Application of solid-state fermentation to food industry—A review
Susana Rodrı´guez Couto
a,b
, M
a
A
´ ngeles Sanroma´n
a,*
a
Department of Chemical Engineering, Isaac Newton Building, University of Vigo, Lagoas Marcosende, 36310 Vigo, Spain
b
Department of Chemical Engineering, Chemical Engineering School, Rovira i Virgili University, 43007 Tarragona, Spain
Received 9 November 2004; accepted 19 May 2005
Available online 20 July 2005
Abstract
Solid state fermentation (SSF) has become a very attractive alternative to submerged fermentation (SmF) for specific applications
due to the recent improvements in reactor designs. This paper reviews the application of SSF to the production of several metabo-
lites relevant for the food processing industry, centred on flavours, enzymes (a-amylase, fructosyl transferase, lipase, pectinase),
organic acids (lactic acid, citric acid) and xanthan gum. In addition, different types of biorreactor for SSF processes have been
described.
Ó 2005 Elsevier Ltd. All rights reserved.
Keywords: Bioreactors; Enzyme production; Food processing industry; Solid-state fermentation
1. Introduction
Microorganisms have long played a major role in the
production of food (dairy, fish and meat products) and
alcoholic beverages. In addition, several products of
microbial fermentation are also incorporated into food
as additives and supplements (antioxidants, flavours,
colourants, preservatives, sweeteners, . . .). There is great
interest in the development and use of natural food and
additives derived from microorganisms, since they are
more desirable than the synthetic ones produced by
chemical processes.
Solid-state fermentation (SSF) reproduces the natural
microbiological processes like composting and ensiling.
In industrial applications this natural process can be uti-
lised in a controlled way to produce a desired product.
SSF is defined as any fermentation process performed
on a non-soluble material that acts both as physical sup-
port and source of nutrients in absence of free flowing
liquid (
). The low moisture content means
that fermentation can only be carried out by a limited
number of microorganisms, mainly yeasts and fungi,
although some bacteria have also been used (
). Some examples of SSF pro-
cesses for each category of microorganisms are reported
in
.
SSF offers numerous advantages for the production
of bulk chemicals and enzymes (
Pandey, Selvakumar, Soccol, & Nigam, 1999a; Soccol,
Iloki, Marin, & Raimbault, 1994
). This process is known
from ancient times and different fungi have been culti-
vated in SSF for the production of food. Typical exam-
ples of it are the fermentation of rice by Aspergillus
oryzae to initiate the koji process and Penicillium roque-
fortii for cheese production. Also, in China, SSF has
been used extensively to produce brewed foods (such as
Chinese wine, soy sauce and vinegar) since ancient time
(
). Also, in Japan SSF is used commercially
to produce industrial enzymes (
Since 1986 in Brazil a series of research projects for the
value-addition of tropical agricultural products and
sub-products by SSF has been developed due to the high
0260-8774/$ - see front matter
Ó 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jfoodeng.2005.05.022
*
Corresponding author. Tel.: +34 986 812383; fax: +34 986 812380.
E-mail address:
(M
a
. A
´ . Sanroma´n).
www.elsevier.com/locate/jfoodeng
Journal of Food Engineering 76 (2006) 291–302
amounts of agricultural residues generated by this coun-
try (
). Thus, the produc-
tion of bulk chemicals and value-added fine products
such as ethanol, single-cell protein (SPC), mushrooms,
enzymes, organic acids, amino acids, biologically active
secondary metabolites, etc. (
) has been produced from these raw materials
by means of SSF technique.
In recent years, SSF has received more and more
interest from researchers, since several studies for en-
zymes (
), flavours (
), colourants (
) and other substances of interest to the food indus-
try have shown that SSF can give higher yields
(
) or better product characteristics
(
Acun˜a-Arguelles, Gutierrez-Rojas, Viniegra-Gonza´lez,
) than submerged fermentation
(SmF). In addition, costs are much lower due to the effi-
cient utilisation and value-addition of wastes (
).
Castilho, Alves, and Medronho (2000)
,
have performed a detail economic analysis of the pro-
duction of Penicillium restrictum lipase in both SmF
and SSF. They found that for a production scale of
100 m
3
lipase concentrate per year, total capital invest-
ment needed for SmF was 78% higher than that needed
for SSF. Also, SSF unitary product cost was 47% lower
than the selling price. These studies pointed out that the
great advantage of SSF processes is the extremely cheap
raw material used as main substrate. Therefore, SSF is
certainly a good way of utilising nutrient rich solid
wastes as a substrate. Both food and agricultural wastes
are produced in huge amounts and since they are rich in
carbohydrates and other nutrients, they can serve as a
substrate for the production of bulk chemicals and en-
zymes using SSF technique.
The nature of the solid substrate employed is the
most important factor affecting SSF processes and its
selection depends upon several factors mainly related
with cost and availability and, thus, may involve the
screening of several agro-industrial residues. In SSF pro-
cess the solid substrate not only supplies the nutrients to
the culture but also serves as an anchorage for the
microbial cells. Among the several factors, which are
important for microbial growth and activity in a partic-
ular substrate, particle size and moisture level/water
activity are the most critical (
Generally, smaller substrate particles provide a larger
surface area for microbial attack but if they are too
small may result in substrate agglomeration as well as
poor growth. In contrast, larger particles provide better
aeration but a limited surface for microbial attack.
Therefore, a compromised particle size must be selected
for each particular process (
Research on the selection of suitable substrates for
SSF has mainly been centred around agro-industrial resi-
dues due to their potential advantages for filamentous
fungi, which are capable of penetrating into the hardest
of these solid substrates, aided by the presence of turgor
pressure at the tip of the mycelium (
). In addition, the utilisation of these agro-
industrial wastes, on the one hand, provides alternative
substrates and, on the other, helps in solving pollution
problems, which otherwise may cause their disposal
(
).
SSF offers numerous advantages over SmF such as
simpler technique and lower cost (
). However,
there are few designs available in the literature for biore-
actors operating in solid-state conditions. This is princi-
Table 1
Main groups of microorganisms involved in SSF processes (extracted
from
)
Microflora
SSF process
Bacteria
Bacillus sp.
Composting, natto, amylase
Pseudomonas sp.
Composting
Serratia sp.
Composting
Streptococcus sp.
Composting
Lactobacillus sp.
Ensiling, food
Clostridium sp.
Ensiling, food
Yeast
Endomicopsis burtonii
Tape cassava, rice
Saccharomyces cerevisiae
Food, ethanol
Schwanniomyces castelli
Ethanol, amylase
Fungi
Altemaria sp.
Composting
Aspergillus sp.
Composting, industrial, food
Fusarium sp.
Composting, gibberellins
Monilia sp.
Composting
Mucor sp.
Composting, food, enzyme
Rhizopus sp.
Composting, food, enzymes,
organic acids
Phanerochaete chrysosporium
Composting, lignin degradation
Trichoderma sp.
