies (Table
5) [94]. This is mainly due to that all substrate is used in the
batch mode, whereas a residual concentration remains in the
continuous one. On the other hand, the continuous mode
generally resulted in higher productivities (Table 5) [127].
The major reason is probably that the continuous cultures
were run at a high dilution rate, where the advantage over
the batch mode is most pronounced [128]. Varying the dilution
rate (D) in continuous culture affects both the substrate and
nutrient concentrations. However, the effects on the yields and
productivities were inconclusive [25]. Fed-batch [129], semi-
continuous [130] and repeated batch mode [126] gave higher

yields than the batch mode (Table 5).
3.5. Immobilization and recirculation of cells
LAB cells can be recirculated or immobilized/supported
by solids in different ways to increase cell density (Table 6).
Immobilization of cells has not been very successful in
terms of increasing the LA yield and productivity [42,64,
67,73,131140]. In about half of the studies better results
were obtained using free cells. On the other hand, recirculation of cells gave higher LA concentrations and higher or
equal yields (Table 6) [72,141143].
3.6. pH
The fermentation pH is either set at the beginning and
then left to decrease due to acid production, or it is controlled by base titration, or by extraction, adsorption, or
electrodialysis of LA. The effect of pH has been studied by
fermenting at various pH values (Table 7). In all cases,
titration to a constant pH resulted in higher or equal LA
concentration, yield and productivity in comparison with
no pH control (Table 7) [130]. Removing LA by electrodialysis and extraction, including aqueous two-phase
systems, were successful in some of the studies [42],
whereas in others titration gave the same or better results
[144]. The optimal pH for LA production varies between
5.0 and 7.0. A pH below 5.7 was only optimal for Lb.
strains, which are known to tolerate lower pH than lactococci [145].
3.7. Temperature
The effect of temperature on the production of LA has
only been studied in a few reports (Table 8). The temperature giving the highest productivity was in some cases lower
than the temperature resulting in highest LA concentration
and yield [105,146], whereas in others the same temperature
gave the best results in all categories [20,146]. For Lb.
amylophilus, which is known to grow at 15C but not at
97
Table 7
Influence of initial pH and pH control on lactic acid production
Organism
Substrate
pH
pH control
Ent. faecium
alfalfa
alfalfa
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glocose
glucose
glucose
glucose
glucose
glucose
cellulose
cellulose
cellulose
hydr
maize barley
hydr
maize barley
hydr
maize barley
sorghum
sorghum
sorghum
whey
whey
whey
whey
whey
whey
lactose
lactose
lactose
cellulose
cellulose
cellulose
whey
whey
sorghum
sorghum
sorghum
sorghum
sorghum
sucrose
sucrose
glucose
glucose
glucose
glucose
glucose
glucose
gtlucose
glucose
cellulose
cellulose
6.2
5.8
5.4
6.0
6.5
7.1
7.8
4.2
6.2
6.2
init
NH4OH
NaOH titr
NaOH titr
NaOH titr
NaOH titr
NaOH titr
NaOH
NaOH
init, extract
Lb. delbrueckii IAM 1928
Lb. delbrueckii MIX several strains

