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Fermentative Production of Lactic Acid From Renewable Materials: Recent Achievements, Prospects, and Limits
Fermentative Production of Lactic Acid From Renewable Materials: Recent Achievements, Prospects, and Limits
REVIEW
The development and implementation of renewable materials for the production of versatile chemical resources have
gained considerable attention recently, as this offers an alternative to the environmental problems caused by the petroleum industry and the limited supply of fossil resources. Therefore, the concept of utilizing biomass or wastes from
agricultural and industrial residues to produce useful chemical products has been widely accepted. Lactic acid plays an
important role due to its versatile application in the food, medical, and cosmetics industries and as a potential raw
material for the manufacture of biodegradable plastics. Currently, the fermentative production of optically pure lactic
acid has increased because of the prospects of environmental friendliness and cost-effectiveness. In order to produce
lactic acid with high yield and optical purity, many studies focus on wild microorganisms and metabolically engineered
strains. This article reviews the most recent advances in the biotechnological production of lactic acid mainly by lactic
acid bacteria, and discusses the feasibility and potential of various processes.
2014, The Society for Biotechnology, Japan. All rights reserved.
[Key words: Lactic acid; Renewable materials; Microbial fermentation; Lactic acid bacteria; Metabolic pathways]
1389-1723/$ e see front matter 2014, The Society for Biotechnology, Japan. All rights reserved.
http://dx.doi.org/10.1016/j.jbiosc.2014.06.003
11
Bacteria
Bacteria
Bacteria
Fungi
e, not determined. ldh, lactate dehydrogenase gene; pdc, pyruvate decarboxylase gene; cupdc1, pyruvate decarboxylase gene; p, pyruvate formate lyase gene; msg, methylglyoxal synthase gene; cydAB, cyoABCD, cbdAB, cytochrome
oxidases genes; glpK-glpD, glycerol kinase and glycerol-3-phosphate dehydrogenase gene; dld, D-lactate dehydrogenase gene; xylA, xylRAB, xylose isomerase genes; xylB, xylulokinase gene; bgl, b-glucosidase gene; als, acetolactate
synthase gene; xpk, phosphoketolase gene; pta, phosphate acetyltransferase gene; adhE, alcohol/acetaldehyde dehydrogenase gene; frdA, fumarate reductase gene; tkt, transketolase gene; LDH, lactate dehydrogenase.
31
e
77.5
Glucose
Increase LDH activity
Bacteria
Bacteria
E. coli ATCC 11303 TG114
E. coli ATCC 700926 LA02Ddld
expression
L-ldhA
29
32
42
51
50.1
38.6
Glucose
Xylose
Arabinose
Improved yield of D-lactic acid
L-Lactic acid fermentation from xylose
D-Lactic acid fermentation from arabinose
0.97e0.99
0.79
0.82
40
41
Yeast
Yeast
Yeast
Yeast
Yeast
Bacteria
Bacteria
Bacteria
Bacteria
Candida utilis NBRC0988 cupdc1 D4-ldh2
Kluyveromyces lactis CBS2359 pEPL2
Pichia stipitis CBS6054 VTT-C-04590
Saccharomyces cerevisiae CEN.PK182 RWB850-2
S. cerevisiae OC2 YILM-2B
Bacillus coagulans P4e102B QZ19
Corynebacterium glutamicum X5C1
C. glutamicum R DldhA/pCRB204
Escherichia coli ATCC 700926 ECOM3
118
32
Glucose
Glycerol
msgA deletion
pta, adhE, frdA, dld deletion and glpK-glpD
expression
pB mutation
xylRAB expression
D-ldhL1and xpk1 deletion, tkt expression
0.98
0.85
0.95
0.60
0.58/0.44
0.83 0.02
0.61
e
e
0.87
0.8
103.3
109
58/41
122
61.5
90
z40.5
120
e
Glucose
Glucose
Xylose/Glucose
Glucose
Glucose
Glucose
Cellobiose/Glucose/Xylose
Glucose
Glucose
Genus
L-ldhL
Yield
Y (g g1)
Lactic acid
g (g L1)
Substrate
Outcome
Approach
J. BIOSCI. BIOENG.,
TABLE 1. Metabolic engineering approaches for lactic acid production by several microorganisms.
33
34
35
28
36
37
38
29
39
WANG ET AL.