Composting, biological control,
bioinsecticide
Beauveria sp., Metharizium sp.
Biological control, bioinsecticide
Amylomyces rouxii
Tape cassava, rice
Aspergillus oryzae
Koji, food, citric acid
Rhizopus oligosporus
Tempeh, soybean, amylase, lipase
Aspergillus niger
Feed, proteins, amylase,
citric acid
Pleurotus oestreatus, sajor-caju
Mushroom
Lentinus edodes
Shii-take mushroom
Penicilium notatum, roquefortii
Penicillin, cheese
292
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
pally due to several problems encountered in the control
of different parameters such as pH, temperature, aera-
tion and oxygen transfer and moisture. SSF lacks the
sophisticated control mechanisms that are usually asso-
ciated with SmF. Control of the environment within the
bioreactors is also difficult to achieve, particularly tem-
perature and moisture.
The aim of this paper is to review the potential appli-
cation of SSF for the production of several metabolites
of great interest to the food industry. In addition, differ-
ent types of biorreactor for SSF processes are described.
2. Some examples of applications of SSF to food industry
2.1. Flavours
Flavours comprise over a quarter of the world mar-
ket for food additives. Most of the flavouring com-
pounds are produced via chemical synthesis or by
extraction from natural materials. However, recent mar-
ket surveys have shown that consumers prefer foodstuff
that can be labelled as natural. Plants have been major
sources of essential oils and flavours but their use de-
pends on natural factors difficult to control such as
weather conditions and plant diseases. An alternative
route for flavour synthesis is based on microbial biosyn-
thesis or bioconversion (
). Several microorganisms,
including bacteria and fungi, are currently known for
their ability to synthesise different aroma compounds.
Attempts to use these microorganisms in SmF resulted
in low productivity of aroma compounds (
), which hampered their industrial applica-
tion. SSF could be of high potential for this purpose
(
). Thus,
reviewed the
prospects of microbial production of food flavours
and
the
recommended
SSF
processes
for
their
production.
Several researchers have studied the production of ar-
oma compounds by SSF from several microorganisms
such as Neurospora sp. (
Zygosaccharomyces
rouxii
), Aspergillus sp. (
Yoshida, Ishikawa, & Kobayashi, 1990
), using pre-
gelatinised rice, miso and cellulose fibres, respectively.
Bramorski, Soccol, Christen, and Revah (1998)
com-
pared fruity aroma production by Ceratocystis fimbriata
in solid-state cultures using several agro-industrial
wastes (cassava bagasse, apple pomace, amaranth and
soybean), determining that the media with cassava ba-
gasse, apple pomace or soybean produced a strong fru-
ity aroma.
Soares, Christen, Pandey, and Soccol (2000)
also reported the production of strong pineapple aroma
when SSF was carried out using coffee husk as a sub-
strate by this strain.
Bramorski, Christen, Ramirez, Soc-
and
described the production of volatile
compounds such as acetaldehyde and 3-methylbutanol
by the edible fungus Rhizopus oryzae during SSF on
tropical agro-industrial substrates.
Kluyveromyces marxianus produced aroma com-
pounds, such as monoterpene alcohols and isoamyl ace-
tate (responsible for fruity aromas), in SSF using
cassava bagasse or giant palm bran as a substrate
(
).
Esters are the source of the aromas and among them
pyrazines, which possess a nutty and roasty flavour, are
used as a food additive for flavouring (
).
son, Creuly, Gros, and Larroche (1997)
and
studied the production of
2,5-dimethylpyrazine
(2,5-DMP)
and
tetramethyl-
prazine (TTMP) using B. natto and B. subtilis, respec-
tively, on soybeans in SSF. They found that SSF
was
very
suitable
for
the
production
of
these
compounds.
2.2. Enzyme production
Recently,
dos Santos, Souza da Rosa, DalÕBoit,
evaluated whether SSF is
the best system for producing enzymes. They found that
SSF is appropriate for the production of enzymes and
other thermolabile products, especially when higher
yields can be obtained than in SmF.
2.2.1. a-Amylase
a
-Amylases (endo-1,4-a-
D
-glucan glucanohydrolase
EC 3.2.1.1) are extra-cellular endo enzymes that ran-
domly cleave the 1,4-a linkages between adjacent glu-
cose units in the linear amylose chain and ultimately
generates glucose, maltose and maltotriose units. Since
the 1950s, fungal amylases have been used to manufac-
ture sugar syrups containing specific mixtures of sugars
Table 2
Advantages and disadvantages of SSF over SmF
Advantages
Disadvantages
Higher productivity
Difficulties on scale-up
Better oxygen circulation
Low mix effectively
Low-cost media
Difficult control of process
parameters (pH, heat, moisture,
nutrient conditions, . . .)
Less effort in downstream
processing
Problems with heat build-up
Reduced energy and cost
requirements
Higher impurity product, increasing
recovery product costs
Simple technology
Scarce operational problems
It resembles the natural
habitat for several
microrganisms
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
293
that could not be produced by conventional acid hydro-
lysis of starch. Amylases are extensively employed in
processed-food industry such as baking, brewing, prepa-
ration of digestive aids, production of cakes, fruit juices,
starch syrups, etc.
The production of a-amylases has generally been car-
ried out using SmF; however, SSF systems appear as a
promising technology. Recently,
used spent brewing grains in SSF for the production
of a-amylase and determined that the supplement of fer-
mentation media with Tween-80 or calcium ions en-
hanced a-amylase activity.
Krishna and Chandrasekaran (1996)
used banana
fruit stalk as a substrate in SSF with Bacillus subtilis.
Different factors such as initial moisture content, parti-
cle size, thermal treatment time and temperature, pH,
incubation temperature, additional nutrients, inoculum
size and incubation period on the production of a-amy-
lase were characterised. Results obtained for the optimi-
sation of process parameters clearly shown their impact
on the gross yield of enzymes as well as their indepen-
dent nature in influencing the organismÕs ability to syn-
thesise the enzyme. It is known that particle size (specific
surface area) is a critical factor in SSF. Banana fruit
stalk particles of 400 lm favoured maximal a-amylase
production compared to larger particles. A similar
trend was reported for the production of glucoamylases
with wheat bran (
) and cellulases with coir
pith of small particle size (
Nowadays, gelatinisation is coupled with liquefac-
tion, which is possible by the action of thermostable
amylases, which have been reported in both SmF (
ford, Stamford, Coelho, & Araujo, 2001
) and SSF
(
determined that the productivity of
thermostable amylases from Bacillus sp. was affected
by the nature of the solid substrate (wheat bran, rice
bran, corn bran and combination of two brans), nature
of the moistening agent, level of moisture content,
incubation temperature, presence or absence of sur-
factant, carbon, nitrogen, mineral, amino acid and
vitamin supplements. Maximum enzyme production
was obtained on wheat bran supplemented with glycerol
(1.0%, w/w), soyabean meal (1.0%, w/w),
L
-proline
(0.1%, w/w), vitamin B-complex (0.01%) and moistened
with tap water containing 1% Tween-40.