Lb. delbrueckii sp. bulgaricus CNRZ 369
Lb. helveticus NCDO 1844



LA
g/l
8.3
27
YLA/tot
g/g
Qv
g/(lh)
0.27
0.90
5.5
5.0
5.5
5.5
6.0
6.5
7.0
4.2
5.0
5.9
5.0
CaCO3
titr, dial
NaOH titr
NaOH titr
titr, dial
NaOH titr
NaOH titr
NaOH titr
CaCO3
CaCO3
CaCO3
Na2CO3
10
36
27
5.6
55
53
81
92
88
81
54
49
15
26
18
59
0.12
0.42
0.32
0.06
0.55
0.53
0.79
0.89
0.81
0.78
0.52
0.47
0.30
0.52
0.36
0.61
5.5
Na2CO3
87
0.90
5.8
Na2CO3
85
0.87
5.5
6.0
6.5
5.4
6.0
3.5
4.5
5.5
6.5
4.5
5.0
5.6
4.2
5.0
5.8
5.6
5.6
5.5
6.0
6.5
7.0
7.5
6.0
6.0
5.0
5.5
6.0
6.5
4.2
4.2
6.0
6.0
4.3
4.3
NH3
NH3
NH3
NH4OH
NH4OH
init
init
init
init
NH4OH titr
NH4OH titr
NH4OH titr
NH4OH
NH4OH
NH4OH
NaOH titr
NaOH titr, dial
NH3
NH3
NH3
NH3
NH3
NaOH titr
NaOH titr, extract
NaOH
NaOH
NaOH
NaOH
yes
yes, extract
NH4OH
extract
NH4OH
NH4OH, extract
130
0.70
1.6
1.2
1.0
0.90
1.0
6.5
2.9
2.3
5.3
5.3
2.4
1.9
4.3
4.5
1.5
1.4
77
80
65
78
79
78
25
19
71
771
45
28
0.45
0.64
0.48
0.48
0.60
0.67
0.50
0.90
0.90
0.27
0.58
0.33
0.78
1.2
0.73
0.74
0.65
0.78
0.79
0.78
0.91
0.79
104
144
127
164
114
95
3.5
4.5
2.3
35
50
25
25
31
35
25
45
45
27
52
33
31
47
Ref
100
36
159
0.68
3.1
11
0.23
0.43
134
1.3
1.1
2.1
2.0
2.8
5.1
1.9
1.7
8.0
2.3
3.9
4.9
4.9
2.5
0.91
1.3
5.4
0.31
0.96
43
117
100
185
153
188
42
112
98
Table 7
(continued)
Organism
Substrate
pH
pH control
glucose
glucose
glucose
glucose
glucose
soy molasses
soy molasses
soy molasses
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
maltose
maltose
maltose
hydr wheat
flour
hydr wheat
flour
hydr wheat
flour
hydr wheat
flour
hydr wheat
flour
lactose citrate
lactose citrate
lactose citrate
lactose citrate
5.5
6.3
7.5
6.2
6.2
5.6
6.0
6.4
6.5
6.5
6.5
6.5
6.5
6.0
6.0
6.0
6.0
6.0
6.0
6.0
6.0
5.0
5.8
6.5
5.0
5.8
6.5
6.0
NH4
NH4
NH4
init
CaCO3
NaOH titr
NaOH titr
NaOH titr
init
init, extract
init
NaOH
init, extract
NaOH
NaOH, deal
yes
yes, dial
init, dial
NaOH, dial
NaOH
NaOH, dial
NaOH titr
NaOH titr
NaOH titr
NaOH titr
NaOH titr
NaOH titr
init
7.5
26
5.5
5.4
4.9
5.1
5.7
3.5
19
14
45
35
60
66
60
60
50
62
5.4
5.3
4.9
5.1
4.2
3.2
3.3
0.21
0.81
0.85
0.79
0.82
1.0
1.1
0.18
0.81
0.70
0.90
0.70
0.80
0.88
0.75
0.75
0.85
0.88
0.92
0.90
0.86
1.0
0.82
0.70
0.02
6.0
NaOH
96
0.76
3.0
4.0
NaOH titr
0.041
0.23
5.0
NaOH titr
20
0.11
0.42
6.0
NaOH titr
105
0.58
2.9
5.0
5.5
6.0
6.5
NaOH
NaOH
NaOH
NaOH
0.13
0.71
0.71
0.73
0.83
1.9
4.5
7.7

Lc. lactis 65.1

Lc. lactis 65.1

Lc. lactis sp. lactis biovar diacetylactis CNRZ 2125
titr
titr
titr
titr
LA
g/l
7.0
7.0
37
37
38
YLA/tot
g/g
Qv
g(lh)
Ref
16
23
17
0.30
2.6
132
180
87
189
0.87
1.7
10
15
3.0
4.0
2.4
5.1
187
143
190
191
136
1.7
3.4
2.5
0.37
1.2
1.0
0.47
25
25
18
20
152
Abbreviations as in Table 5 and: init initial pH set, then uncontrolled; titr titration.
45C [147], the optimal temperatures were 25 and 35C for