Reference
12
Cellobiose
Cellooligosaccharides
Arabinose
Galactose
(1)
13
Xylose
(3)
(2)
-glucosidase
Glucose1-P
Ribulose
ATP
ATP
(20)
ADP
ADP
NAD(P)H
ADP
NAD(P)
6-Phosphogluconate
ADP
(8)
NAD(P)+
(5)
NAD(P)H
ATP
(21)
Ribulose 5-P
(19)
(17)
ATP
Xylulose
UDP-glucose
(18)
ADP
Glucose
Galactose1-P
UDP-galactose
ATP
(7)
CO2
Xylulose 5-P
Glucose 6-P
(6)
Mannose
Ribose 5-P
(22)
(9)
(16)
(4)
Xylulose 5-P
Xylulose 5-P
PEP
Pyr
GAP
Mannose 6-P
(15)
Sedoheptulose 7-P
(10)
Fructose 6-P
ATP
(9)
(11)
Erythrose 4-P
ADP
Acetyl-P
GAP
NAD+
NADH
ATP
ADP
CoA
(24)
ATP
Pyruvate
(12)
Dihydroxyacetone
phophate
(13)
NAD
ADP
(14)
NAD
Acetic acid
Acetyl-CoA
NADH
GAP
(23)
Pi
H20
Fructose1,6-P
ADP
NADH
(25)
NAD
ATP
NADH
H20
Pyruvate
Lactic acid
Acetaldehyde
NADH
NADH
NAD+
NAD
(14)
(26)
Lactic acid
PP/Glycolytic Pathway
(Homo-lactic acid metabolism)
Ethanol
PK Pathway
(Hetero-lactic acid metabolism)
FIG. 1. Pathways for lactic acid production from renewable materials-derived sugars (glucose, xylose, cellobiose, arabinose, mannose, and galactose) by lactic acid bacteria (4,7,16,19).
GAP, glyceraldehyde 3-phosphate; PEP, phosphoenolpyruvate; Pyr, pyruvate. Enzymes: 1, galactokinase; 2, arabinose isomerase; 3, xylose isomerase; 4, mannose phosphotransferase system; 5, hexokinase; 6, glucose-6-phosphate dehydrogenase; 7, 6-phosphogluconate dehydrogenase; 8, ribulose-5-phosphate 3-epimerase; 9, transketolase;
10, transaldolase; 11, 6-phosphofructokinase; 12, fructose-bisphosphate aldolase; 13, triosephosphate isomerase; 14, lactate dehydrogenase; 15, phosphomannose isomerase;
16, phosphoglucose isomerase; 17, phosphoglucomutase; 18, galactose-1-phosphate uridyl transferase; 19, glucosyltransferase; 20, ribulokinase; 21, xylulokinase; 22, phosphoketolase; 23, acetate kinase; 24, phosphotransacetylase; 25, aldehyde dehydrogenase; 26, alcohol dehydrogenase.
14
WANG ET AL.
(55), wheat bran extract (56), and silkworm larvae (57). By using
corn steep liquor as the cheap nitrogen sources, the lactic acid
productivity increased with decreasing the C/N ratio (mol/mol)
from 37:1 to 19:1, while the further decrease to 12:1 had no positive impact on lactic acid production by Lactobacillus sp. MKTLC878 (58).
In addition to carbon sources and nitrogen sources, vitamins and
minerals have signicant effects on lactic acid fermentation by LAB.
For example, pyridoxine stimulated the growth of some LAB, while
neither 2,4,5-trimethyl-3-hydroxypyridine nor 2,4-dimethyl-3hydroxy-5-hydroxymethyl pyridine showed the same effect (59).
Chauhan et al. (60) screened media components for high lactic acid
production with Lactobacillus sp. KCP01. They found that
MgSO4$7H2O, KH2PO4, sodium citrate, NaCl, and sodium succinate
had less of an effect on lactic acid production, and that sodium
sulphate, sodium acetate, K2HPO4, MnSO4$4H2O and FeSO4$7H2O
were found to be signicant.
Effects of fermentation condition on lactic acid
fermentation by LAB In the eld of fermentation technology,
fermentation conditions such as pH, temperature, and inoculum
size were considered important factors for cell growth, lactic acid
concentration, lactic acid productivity, and lactic acid yield.
The initial or controlled optimal pH value for lactic acid production varies between 5.0 and 7.0, depending on the used LAB.
Without pH control, the pH of the fermentation broth decreases
with increasing lactic acid production, resulting in the inhibition of
cell growth and its production. Idris and Suzana (61) reported the
effect of initial pH from 4.5 to 8.5 on lactic acid production during
batch fermentation by Lactobacillus delbrueckii ATCC 9646. An
initial pH of 6.5 caused the early induction of sugar consumption,
maximal rate of sugar consumption, and maximal lactic acid concentration. Guyot et al. (62) studied lactic acid production in batch
fermentation with pH control by Lactobacillus manihotivorans LMG
18010T at different pH values (4.5). Compared to non-pHcontrolled batch fermentation, pH-controlled batch fermentation
at 6.0 reduced the fermentation period by half, and increased the
lactic acid concentration to 12.6 g L1 and the OD600 value to 5.1
from 7.6 g L1 and 1.65, respectively.