Recently,
reported the
use of coconut oil cake (COC) as a substrate for the pro-
duction of a-amylase by A. oryzae under SSF condi-
tions. Raw COC supported the growth of the culture,
resulting in the production of 1372 U/gds a-amylase in
24 h. Supplementation with 0.5% starch and 1% peptone
to the substrate positively enhanced the enzyme synthe-
sis producing 3388 U/gds, proving COC a promising
substrate for a-amylase production.
2.2.2. Fructosyl transferase
Fructosyl transferase (EC 2.4.1.10) catalyses the
formation of fructo-oligosaccharides from sucrose.
Fructo-oligosaccharides are present in various com-
monly consumed foods like fruits, vegetables, cereals
and honey in trace amounts. The production of fructosyl
transferase derived from microorganisms has attracted
attention in recent years by SmF using Aspergillus spp,
Penicillium spp and Aureobasidium spp (
) and SSF using both Aspergil-
lus foetidus and A. oryzae (
1995; Sangeetha, Ramesh, & Prapulla, 2004
).
Recently,
have studied the
production of fructosyl transferase by A. oryzae employ-
ing a great variety of agricultural by-products as sub-
strates: Cereal brans (wheat bran, rice bran and oat
bran), corn products (corn cob, corn bran, corn germ,
corn meal, corn grits and whole corn powder), coffee-
and tea-processing by-products (coffee husk, coffee pulp,
spent coffee and spent tea), sugarcane bagasse and cas-
sava bagasse. They found that, among them, the best re-
sults were obtained when rice bran, wheat bran, corn
germ, spent coffee and tea were used supplemented with
yeast extract and complete synthetic media.
2.2.3. Lipase
Lipases (triacylglycerol acylhydrolases, EC 3.1.1.3)
are well known as efficient biocatalysts for the hydroly-
sis of water-insoluble fatty-acid esters, being triacylgly-
cerols of long chain fatty acids their natural substrates.
Lipases are nowadays widely used at industrial scale
with applications in food, detergent, cosmetic and phar-
maceutical industries (
Most studies on lipolytic enzymes production by bac-
teria, fungi and yeasts have been performed in sub-
merged cultures; however, there are few reports on
lipase synthesis in solid state cultures. In recent years,
increasing attention has been paid to the conversion of
processing industry wastes in lipase by solid state
cultures.
There are several reports dealing with extracellular li-
pase production by fungus such as Rizhopus sp., Asper-
gillus sp., Penicillium sp. on different solid substrates
(
Christen, Angeles, Corzo, Farres, & Revah, 1995; Cor-
) under submerged conditions. However, few
researchers have investigated the synthesis of lipase by
yeasts using SSF technique. Among them,
Jayaraman, and Lakshmanan (1993)
determined that
the C/N ratio of the medium is an important parameter
for lipase production by the yeast Candida rugosa.
Farres (1991), Ohnishi, Yoshida, and Sekiguchi (1994),
Christen et al. (1995)
and
, compared SmF and SSF
294
S.R. Couto, M
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.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
systems for lipase production. All they found that en-
zyme yields were higher and stable in the latter.
Several factors can affect extracellular lipase produc-
tion such as pH, temperature, aeration and medium
composition. Furthermore, the presence of triglycerides
or fatty acids has been reported to increase lipolytic en-
zyme secretion by a certain number of microorganisms
(
). Therefore, in SSF the type
of substrate could be used to enhance the production of
enzymes, as several food and agroindustrial wastes are
rich in fatty acids, triglycerides and/or sugars.
Dominguez, Costas, Longo, and Sanroman (2003)
have reported the great potential of food-agroindustrial
wastes (ground nut and barley bran) as support-sub-
strates for lipase production in solid state cultures of
the yeast Y. lipolytica, since they led to much higher
activities than those found using an inert support.
2.2.4. Pectinases
They constitute a heterogeneous group of enzymes
that catalyse the degradation of pectins, which are the
structural polysaccharides present in vegetable cells
and are responsible for maintaining the plant tissues
integrity (
Alkorta, Garbisu, Llama, & Serra, 1998
). Pec-
tinases are widely used in the food industry to clarify
fruit juices and wine, to improve oil extraction, to re-
move the peel from citrus fruit, to increase the firmness
of several fruits and to degum fibres (
1996; Chang, Siddiq, Sinha, & Cash, 1994
).
Commercial pectinase preparations are produced
from fungal microorganisms, mainly by Aspergillus
niger strains. The use of SSF for pectinase production
has been proposed using different solid agricultural
and agro-industrial residues as substrates such as wheat
bran (
Castilho, Alves, & Medronho, 1999; Singh, Platt-
), soy bran (
),
cranberry and strawberry pomace (
), coffee pulp and coffee husk (
Roussos, & Viniegra-Gonzalez, 1993
), husk (
Minjares, Roussos, Raimbault, & Viniegra-Gonza´lez,
1993
Boccas, Roussos, Gutierrez, Serrano, & Viniegra,
), cocoa (
Schwan, Cooper, & Wheals, 1997
), lemon
and orange peel (
Garzo´n & Hours, 1991; Ismail, 1996;
Maldonado, Navarro, & Callieri, 1986
), orange bagasse,
sugar cane bagasse and wheat bran (
), sugar cane bagasse (
) and apple pomace (
). Also,
produced pectinase from A. niger by SSF
using sugar beet pulp as a carbon source and wastewater
from monosodium glutamate production as nitrogen
and water source. This allowed not only reducing pro-
duction costs but also decreasing the pollution source.
It was found that SSF was more productive than
SmF and, in addition, the pectinases produced by SSF
showed more stable properties: they had a higher
stability to pH and temperature and they were less
affected by catabolic repression than pectinases pro-
duced by SmF (
2.3. Organic acids
Organic acids have been utilised for long time by the
food industry as food additives and preservatives for
prevent deterioration and extending the shelf life of
perishable food ingredients. Here, two common organic
acids widely used on the food industry have been
considered.
2.3.1. Lactic acid
Lactic acid fermentation has received extensive atten-
tion since long time (
). It
has wide applications in food, pharmaceutical, leather
and textile industries and as a chemical feed stock. It
has two enantiomers
L
(+) and
D
( ) of which
L
(+) is
used by human metabolism due to the presence of
L
lac-
tate dehydrogenase and is preferred for food. Nowa-
days, lactic acid is in great demand due to its use as
starting material to produce biodegradable polymers
used in medical, industrial and consumer products
(
Bohlmann & Yoshida, 2000; Gross & Kalra, 2002;
Lichtfield, 1996; Malhotra, Raina, & Sanjay, 2000
Soccol, Marin, Rimbault, and Labeault (1994)
stud-
ied the production of
L
(+)-lactic acid by Rhizopous ory-
zae in solid-state conditions operating with sugarcane
bagasse as a support. They obtained a slightly higher
productivity than in submerged cultivation. Also,
investigated the
L
(+)-lactic acid
production by Lactobacillus paracasei in solid-state con-
ditions using sweet sorghum as a support. More re-
cently,
Naveena, Altaf, Bhadrayya, Madhavendra, and
and
Naveena, Altaf, Bhadriah, and Reddy
have reported the production of
L
(+) lactic acid
by Lactobacillus amylophilus GV6 under SSF conditions
using wheat bran as both support and substrate.