maximum productivity and yield, respectively [105]. For
Lb. casei and Lb. paracasei the optimal temperature was
reported to be between 37 and 44C [99,101,146], which is
contradictory to the information that the strains grow at 15
but not at 45C [147]. In agreement with previous observations [147,148], Lc. lactis and Lb. rhamnosus exhibited the
highest yields and productivities at 33 to 35C [20] and 41
to 45C [146], respectively.
4. Other effects on the processes
4.1. By-product formation
In addition to LA, LAB produce by-products, including
acetic acid, formic acid, carbon dioxide, and ethanol, de-
pending on the metabolic pathway used. For efficient industrial production of LA, by-product formation should be
avoided, or kept to a minimum. The ratio of g LA per g
total product (LA/tot) was higher in batch culture than in
fed-batch [112], and also under anaerobic conditions
compared with aerobic conditions [149] (Table 9). When
NaCl [149] and substrate concentrations [150] increased,
LA/tot also increased (Table 9). However, there was no
change in LA/tot when nutrients were varied (Table 9)
[18,151].
Different carbon sources give varying amounts of byproducts, so that maltose fermentation in Lc. lactis resulted
in values of LA/tot of 0.67, compared to 0.93 for glucose
fermentation (Table 9) [25]. Also, lactose and galactose
result in lower LA/tot-values than glucose in some strains of
99
Table 8
Influence of temperature on lactic acid production
Organism
Temp
C
Substrate
LA
g/l
YLA/tot
g/g
Qv
g/(lh)
Ref
25
28
35
30
37
41
45
37
41
30
36
44
30
37
41
45
30
35
37
40
30
35
37
30
34
37
40
starch
starch
starch
glucose
glucose
glucose
glucose
hydr barley starch
hydr barley starch
sweet sorghum
sweet sorghum
sweet sorghum
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
maltose
maltose
maltose
glucose
glucose
glucose
glucose
26
29
30
80
80
82
42
140
117
0.52
0.58
0.60
0.89
0.89
0.91
0.47
0.98
0.82
0.54
0.44
0.33
3.2
5.6
5.6
1.2
105

Lb. paracasei No 8
67
70
68
75
4.9
5.2
5.2
1.2
3.2
3.7
4.0
60
65
60
50
0.74
0.78
0.76
0.83
0.86
0.88
0.88
0.20
0.70
0.73
0.80
1.3
1.5
1.5
1.2
146
101
1.5
1.9
2.2
3.3
3.3
3.5
3.3
2.5
2.9
1.8
1.0
1.2
1.1
2.2
2.8
2.3
1.5
99
146
25
25
20
Abbreviations as in Table 2.
Lc. lactis [23,24]. Pentose fermentation results in the production of acetate or ethanol and LA in equimolar amounts,
and values of LA/tot of 0.57 0.79 have been reported (Table 9) [150]. Recirculation [53] of cells gave lower or
similar LA/tot-values as free cells, whereas the effects of pH
[151153] and temperature [20,105] were inconclusive (Table 9).
4.2. LA isomery
Most LAB produce only one isomeric form of LA, but
sometimes there is a slight production of the other isomer.
Lb. helveticus and Lb. plantarum produce a racemic mixture, the composition of which varies. The lactate dehydrogenase (LDH) is stereospecific, giving either D- or
L-LA [14]. Which isomeric form(s) of the enzyme present
in the LAB mainly determines the isomery of the LA
produced. For some applications, such as PLA synthesis,
an optically pure product or a racemic mixture of constant composition is desirable. For the L-LA-producing
Lb. amylophilus, Lb. delbrueckii and Lb. rhamnosus, no
D-isomer was produced when the pH was varied [36,153]
or when the amount of nutrients was changed [18,36]
(Table 10). On the other hand, only D-LA was formed by
Lb. delbrueckii spp. bulgaricus in batch and continuous
culture [34], from glucose and lactose [34], and when the
amount of nutrients was changed [36] (Table 10). The