Since LAB are mesophilic bacteria, LAB can grow and produce
lactic acid at a maximum temperature of approximately 50 C. In
addition, the growth rates of the LAB used differed based on the
fermentation temperature, and the optimum temperature for lactic
acid concentration did not correspond to lactic acid yield or lactic
acid productivity. For Lactobacillus amylophilus ATCC 49845,
Yumoto and Ikeda (63) reported optimal temperatures of 25 C and
35 C for maximum lactic acid productivity and yield, respectively.
Abdel-Rahman et al. (17) examined the effect of temperature on the
production of lactic acid from 20 g L1 cellobiose by E. mundtii QU
25. Both the maximum lactic acid concentration and the highest
yield were obtained at 30e43 C.
An inoculum size of 5e10% (v v1) is desirable to arrest heterolactic fermentation and to reduce the duration of the lag phase (64).
Because an inoculum size of more than 5% is costly and constrains
the operation, the inoculum size should be decreased. To investigate the inuence of inoculum size on lactic acid production,
different amounts (1e5% v v1) were inoculated into the fermentation medium. The maximum lactic acid concentration of
33.72 g L1 by Lactobacillus casei NBIMCC 1013 was observed with
an inoculum size of 2e4% (v v1), whereas a low lactic acid production was attributed to a low density of starter culture (1% v v1)
(65). Qi and Yao (66) studied the effect of inoculum size from 3% to
8% on lactic acid fermentation from rice straw with Lactobacillus
casei GIM 1.159. An inoculum size of 6% was found to be optimal for
the production of lactic acid. Lower inoculum sizes resulted in
insufcient biomass for lactic acid conversion, whereas higher
J. BIOSCI. BIOENG.,
inoculum sizes caused excessive depletion of the nutrients necessary for cell growth.
DIFFICULTIES AND PROBLEMS IN LACTIC ACID FERMENTATION
BY LAB
Although approximately 90% of worldwide lactic acid is produced by microbial fermentation, there are still many challenges for
more effective and economical production. One of the obstacles in
the process of co-culture using mixed sugars is carbon catabolite
repression when using renewable materials containing various
components (16,67). A few wild-type and metabolically engineered
strains have been shown to simultaneously consume different
sugars such as glucose/cellobiose (17), xylose/arabinose/glucose
(68), and cellobiose/glucose/xylose (38). Moreover, the strategy of
mixed cultures with different microorganisms has been studied for
improving product yield and sugar consumption (50,69).
Another obstacle is end-product inhibition in the lactic acid
production process by LAB. As fermentation progresses along with
substrate utilization, the concentration of lactic acid gradually increases, causing acidication of fermentation broth and leading to
slowing of the fermentation process, including cell growth, substrate utilization, and lactic acid production. Thus, to alleviate the
end-product inhibition effect caused by lactic acid during fermentation, lactic acid should be removed selectively in situ from the
fermentation broth by a separation method such as electrodialysis
(70), or lactic acid-resistant LAB should be selected (7).
By-products such as ethanol and acetic acid are usually formed in
hetero-fermentation using pentoses, which results in a theoretical
yield of only 0.60 g g1 of lactic acid from pentose (16,42,43). Cost of
separation and purication also increases with the lowered purity of
the lactic acid product. Some potential pentose-utilizing strains are
reported to overcome the problem, e.g., the wild-type of E. mundtii QU
25 can consume xylose homofermenatively with little or no acetate
production and with high yield (16). A metabolically engineered
Lactobacillus plantarum DldhL1-xpk1::tkt was also reportedly able to
convert xylose (46) and arabinose (42) to lactic acid in homofermentation.
The most serious obstacle for lactic acid production commercially is the availability and cost of feedstock for lactic acid
fermentation (4,7). Particularly, carbon sources are signicant,
should be consumed by the LAB, and are divided into 2 groups:
puried sugars such as glucose, xylose, sucrose, and sugar-containing materials. However, the use of puried sugars as feedstock
for lactic acid production is very expensive. From the perspective of
the economics of the feedstock in terms of its geographic availability, domestic renewable materials such as wastes and unutilized
resources from the agro-industry and forestry are preferred due to
their availability and low cost.
LACTIC ACID PRODUCTION FROM RENEWABLE MATERIALS
In most cases, glucose is the preferred carbon source for lactic
acid fermentation by LAB. However, as cheap and widely existing
renewable materials, starch (e.g., wheat starch, corn starch, sago
starch, potato starch) and lignocellulose (e.g., woody materials, crop
residues) are recognized to meet the requirements for the biotechnological production of lactic acid economically and efciently.