2.4. Citric acid
Citric acid is one of the most commonly used organic
acids in food and pharmaceutical industries. The food
industry is the largest consumer of citric acid, using al-
most 70% of the total production, followed by about
12% for the pharmaceutical industry and 18% for other
applications (
Shah, Chattoo, Baroda, & Patiala, 1993
Its pleasant taste, high solubility and flavour-enhancing
properties have ensured its dominant position in the
market. Although citric acid can be obtained by chemi-
cal synthesis, the cost is much higher than using fermen-
tation. It is mainly produced by SmF, by the filamentous
fungus A. niger. Recently, in order to increase the
S.R. Couto, M
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A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
295
efficiency of citric acid production using A. niger, SSF
has been studied as a potential alternative to SmF.
The production of citric acid depends strongly on an
appropriate strain and on operational conditions. Oxy-
gen level is an important parameter for citric acid fer-
mentation. Several researchers (
Gaime-Perraud, & Roussos, 1998; Prado et al., 2004
have studied the influence of forced aeration on citric
acid production and the metabolic activity of A. niger
in SSF by respirometric analysis. They showed that
citric acid production was favoured by a limited biomass
production, which occurred with low aeration rates.
Both works showed the feasibility of using the strain
A. niger for citric acid production by SSF.
Different agro-industrial residues such as apple
pomace, coffee husk, wheat straw, pineapple waste,
mixed fruit, maosmi waste, cassava bagasse, banana,
sugar beet cosset and kiwi fruit peel have been investi-
gated for their potential to be used as substrates (
& Woodams, 1985; Khare, Krishana, & Gandhi, 1995
Kumar, Jain, Shanker, & Srivastava, 2003a, 2003b
Shojaosadati & Babaripour, 2002
). In addition, SSF
gave high citric acid yield without inhibition related to
presence of certain metal ions such as Fe
2+
, Mn
2+
,
Zn
2+
, etc. (
Gutierrez-Rozas, Cordova, Auria, Revah,
), although
reported that addition of these minerals
into the production media to a certain level enhanced
citric acid production by 1.4–1.9 fold with respect to
SmF. Therefore, SSF is a good way of using nutrient
rich solid waste as a substrate.
2.5. Xanthan gum
Xanthan gum is a hetero-polysaccharide produced
industrially by the bacterium Xanthomonas campestris,
fermenting commonly glucose or sucrose. It is the most
important microbial polysaccharide from the commer-
cial point of view, with a worldwide production of about
30,000 tons per year, corresponding to a market of $408
million (
Demain, 2000; Sutherland, 1998
). This water-
soluble microbial polysaccharide gives aqueous solu-
tions with several industrial applications in the food,
cosmetic, textile and pharmaceutical industries due to
their rheological properties. Because of these properties,
they have been used as emulsifiers, as stabilisers and as
texture enhancers in the food industry.
Recently, the feasibility of this exopolysaccharide
production using SSF has been reported by the group
of
Navarini, and Bertocchi (1999)
. X. campestris strains
were cultivated on a great variety of solid substrates or
by-products such as spent malt grains, apple pomace,
grape pomace and citrus peels, easily available and
low cost substrates, in order to evaluate their ability to
produce the exopolysaccharide xanthan. With most of
the substrates, xanthan yields were comparable to those
obtained from conventional submerged cultivation. In
addition, the products were analysed by NMR spectros-
copy, revealing a composition consistent with that of
commercial xanthan.
2.6. SSF bioreactors
The design of an efficient industrial-level reactor for
SSF is of significance because SSF is more environmen-
tally friendly than SmF. However, it shows considerable
drawbacks such as transfer resistance, steep gaseous
concentration and heat gradients that develop within
the medium bed, which may adversely affect solid-state
fermentor performances (
). Agitation and rotation in
SSF were often carried out to improve mass and heat
transfers, but the shearing force caused by agitation
and rotation has adverse effects on medium porosity
and disrupts fungal mycelia.
There are four types of reactors to perform SSF pro-
cesses and each in their own design tries to make condi-
tions more favourable for fermentation under solid state
conditions. The bioreactors commonly used, which can
be distinguished by the type of aeration or the mixed
system employed, include the following:
Tray: It consists of flat trays. The substrate is spread
onto each tray forming a thin layer, only a few centime-
tres deep. The reactor is kept in a chamber at constant
temperature through which humidified air is circulated
(
). The main disadvantage of this configuration
is that numerous trays and large volume are required,
making it an unattractive design for large-scale produc-
tion (
Pandey, Soccol, Rodriguez-Leon, & Nigam, 2001
).
Packed-bed: It is usually composed of a column of
glass or plastic with the solid substrate retained on a per-
forated base. Through the bed of substrate humidified
air is continuously forced (
). It may be fitted with a jacket for
circulation of water to control the temperature during
fermentation (
). This is the configuration usually
employed in commercial koji production. The main
drawbacks associated with this configuration are: diffi-
culties in obtaining the product, non-uniform growth,
poor heat removal and scale-up problems.
Horizontal drum: This design allows adequate aera-
tion and mixing of the substrate, whilst limiting the
damage to the inoculum or product. Mixing is per-
formed by rotating the entire vessel or by various agita-
tion devices such as paddles and baffles (
296
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
Mitchell, Johns, & Litster, 1999
). Its main disad-
vantage is that the drum is filled to only 30% capacity,
otherwise mixing is inefficient.
Fluidised bed: In order to avoid the adhesion and
aggregation of substrate particles, this design supplies
a continue agitation with forced air (
). Although
the mass heat transfer, aeration and mixing of the sub-
strate is increased, damage to inoculum and heat
build-up through sheer forces may affect the final
product yield.
The different disadvantages detected in the above-
mentioned bioreactor designs to perform SSF processes
have promoted the necessity of developing new bioreac-
tor configurations or modifying the already existing de-
signs. These bioreactor configurations should be able to
operate in continuous mode with high productivity for
prolonged periods of time without operational problems
as well as permit the scale-up of the process. Our re-
search group has been working in this field, resulting
in the design of a new bioreactor, called immersion bio-
reactor. This bioreactor consists of a jacketed cylindrical
glass vessel with a round bottom, inside which several
wire mesh baskets filled with support colonised by the
fungus are placed. They moved upwards and down-
wards by means of a pneumatic system, remaining more
time outside than inside the medium (
) (
). It is noteworthy
that this bioreactor configuration was also able to run
in continuous mode without operational problems,
attaining high ligninolytic enzyme activities (
Couto, Barreiro, Rivela, Longo, & Sanroman, 2002
Different studies were carried out for the production
of natural food and additives derived from micro-
organisms in different bioreactor configurations. For
Trays
Gas exit
Air inlet
Solid substrate
Fig. 1. Scheme of a tray bioreactor (passive aeration; static).