composition of the racemate formed by Lb. plantarum
changed with aeration and amount of NaCl [149]. Comparing batch with continuous culture, the amount of the
predominant isomer was higher in the former [51]. The
amount of the predominant isomer also increased with
increasing pH [18,20] and amount of substrate [20], but
decreased with increasing temperature [20] and when the
pH was uncontrolled [18].
4.3. Cell density
The highest cell densities (48 103 g/l) were achieved by
recirculation [53,79,113,142,154], but also fermentations
without recirculation resulted in 60 and 77 g/l cells [27,155,
156]. Fermentation by LAB is normally accompanied by
increased cell mass that constitutes an undesired by-product
if the aim of the process is LA production. On the other
hand, the bacteria are the primary product in the production
of probiotics. At pH below 6.5 and temperatures above
30C, lower cell mass and cell yield were determined for Lc.
lactis fermenting glucose or maltose [25]. Similar results
were obtained for Lb. delbrueckii [157]. Maltose [25], lactose [158], and mannose [158] gave more cells than glucose,
whereas fructose [158], cellulose [159], and xylose [159
100
Table 9
Effect of process parameters on by-product formation
Parameter Organism
studied
Fermentation
mode
Substrate
Substr g/l/
Nutrients
carbon,
nutr
Lb. IMET 11466
batch
glucose
carbon
Lb. rhamnosus ATCC

10863
batch
fed-b, recirc
glucose
alpha-cellulose
hydr rye
1/20 MRS
MRS
fed-b, recirc
switch grass
cellulose
glucose
carbon
conc
conc
conc
nutr
nutr, O2
nutr, pH
mode
recirc
pH
pH
pH
pH
Lc. lactis sp. lactis ATCC batch

19435
batch
Lb. amylophilus ATCC
batch
49845
batch
batch
Lc. lactis IO-l JCM 7638 batch
batch
batch
batch
Lb. helveticus
cont
cont
Lb. delbrueckii sp.
batch
bulgaricus
ATCC 11842
batch
Lb. plantarum H4
batch, aer
batch, aer
batch, aer
batch, ana
batch, ana
batch, ana
Lb. plantarum MOP 3
batch
batch
batch
batch
batch
Lb. rhamnosus ATCC
batch
10863
fed-b, recirc
Lb. rhamnosus ATCC
cont
10863
cont, recirc 59%
cont, recirc 79%
cont, recirc 78%
Lb. rhamnosus ATCC
batch
10863
batch
batch
batch
batch
ATCC 19435
batch
batch
batch
ATCC 19435
batch
batch
batch
19435
batch
batch
maltose
starch
pH/
LA
TempC g/l
HAc
g/l
trace
35
93
45
0
1.0
1.0
0.98
28
0.50
0.98
0.93
25
3.2 0.49
29 0
0.36
0
0.72
0
0.67
1.0
105
0
0
0
0
1.0
1.0
0.57
0.72
0.74
0.70
70
100
29
49
130
160
37
127
hydr wheat flour
45
53
12
25
22
27
35
48
18
starch
glucosecitrate
glucosecitrate
glucosecitrate
glucosecitrate
glucosecitrate
glucosecitrate
glufrumalate
glufrumalate
glufrumalate
glufrumalate
glufrumalate
alpha-cellulose
hydr wheat flour YE
26
3.2
4.2
4.2
10.0
4.5
5.5
9.1
8.6
6.6
7.3
13
13
3.1
1.7
0.79
1.3
0.58
0.56
0
0
0
0
0
0
45
32
1.0
0.44
5.0
32
31
32
65
0.66
0.90
0.49
0.35
glucose
glucose
glucose
glucose
5.5
6.0
6.5
5.0
78
79
78
5.4
0.33
0.35
0.39
0.028 0.20
glucose
glucose
maltose
5.8
6.5
5.0
maltose
maltose
glucose
5.8
6.5
4.0
4.2 0.33
3.2 0.49
18 0.90
0.24
0.36
15
glucose
glucose
5.0
6.0
68
56
21
2.4
alpha-cellulose
glucose
glucose
glucose
glucose
glucose
NaCl 6%
NaCl 8%
HAc 49 mM
4.1
HAc 20 mM NaCl 3% 4.5
HAc 20 mM NaCl 6% 4.5
NaCl 6%
5.5
HAc 0.7 mM
6.0
40
40
40
40
112
0.065 0.20
starch
starch
xylose
xylose
xylose
xylose
lactose
lactose
starch
NaCl 6%
NaCl 8%
LA/tot Ref
g/g
92
4.9 0.10
50
EtOH HFo
g/l
g/l
0
0
9.1
9.7
7.9
7.3
1.0
1.5
10
present
present
0.64
18
15
0
0
0
0
0
150
56
0.63
0.51
0.71
0.84
0.88
0.89
0.91
1.0
1.0
1.0
1.0
1.0
1.0
112
149
151
0.29
0.66
0.98
0.96
142
0.45
0.64
0.30
1.1
1.5
0.74
0.10
0.94
0.91
0.95
0.99
153
0.40
0.40
0.51
0
0.99
0.99
0.99
0.96
25
5.3 0.067 0
0.16
4.9 0.10 0.065 0.20
5.1 0.098 0.065 0.19
0.96
0.93
0.94
25
0.54
0.72
1.0
0.79
0.67
0.46
106
0.90
0.60
0.93
0.93
2.0
1.5