Starch from various plant products is a potentially renewable
material in terms of cost and availability. As a type of polysaccharide
of glucose, starch mainly exists in tubers such as potatoes and
cassava, and seeds of grains including wheat, corn, and rice. On the
other hand, lignocellulose, another carbohydrate source, is available
in large quantities, with widespread distribution and comparably
lower cost (6,71). The major constituents of lignocellulose are
15
TABLE 2. Recent investigations on lactic acid production from renewable materials by different fermentation modes and methods.
Microorganism
Substrate
0.50
0.93
e
0.76
0.93
0.83
0.38
1.105 0.06
1.56
3.04
1.7
1.2
68
15
39.1
83.8
141.5
162.5
180
35.4
90.0
48.7
44.2
166
28.0
37.11
24.0
46.4
32.5
57.61 1.37
118
0.70
0.96
0.936
0.897
0.903
0.35
0.90
0.95
0.92
0.87
0.28
0.46
0.76
0.46
0.88
0.46
0.94
0.81
1.40
4.7
1.69
2.14
0.75
2.3
1.01
5.7
4.2
0.78
1.03
0.51
0.64
5.4
1.60 0.12
1.71
88
89
13
90
72
91
53
92
97
129
27.0
10.9
0.89
0.95
0.90
0.36
1.74
3.1
6.7
0.17
93
94
18
Fed-batch
Batch
Continuous
Continuous (with high cell density)
Batch
Batch
74.7
16.6 0.8
10.4
11.7
93.0
62.8
Corncob molasses
Sago starch
E. faecalis RKY1
Lactobacillus bifermentans DSM
20003T
L. brevis S3F4
L. casei NCIMB 3254
L. casei G-02
L. casei LA-04-1
Wood hydrolyzate
Wheat bran hydrolyzate
Alfalfa bers
Cellobiose and cellotriose
Corncob residue
Molasses
Rice bran
Green microalga
Trimming vine shoots
Alfalfa bers
Apple pomace
Hydrolyzed acorn starch
Wheat starch
Batch
SSF
Fed-batch, SSF
Fed-batch (fed with CSL)
Fed-batch (fed with YE)
SSF
Batch
Batch
Continuous
Batch
SSF
SSF
Batch
SSF
Batch
Batch
SHF
SSF
Batch
Cell-recycle
Batch
Reference
Lactic acid
g (g L1)
Corncob
Cassava bagasse
Jerusalem artichoke tubers
Soybean meal hydrolyzate
Productivity
P (g L1 h1)
Fermentation mode
Yield
Y (g g1)
81
82
83
79
84
85
72
86
87
SSF, simultaneous saccharication and fermentation; SHF, separate hydrolysis and fermentation; CSL, corn steep liquor; YE, yeast extract; e, not determined.
cellulose (linear b-D-glucan), hemicellulose (hetero-polysaccharides including xylose, glucose, mannose, galactose, and
arabinose), and lignin (a polymer of three closely-related phenylpropane moieties). Basically, cellulose forms a skeleton, which is
surrounded by hemicellulose and lignin (6).
Pretreatment of raw renewable materials
Enzymatic hydrolysis of raw renewable materials easily exposes the fermentative
sugars consumed by the used strain, and the hydrolyzates can then be
used in SSF, or direct fermentation (72). However, enzymatic
hydrolysis of lignocellulosic biomass without pretreatment is usually
not very effective because of the high stability caused by the
association of cellulose and hemicellulose with lignin (73). Therefore,
pretreatment of renewable materials is performed to remove lignin,
separate cellulose and hemicellulose, increase the accessible surface
area, partially depolymerise cellulose, and increase the porosity of
the material to aid in the subsequent access of the hydrolytic
enzymes. Pretreatment methods include physical (mechanical),
physico-chemical (steam pretreatment, hydrothermolysis, wet
oxidation, ammonia ber expansion), chemical (dilute acid, alkaline,
ionic liquids), and biological methods (microorganisms, enzymes)
(5,73). As the result of pretreatment, the size of the material is
reduced and its physical structure is opened. On the other hand,
when starchy materials are used as the substrate for lactic acid
production, the bioconversion requires a pretreatment process
including the gelatinization and liquefaction of starch, which is
carried out at a temperature between 90 C and 130 C for 15 min (74).
Acid pretreatment is extensively applied in the hydrolysis of
starchy and lignocellulose materials (6). Pretreatment with dilute
acid at high temperature, or strong acid (H2SO4, SO2, H3PO4) at low
pH, usually results in the hydrolysis of hemicellulose to monomeric
sugars (e.g., xylose, arabinose) and minimizes the need for hemicellulases (75). However, the utilization of various chemicals in the
pretreatment procedures is a major drawback and affects the total
cost of the fermentation. Physical or physico-chemical methods of
pretreatment such as milling, steam explosion, and irradiation
reduce the particle size, thereby increasing the available surface area
16
WANG ET AL.
J. BIOSCI. BIOENG.,
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