Fig. 2. Scheme of a packed-bed bioreactor (humidified air; static).
Air inlet
Gas exit
Sampling port
Culture medium level
Support susbstrate
Air diffuser
3 rpm
Fig. 3. Scheme of a horizontal drum bioreactor (humidified air;
mechanical agitation).
Fig. 4. Scheme of a fluidised-bed bioreactor (humidified air; pneu-
matic agitation).
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
297
example, the production of aroma compounds by K.
marxianus grown on cassava bagasse in solid state fer-
mentation using packed bed reactors, testing two differ-
ent aeration rates was studied by
Headspace analysis of the culture by gas chromatogra-
phy showed the production of 11 compounds. The pre-
dominant compounds were ethyl acetate, ethanol and
acetaldehyde. The fruity aroma was attributed to the
productions of esters.
Recently,
Navarrete-Bolan˜os, Jime´nez-Islas, Botello-
Alvarez, Rico-Martı´nez, and Paredes-Lo´pez (2004)
have
employed
a
modular
rotating
drum
bioreactor
(equipped with inlet air injection, variable speed pumps,
humidifier, and gas analyser) for xanthophylls extrac-
tion from marigold flowers. Marigold extracts have been
commercialised internationally and are used as additives
for poultry feed, as they provide bright colours in egg
yolks, skin, and fatty tissues. For this reason, they are
used as an additive in several food and pharmaceutical
industries. Based on experimental design strategies, opti-
mum operation values were determined for aeration,
moisture, agitation and marigold-to-inoculum ratio in
SSF of marigold flowers by mixed culture of three
microorganisms (Flavobacterium IIb, Acinetobacter anit-
ratus, and Rhizopus nigricans), leading to a xanthophylls
yield of 17.8-g/kg dry weight. This value represented a
65% increase in relation to the control.
Milagres, Santos, Piovan, and Roberto (2004)
have
shown that Thermoascus aurantiacus was able to pro-
duce a high level of thermostable xylanase when sugar
cane bagasse was used as a substrate in a glass-column
reactor with forced aeration. The airflow rate had a sig-
nificant effect on enzyme activity, whereas initial mass of
bagasse had none. The highest yield of xylanase
(1597 U/g) was obtained operating in the bioreactor at
the optimal conditions: airflow rate (6 l/h g) and sub-
strate (8 g).
A packed-bed bioreactor with four stages was con-
structed and operated for microbial production of citric
acid by A. niger using apple pomace as a substrate.
Under the optimised conditions, 124 g citric acid was
produced from 1 kg dry apple pomace with yield of
80% based on total sugar (
).
3. Conclusion
Critical analysis of the literature shows that produc-
tion of relevant compounds for the food processing
industry by SSF offers several advantages. It has been
well established that enzyme titres produced in SSF sys-
tems are much higher than the achieved in SmF ones.
Although the reasons for this are not clear, this fact is
kept in mind while developing novel bioreactors for
SSF processes.
Acknowledgments
The authors are grateful to Xunta de Galicia (local
government of Spain) for the financial support of the re-
search post of Susana Rodrı´guez Couto under the pro-
gramme Isidro Parga Pondal.
References
Acun˜a-Arguelles, M. E., Gutierrez-Rojas, M., Viniegra-Gonza´lez, G.,
& Favela-Torres, E. (1994). Effect of water activity on exo-
pectinase production by Aspergillus niger CH
4
on solid state
fermentation. Biotechnology Letters, 16, 23–28.
Acun˜a-Arguelles, M. E., Gutierrez-Rojas, M., Viniegra-Gonza´lez, G.,
& Favela-Torres, E. (1995). Production and properties of three
pectinolytic activities produced by Aspergillus niger in submerged
and solid-state fermentation. Applied and Microbiology Biotech-
nology, 43, 808–814.
Alkorta, I., Garbisu, G., Llama, M. J., & Serra, J. L. (1998). Industrial
applications of pectic enzymes: a review. Process Biochemistry, 33,
21–28.
Antier, P., Minjares, A., Roussos, S., Raimbault, M., & Viniegra-
Gonza´lez, G. (1993). Pectinase-hyperproducing mutants of Asper-
gillus niger C28B25 for solid-state fermentation of coffee pulp.
Enzyme and Microbial Technology, 15, 254–260.
Antier, P., Minjares, A., Roussos, S., & Viniegra-Gonzalez, G. (1993).
New approach for selecting pectinase producing mutants of
Aspergillus niger well adapted to solid state fermentation. Biotech-
nology Advances, 11, 429–440.
Auria, R., Palacios, J., & Revah, S. (1992). Determination of the
interparticular effective diffusion coefficient for CO
2
and O
2
in solid
state fermentation. Biotechnology Bioengineering, 39, 898–902.
Babu, K. R., & Satyanarayana, T. (1995). a-Amylase production by
thermophilic Bacillus coagulans in solid state fermentation. Process
Biochemistry, 30, 305–309.
Bai, Z. H., Zhang, H. X., Qi, H. Y., Peng, X. W., & Li, B. J. (2004).
Pectinase production by Aspergillus niger using wastewater in solid
state fermentation for eliciting plant disease resistance. Bioresource
Technology, 95, 49–52.
Water cooling
Sampling port
Pneumatic system
Air inlet
Compressor
Mesh with substrate
Culture medium level
Gas exit
Compressor
Fig. 5. Scheme of an immersion bioreactor (humidified air; mechanical
agitation).
298
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
Baker, R. A., & Wicker, L. (1996). Current and potential applications
of enzyme infusion in the food industry. Trends Food Science
Technology, 7, 279–284.
Barrios-Gonzalez, J., Gonzalez, H., & Mejia, A. (1993). Effect of
particle size, packing density and agitation on penicillin production
in solid state fermentation. Biotechnology Advances, 11, 539–
547.
Benjamin, S., & Pandey, A. (1996a). Lipase production by Candida
rugosa on copra waste extract. Indian Journal of Microbiology, 36,
201–204.
Benjamin, S., & Pandey, A. (1996b). Optimization of liquid media for
lipase production by Candida rugosa. Bioresource Technology, 55,
167–170.
Benjamin, S., & Pandey, A. (1997a). Coconut cake—a potent substrate
for the production of lipase by Candida rugosa in solid state
fermentation. Acta Biotechnologica, 17, 241–251.
Benjamin, S., & Pandey, A. (1997b). Enhancement of lipase produc-
tion during repeated batch cultivation using immobilised Candida
rugosa. Process Biochemistry, 32, 437–440.
Benninga, H. (1990). A history of lactic acid making. Boston: Kluwer
Academic Publishers.
Berger, RG. (1995). Aroma biotechnology. Berlin: Springer-Verlag.