Table 9
(continued)
Parameter
studied
Organism
Fermentation
mode
Substrate
Lc. lactis sp. lactis ATCC

19435
batch
glucose
30
batch
batch
batch
batch
glucose
glucose
glucose
maltose
batch
batch
batch
101

19435

19435
Lb. amylophilus ATCC

49845
Substr g/l/
Nutrients
pH/
TempC
LA
g/l
HAc
g/l
EtOH
g/l
HFo
g/l
LA/tot
g/g
Ref
4.9
0.10
0.065
0.20
0.93
25
35
37
40
30
5.2
5.2
1.2
3.2
0.074
0.085
0.030
0.49
0.080
0.050
0
0.36
0.18
0.11
0
0.72
0.94
0.95
0.98
0.67
25
maltose
maltose
glucose
35
37
30
3.7
4.0
60
0.48
0.36
1.0
0.38
0.26
1.0
0.78
0.60
0.69
0.77
0.97
20
batch
batch
batch
batch
glucose
glucose
glucose
starch
34
37
40
25
65
60
50
26
1.5
6.0
7.5
0
1.5
4.0
6.0
0
0.96
0.86
0.79
1.0
105
batch
batch
starch
starch
28
35
29
30
0
0
0
0
0
0
1.0
1.0
Abbreviations as in Table 6 and: carbon carbon source; nutr nutrients; O2 aeration; mode fermentation mode; T temperature; ana anaerobic;
aer aerobic; fru fructose; HAc acetic acid; EtOH ethanol; HFo formic acid; La/tot g LA per g total products.
161] fermentation resulted in lower cell masses than glucose.

5. Process considerations
The best fermentation conditions are not always the most
favorable for the whole process from an economical point of
view, because the costs of substrate and downstream processing are proportionally high. The substrate is usually
a question of geographic availability. Wastes from, for
example, agriculture and forestry are preferable to expensive, pure sugars for the low-price product LA. In
fermentative production of LA, renewable resources that
do not contribute to the greenhouse effect can be utilized.
A disadvantage is the infections, as observed when WWF
was fermented by Lc. lactis [20]. In Northern Europe
wheat is a major crop, whereas corn is used in the USA
and tapioca in Africa. Fractions lacking suitable polymer
qualities to be used for other purposes can be given added
value. The mode of operation has to be chosen to release
nutrients available in the substrate, e.g. enzymatic hydrolysis of starchy material.
For lignocellulosic substrates, a strain fermenting pentoses as well as hexoses such as Lb. pentosus is required
to maximize the yield [111]. Starch can either be both
hydrolyzed and fermented by certain amylase-producing
Lb. strains, such as Lb. amylovorus [102], or, after being
hydrolyzed to glucose, fermented by a number of organisms [18]. A homofermentative strain maximizes the proportion of LA produced. In contrast to the synthetic
racemate, an optically pure product can be produced by
fermentation by choosing the proper organism and
growth conditions.
Batch fermentation was superior to continuous fermentation in all respects but the volumetric productivity [127,
138]. Repeated or semicontinuous batch modes increase the
yield further [44]. If the substrate is expensive the yield
should be maximized, as in batch or semicontinuous operation [128], whereas the volumetric productivity is maximized by continuous operation if investment costs are high.
A high productivity is achieved by recycling the cells,
resulting in a high cell mass without reducing the yield
[47]. When using starch or lignocellulose, which must be
hydrolyzed before fermentation, the two steps can be
performed separately or simultaneously (SSF). In SSF the
hydrolyzing enzymes are not inhibited due to the continuous removal of the produced glucose, and less enzyme is
required. The time was only marginally reduced in SSF
of WWF by Lc. lactis [106]. Only one vessel is needed,
and only one temperature and pH has to be adjusted. In
SSF recirculation of cells is hampered by solid substrate
residues fouling the equipment. SSF of WWF requires
nutrient supplementation that is not demanded in separate
fermentation [106].
pH control is traditionally performed with calcium hydroxide, but the regeneration of LA results in the production
of large amounts of solid calcium sulfate [3]. Better alternatives are ammonia or calcium carbonate, leading to production of the fertilizer ammonium sulfate [162] or gaseous
carbon dioxide, respectively. Continuous removal of the
acid with extraction or electrodialysis results in even higher
LA concentrations and yields. The extracting material must
be bio-compatible so as not to harm the organism, and one
way of achieving this is aqueous two-phase systems, which
provide good separation of LA and cells when combined
with a tertiary amine [163]. Solid resins have also been
102
Table 10
Effect of process parameters on the isomeric form of LA produced
Parameter
studied
Organism
carbon, conc
Lb. delbrueckii sp. bulgaricus CBS 743.84
Fermentation
mode
batch
batch
batch
carbon, nutr, T Lb. delbrueckii sp. bulgaricus CBS 743.84 batch
batch
carbon
Lb. delbrueckii sp. bulgaricus DSM 2129
batch
batch
conc
batch
batch
batch
conc
batch
batch
conc, recirc
cont, recirc
cont, recirc
conc
batch
batch
nutr
Lb. delbrueckii sp. delbrueckii ATCC 9649 batch
batch
nutr
Lb. delbrueckii sp. delbrueckii ATCC 9649 batch
batch
nutr
Lb. delbrueckii sp. bulgaricus ATCC 11842 batch
batch
nutr
batch
nutr, O2
Lb. plantarum H4
batch, aer
batch, aer
batch, aer
batch, ana
batch, ana
batch, ana
nutr
batch
batch
batch
batch
batch
batch
nutr
batch
batch
batch
nutr
batch
batch
nutr
batch
batch
nutr
batch
batch
mode
Lb. delbrueckii sp. bulgaricus DSM 2129
batch
cont
mode
batch
cont
recirc
cont, recirc
cont, recirc
recirc
cont, recirc
cont, recirc
pH
batch
pH
batch
batch
batch
batch
pH
batch
batch
78%
96%
51%
79%
78%
96%
Substrate
Substr g/l/
Nutrients
whey UF perm
centr whey
centr whey
lactose
lactose
glucose
lactose
starch
starch
starch
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
starch
starch
starch
starch
lactose
lactose
glucose citrate
glucose citrate
glucose citrate
glucose citrate
glucose citrate
glucose citrate
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
glucose
starch
starch
glucose
glucose
starch
starch
glucose
glucose lactose
glucose
glucose
glucose
glucose
glucose
glucose
lactose
lactose
glucose
glucose
glucose
glucose
starch
starch
43
93
78
whey milk YE
YE trp
pH/
% L-LA Ref
TempC
37
44
50
70
100
50
70
40
80
85
174
YE 1%
YE 3%
hydr wheat flour
hydr wheat flour YE
hydr wheat flour
hydr wheat flour YE
whey YE
whey soy flour
NaCl 6%
NaCl 8%
NaCl 6%
NaCl 8%
YE 0.25% trp 0.5%
YE 0.5% trp 1%
YE 0.75% trp 1.5%
YE 1% trp 2%
YE 1.5% trp 3%
YE 2% trp 4%
YE 0.2%
YE 1%
hydr wheat flour
hydr wheat flour YE
hydr wheat flour
unhydr wheat flour
hydr wheat flour
hydr wheat flour YE
5.4
6.0
5.0
5.5
6.0
6.5
6.0 init
6.0
1
7
10
1
3.0
0
0
93
93
93
95
95
96
97
96
99
0
0
94
95
91
95
100
100
48
44
43
45
36
33
95
95
95
95
95
95
98
98
97
94
100
99
98
100
100
0
0
90
86
96
95
96
97
100
100
98
98
98
97
97
100
33
33
34
105
156
142
20
49
18
18
36
149
156
180
18
20
18
34
51
142
142
36
153
18