Besson, I., Creuly, C., Gros, J. B., & Larroche, C. (1997). Pyrazine
production by Bacillus subtilis in solid state fermentation on
soybeans. Applied Microbiology and Biotechnology, 47, 489–495.
Boccas, F., Roussos, S., Gutierrez, M., Serrano, L., & Viniegra, G. G.
(1994). Production of pectinase from coffee pulp in solid state
fermentation system: selection of wild fungal isolate of high
potency by a simple three-step screening technique. Journal of Food
Science Technology, 31, 22–26.
Bohlmann, G., & Yoshida, Y. (2000). Biodegradable polymers.
Chemical economic handbook report.
Bramorski, A., Christen, P., Ramirez, M., Soccol, C. R., & Revah, S.
(1998). Production of volatile compounds by the edible fungus
Rhizopus oryzae during solid-state cultivation on tropical agro-
industrial substrates. Biotechnology Letters, 20, 359–362.
Bramorski, A., Soccol, C. R., Christen, P., & Revah, S. (1998). Fruit
aroma production by Ceratocystis fimbriata in static cultures from
solid agro-industrial wastes. Revista de Microbiologia, 28, 208–212.
Castilho, L. R., Alves, T. L. M., & Medronho, R. A. (1999). Recovery
of pectinolytic enzymes produced by solid state culture of Asper-
gillus niger. Process Biochemistry, 34, 181–186.
Castilho, L. R., Alves, T. L. M., & Medronho, R. A. (2000).
Production and extraction of pectinases obtained by solid state
fermentation of agro-industrial residues with Aspergillus niger.
Bioresource Technology, 71, 45–50.
Chang, T. S., Siddiq, M., Sinha, N. K., & Cash, J. N. (1994). Plum
juice quality affected by enzyme treatment and fining. Journal of
Food Science, 59, 1065–1069.
Chen, H. Z. (1992). Advances in solid-state fermentation. Research and
Application of Microbiology (China), 3, 7–10.
Christen, P., Angeles, N., Corzo, G., Farres, A., & Revah, S. (1995).
Microbial lipase production on a polymeric resin. Biotechnology
Techniques, 9, 597–600.
Christen, P., Bramorski, A., Revah, S., & Soccol, C. R. (2000).
Characterization of volatile compounds produced by Rhizopus
strains grown on agro-industrial solid wastes. Bioresource Tech-
nology, 71, 211–215.
Cordova, J., Nemmaoui, M., Ismaili-Alaoui, M., Morin, A., Roussos,
S., Raimbault, M., et al. (1998). Lipase production by solid state
fermentation of olive cake and sugar cane bagasse. Journal of
Molecular Catalysis B: Enzymatic, 5, 75–78.
Demain, A. L. (2000). Small bugs, big business: the economic power of
the microbe. Biotechnology Advances, 18, 499–514.
Dominguez, A., Costas, M., Longo, M. A., & Sanroman, A. (2003). A
novel application of solid state culture: production of lipases by
Yarrowia lipolytica. Biotechnology Letters, 25, 1225–1229.
Domı´nguez, A., Rivela, I., Rodrı´guez Couto, S., & Sanroma´n, M. A.
(2001). Design of a new rotating drum bioreactor for ligninolytic
enzyme production by Phanerochaete chrysosporium grown on an
inert support. Process Biochemistry, 37, 549–554.
dos Santos, M. M., Souza da Rosa, A., DalÕBoit, S., Mitchell, D. A., &
Krieger, N. (2004). Thermal denaturation: is solid-state fermenta-
tion really a good technology for the production of enzymes?
Bioresource Technology, 93, 261–268.
Durand, A., Renaud, R., Almanza, S., Maratray, J., Diez, M., &
Desgranges, C. (1993). Solid-state fermentation reactors: from lab
scale to pilot plant. Biotechnology Advances, 11, 591–597.
Echevarria, J., Leon, J. A. R., Espinosa, M. E., & Delgado, G. (1991).
Optimization of solid state fermentation of sugarcane by Asper-
gullus niger considering the particle size effect. Acta Biotech-
nologica, 11, 15–22.
Ferron, G., Bonnarame, P., & Durand, A. (1996). Prospects of the
microbial production of food flavours. Trends Food Science
Technology, 7, 285–293.
Francis, F., Sabu, A., Nampoothiri, K. M., Ramachandran, S.,
Ghosh, A., Szakacs, G., et al. (2003). Use of response surface
methodology for optimizing process parameters for the production
of a-amylase by Aspergillus oryzae. Biochemical Engineering
Journal, 107–115.
Garzo´n, C. G., & Hours, R. A. (1991). Citrus waste: an alternative
substrate for pectinase production in solid-state culture. Biore-
source Technology, 39, 93–95.
Ghildyal, N. P., Ramalaishna, M., Lonsane, B. K., & Karantb, N. G.
(1992). Gaseous concentration gradients in tray type solid-state
fermentors—Effects on yields and productivities. Bioprocess Engi-
neering, 8, 67–72.
Gombert, A. K., Pinto, A. L., Castilho, L. R., & Freire, D. M. G.
(1999). Lipase production by Penicillium restrictum in solid-state
fermentation using babassu oil cake as substrate. Process Bio-
chemistry, 35, 85–90.
Gross, R. A., & Kalra, B. (2002). Biodegradable polymers for the
environment. Science, 297, 803–807.
Gutierrez-Rozas, M., Cordova, J., Auria, R., Revah, S., & Favela-
Torres, E. (1995). Citric acid and polyols production by Aspergillus
niger at high glucose concentration in solid state fermentation on
inert support. Biotechnology Letters, 17, 219–224.
Hang, Y. D., & Woodams, E. E. (1985). Grape pomace a novel
substrate for microbial production of citric acid. Biotechnology
Letters, 7, 253–254.
Hang, Y. D., Woodams, E. E., & Jang, K. Y. (1995). Enzymatic
conversion of sucrose to kestose by fungal extracellular fructosyl
transferase. Biotechnology Letters, 17, 295–298.
Hesseltine, C. W. (1977). Solid-state fermentation. Part 1. Process
Biochemistry, 12, 24–27.
Ho¨lker, U., Ho¨fer, M., & Lenz, J. (2004). Biotechnology advantages of
laboratory-scale solid-state fermentation with fungi. Applied
Microbiology and Biotechnology, 64, 175–186.
Hours, R. A., Voget, C. E., & Ertola, R. J. (1988a). Some factors
affecting pectinase production from apple pomace in solid-state
cultures. Biological Wastes, 24, 147–157.
Hours, R. A., Voget, C. E., & Ertola, R. J. (1988b). Apple pomace as
raw material for pectinases production in solid state culture.
Biological Wastes, 23, 221–228.
Ismail, A. S. (1996). Utilization of orange peels for the production of
multi-enzyme complexes by some fungal strains. Process Bio-
chemistry, 1, 645–650.
Ito, K., Yoshida, K., Ishikawa, T., & Kobayashi, S. (1990). Volatile
compounds produced by fungus Aspergillus oryzae in rice koji and
their changes during cultivation. Journal of Fermentation and
Bioengineering, 70, 169–172.
Jaeger, K. E., & Reetz, M. T. (1998). Microbial lipases from
versatile tools for biotechnology. Trends in Biotechnology, 16,
396–403.
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
299
Janssens, L., de Pooter, H. L., Vandamme, E. J., & Schamp, N. M.
(1992). Production of flavours by microorganisms. Process Bio-
chemistry, 27, 195–215.
Johns, M. R., & Stuart, D. M. (1991). Production of pigments by
Monascus purpureus in solid culture. Journal of Industrial Micro-
biology, 8, 23–28.
Kamini, N. R., Mala, J. G. S., & Puvanakrishnan, R. (1998). Lipase
production from Aspergillus niger by solid-state fermentation using
gingelly oil cake. Process Biochemistry, 33, 505–511.
Khare, S. K., Krishana, J., & Gandhi, A. P. (1995). Citric acid
production from Okara (soy residue) by solid state fermentation.
Bioresource Technology, 54, 323–325.
Krishna, C., & Chandrasekaran, M. (1996). Banana waste as substrate
for a-amylase production by Bacillus subtilis (CBTK 106) under
solid-state fermentation. Applied and Microbiology Biotechnology,
46, 106–111.
Kumar, D., Jain, V. K., Shanker, G., & Srivastava, A. (2003a).
Utilisation of fruits waste for citric acid production by solid state
fermentation. Process Biochemistry, 38, 1725–1729.
Kumar, D., Jain, V. K., Shanker, G., & Srivastava, A. (2003b). Citric
acid production by solid state fermentation using sugarcane
bagasse. Process Biochemistry, 38, 1731–1738.
Larroche, C., Besson, I., & Gros, J. B. (1999). High pyrazine
production by Bacillus subtilis in solid substrate fermentation on
ground soy-beans. Process Biochemistry, 34, 67–74.
Lichtfield, J. H. (1996). Microbial production of lactic acid. Advances
in Applied Microbiology, 42, 48–95.
Liu, B. L., & Tzeng, Y. M. (1999). Water content and water activity for
the production of cyclodepsipeptide in solid state fermentation.
Biotechnology Letters, 21, 657–661.
Lonsane, B. K., Saucedo-Castuneda, G., & Raimbault, M. (1992).
Scale-up strategies for solid-state fermentation systems. Process
Biochemistry, 27, 259–273.
Maldonado, M. C., Navarro, A., & Callieri, D. A. S. (1986).
Production of pectinases by Aspergillus sp. using differently
pretreated lemon peel as the carbon source. Biotechnology Letters,
8, 501–504.
Malhotra, V. P., Raina, R. K., & Sanjay, R. (2000). Application of
polymers in innovative medical devices. Medical Plastics Data
Service.
Available
from
http://www.medicalplasticsindia.com/
mpds/2000/may/technology01.htm
.
Marek, A., & Bednarski, W. (1996). Some factors affecting lipase
production by yeasts and filamentous fungi. Biotechnology Letters,
18, 1155–1160.
Martins, E. S., Silva, D., Da Silva, R., & Gomes, E. (2002). Solid state
production of thermostable pectinases from thermophilic Ther-
moascus aurantiacus. Process Biochemistry, 37, 949–954.
Medeiros, A., Pandey, A., Christen, P., Fontoura, P. S. G., Freitas, R.
J. S., & Soccol, C. R. (2001). Aroma compounds produced by
Kluyveromyces marxianus in solid-state fermentation on packed
bed column bioreactor. World Journal Microbiology Biotechnology,
17, 767–771.
Milagres, A. M. F., Santos, E., Piovan, T., & Roberto, I. C. (2004).
Production of xylanase by Thermoascus aurantiacus from sugar
cane bagasse in an aerated growth fermentor. Process Biochemistry,
39, 1387–1391.
Miranda, O. A., Salgueiro, A. A., Pimentel, M. C. B., Lima Filho,
J. L., Melo, E. H. M., & Duran, N. (1999). Lipase production
by a Brazilian strain of Penicillium citrinum using an industrial
residue. Bioresource Technology, 69, 145–147.
Muniswaran, P. K. A., & Charyulu, N. C. L. N. (1994). Solid substrate
fermentation of coconut coir pith for cellulase production. Enzyme
and Microbial Technology, 16, 436–440.
Nagel, F. J., Tramper, J., Bakker, M. S., & Rinzema, A. (2001a).
Temperature control in a continuously mixed bioreactor for solid-
state fermentation. Biotechnology and Bioengineering, 72, 219–
230.
Nagel, F. J., Tramper, J., Bakker, M. S., & Rinzema, A. (2001b).
Model for on-line moisture-content control during solid-state
fermentation. Biotechnology and Bioengineering, 72, 231–243.
Navarrete-Bolan˜os, J. L., Jime´nez-Islas, H., Botello-Alvarez, E., Rico-
Martı´nez, R., & Paredes-Lo´pez, O. (2004). An optimization study
of solid-state fermentation: xanthophylls extraction from marigold
flowers. Applied and Microbiology Biotechnology, 65, 383–390.
Naveena, B. J., Altaf, Md., Bhadrayya, K., Madhavendra, S. S., &
Reddy, G. (2005). Direct fermentation of starch to
L
(+) lactic acid
in SSF by Lactobacillus amylophilus GV6 using wheat bran as
support and substrate: medium optimization using RSM. Process
Biochemistry, 40, 681–690.
Naveena, B. J., Altaf, M. d., Bhadriah, K., & Reddy, G. (2005).
Selection of medium components by Plackett–Burman design for
production of
L
(+) lactic acid by Lactobacillus amylophilus GV6 in
SSF using wheat bran. Bioresource Technology, 96, 485–490.
Ohnishi, K., Yoshida, Y., & Sekiguchi, J. (1994). Lipase production of
Aspergillus oryzae. Journal of Fermentation and Bioengineering, 77,
490–495.
Pandey, A. (1991). Aspects of fermenter design for solid-state
fermentations. Process Biochemistry, 26, 355–361.
Pandey, A. (1992). Recent process developments in solid-state
fermentation. Process Biochemistry, 27, 109–117.
Pandey, A. (1994). Solid-state fermentation. In A. Pandey (Ed.) (pp. 3–
10). New Delhi: Wiley Eastern Publishers.
Pandey, A., Ashakumary, L., Selvakumar, P., & Vijayalakshmi, K. S.
(1994). Influence of water activity on growth and activity of A.
niger for glucoamylase production in solid-state fermentation.
World Journal of Microbiology and Biotechnology, 10, 485–486.
Pandey, A., Azmi, W., Singh, J., & Banerjee, U. C. (1999b).
Biotechnology: food fermentation. In V.K. Joshi, A. Pandey
(Eds.) (Vol. I, pp. 383–426). New Delhi: Educational Publishers
& Distributors.