Table 10
103
(continued)
Parameter
studied
Organism
Fermentation
mode
Substrate
pH
batch
batch
batch
batch
batch
batch
batch
batch
batch
batch
glucose
glucose
glucose
starch
starch
starch
glucose
glucose
glucose
glucose
Substr g/l/
Nutrients
pH/
TempC
% L-LA
6.0
5.0
3.0
25
28
35
30
34
37
40
99
99
97
93
93
93
99
90
96
82
Ref
20
105
20
Abbreviations as in Table 9 and: UF perm ultrafiltrated permeate; centr centrifuged; % L-LA % of LA in L-form.
used in combination with immobilized cells in fluidizedbed reactors, resulting in a LA concentration of up to 771
g/l [42]. Electrodialysis can be used in two ways: as a pH
controller producing the lactate anion [164], or in combination with a base, e.g. NaOH, to split the Na-lactate
into NaOH and LA [165]. The cells are removed by
filtration not to foul the membranes. The price of the
membranes is presently a considerable drawback. Both
aqueous two-phase systems and electrodialysis yield LA,
instead of lactate, which potentially could decrease the
purification costs.
Acknowledgments
This work was financially supported by the Swedish
National Board for Industrial and Technical Development
(NUTEK).
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Enzyme and Microbial Technology 26 (2000) 87107

Factors affecting the fermentative lactic acid production from

This paper is based on the Ph.D thesis of Karin Hofvendahl (1998)

Hemicellulose, in contrast to starch and cellulose, contains

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

subdivision of the Gram-positive bacteria; i.e. the GC

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

which in addition to give LA is also metabolized into

2. Effectiveness of various processes

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (0) 87107

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

3. Parameters affecting efficiency

yeast production [60]. In addition it could be used for LA

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

Substrate and treatment

Lb. amylophilus ATCC 49845

Lb. amylovorus ATCC 33620

Lb. amylovorus ATCC 33622

Lb. delbrueckii sp. delbrueckii

Lb. delbrueckii sp. delbrueckii ATCC 9649

Lb. pentosus NRRL B-227

Lb. pentosus NRRL B-473

Lb. plantarum NRRL B-531

Lb. plantarum NRRL B-787

Lb. plantarum NRRL B-788

man, xyl, gal

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

Substrate and treatment

Lb. plantarum NRRL B-813

Lb. plantarum USDA 422

Lb. rhamnosus ATCC 10863

Lb. rhamnosus ATCC 10863

Lc. lactis sp. lactis NRRL B-4449

arabinose, lactose, fructose, and hydrolyzed cellulose

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

Substrate and treatment

Lactobacterium delbrueckii sowjeskij

Lb. helveticus sp. milano

Lb. IMET 11466

Lb. rhamnosus ATCC 10863

Lb. salivarius sp. salivarius ATCC 11742

Lc. lactis sp. lactis AS211

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

Substrate and treatment

Lc. lactis sp. lactis ATCC 19435

Lb. casei L100

Lb. rhamnosus ATCC 10863

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

Lb. rhamnosus ATCC 10863

Abbreviations as in Table 5 and: recirc recirculation of cells.

3.3. Nitrogen source

continuous [130] and repeated batch mode [126] gave higher

K. Hofvendahl, B. HahnHagerdal / Enzyme and Microbial Technology 26 (2000) 87107

Lb. amylophilus ATCC 49845

Lb. delbrueckii IAM 1928

Lb. delbrueckii IFO 3534

Lb. delbrueckii IFO 3534

Lb. delbrueckii IFO 3534

Lb. delbrueckii MIX several strains

Lb. delbrueckii sp. bulgaricus ATCC 11842

Lb. delbrueckii sp. bulgaricus ATCC 55163

Lb. delbrueckii sp. bulgaricus NRRL B-548

Lb. delbrueckii sp. bulgaricus NRRL B-548