Pandey, A., Benjamin, S., Soccol, C. R., Nigam, P., Krieger, N., &
Soccol, V. T. (1999c). The realm of microbial lipases in biotech-
nology. Biotechnology and Applied Biochemistry, 29, 119–131.
Pandey, A., Nigam, P., & Vogel, M. (1988). Simultaneous sacchari-
fication and protein enrichment fermentation of sugar beet pulp.
Biotechnology Letters, 10, 67–72.
Pandey, A., Selvakumar, P., Soccol, C. R., & Nigam, P. (1999a). Solid
state fermentation for the production of industrial enzymes.
Current Science, 77, 149–162.
Pandey, A., Soccol, C. R., & Mitchell, D. (2000a). New developments
in solid state fermentation: I—bioprocesses and products. Process
Biochemistry, 35, 1153–1169.
Pandey, A., Soccol, C. R., Nigam, P., Brand, D., Mohan, R., &
Roussos, S. (2000b). Biotechnological potential of coffee pulp and
coffee husk for bioprocesses. Biochemistry Engineering Journal, 6,
153–162.
Pandey, A., Soccol, C. R., Rodriguez-Leon, J., & Nigam, P. (2001).
Solid-state fermentation in biotechnology. New Delhi: Asiatech
Publishers Inc.
Pandey, A., Soccol, C. R., Selvakumar, P., Soccol, V. T., Krieger, N.,
& Fontana, J. D. (1999d). Recent developments in microbial
inulinase. Applied Biochemistry and Biotechnology, 81, 1–18.
Pastrana, L. M., Gonzalez, M. P., Pintado, J., & Murado, M. A.
(1995). Interactions affecting gibberellic acid production in solid
state fermentation. Enzyme and Microbial Technology, 17, 784–
790.
Pastore, G. M., Park, Y. K., & Min, D. B. (1994). Production of a
fruity aroma by Neurospora from beiju. Mycological Research, 98,
25–35.
Pintado, J., Lonsane, B. K., Gaime-Perraud, I., & Roussos, S. (1998).
On-line monitoring of citric acid production in solid-state culture
by respirometry. Process Biochemistry, 33, 513–518.
Prado, F. C., Vandenberghe, L. P. S., Lisboa, C., Paca, J., Pandey, A.,
& Soccol, C. R. (2004). Relation between citric acid production and
300
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
respiration rate of Aspergillus niger in solid-state fermentation.
Engineering in Life Sciences, 4, 179–186.
Prapulla, S. G., Subhaprada, V., & Karanth, N. G. (2000). Microbial
production of oligosaccharides: a review. Advances in Applied
Microbiology, 47, 299–343.
Raimbault, M. (1998). General and microbiological aspects of solid
substrate fermentation EJB Electronic. Journal of Biotechnology, 1.
Available from
Ramachandran, S., Patel, A. K., Nampoothiri, K. M., Francis, F.,
Nagy, V., Szakacs, G., et al. (2004). Coconut oil cake—a potential
raw material for the production of a-amylase. Bioresource Tech-
nology, 93, 169–174.
Rao, P. V., Jayaraman, K., & Lakshmanan, C. M. (1993). Production
of lipase by Candida rugosa in solid-state fermentation. I: deter-
mination of significant process variables. Process Biochemistry, 28,
385–389.
Richter, K., & Tra¨ger, A. (1994).
L
(+) lactic acid from sweet sorghum
by submerged and solid-state fermentations. Acta Biotechnologica,
14, 367–378.
Rivela, I., Rodrı´guez Couto, S., & Sanroma´n, A. (2000). Extracellular
ligninolytic enzyme production by Phanerochaete chrysosporium in
a new solid-state bioreactor. Biotechnology Letters, 22, 1443–
1447.
Rivera-Mun˜oz, G., Tinoco-Valencia, J. R., Sanchez, S., & Farres, A.
(1991). Production of microbial lipases in a solid-state fermentation
system. Biotechnology Letters, 13, 277–280.
Robinson, T., & Nigam, P. (2003). Bioreactor design for protein
enrichment of agricultural residues by solid state fermentation.
Biochemical Engineering Journal, 13, 197–203.
Rodriguez Couto, S., Barreiro, M., Rivela, I., Longo, M. A., &
Sanroman, A. (2002). Performance of a solid-state immersion
bioreactor for ligninolytic enzyme production: evaluation of
different operational variables. Process Biochemistry, 38, 219–
227.
Rodrı´guez Couto, S., Rivela, I., Mun˜oz, M. R., & Sanroma´n, A.
(2000). Ligninolytic enzyme production and the ability of decolo-
urisation of Poly R-478 in packed-bed bioreactors by Phanero-
chaete chrysosporium. Bioprocess Engineering, 23, 287–293.
Roussos, S., Raimbault, M., Prebois, J. P., & Lonsane, B. K. (1993).
Zymotis, a large scale solid state fermenter-design and evaluation.
Applied Biochemistry and Biotechnology, 42, 37–52.
Sangeetha, P. T., Ramesh, M. N., & Prapulla, S. G. (2004). Production
of fructosyl transferase by Aspergillus oryzae CFR 202 in solid-
state fermentation using agricultural by-products. Applied Micro-
biology and Biotechnology, 65, 530–537.
Sargantanis, J., Karim, M. N., Murphy, V. G., Ryoo, D., & Tengerdy,
R. P. (1992). Effect of operating conditions on solid sub-
strate fermentation. Biotechnology and Bioengineering, 42, 149–
158.
Sarrette, M., Nout, M. J. R., Gervais, P., & Rombouts, F. M. (1992).
Effect of water activity on production and activity of Rhizopus
oligosporus polysaccharides. Applied Microbiology and Biotech-
nology, 37, 420–425.
Schwan, R. F., Cooper, R. M., & Wheals, A. E. (1997). Endopoly-
galacturonase secretion by Kluyveromyces marxianus and other
cocoa pulp-degrading yeasts. Enzyme and Microbial Technology,
21, 234–244.
Seitz, E. W. (1994). Fermentation production of pyrazines and ter-
penoids for flavour and fragrances. In A. Gabeliman (Ed.),
Bioprocess
production
of
flavour,
fragrance,
and
colour
(pp. 95–134). New York: Wiley.
Shah, N. D., Chattoo, B. B., Baroda, R. M., & Patiala, V. M. (1993).
Starch hydrolysate, an optimal and economical source of carbon
for the secretion of citric acid by Yarrow lipolytica. Starch, 45,
104–109.
Shankaranand, V. S., & Lonsane, B. K. (1994). Ability of Aspergillus
niger to tolerate metal ions and minerals in solid state fermentation
system for production of citric acid. Process Biochemistry, 29,
29–37.
Shojaosadati, S. A., & Babaripour, V. (2002). Citric acid production
from apple pomace in multi layer packed bed solid state bioreactor.
Process Biochemistry, 37, 909–914.
Singh, S. A., Plattner, H., & Diekmann, H. (1999). Exopolygalact-
uronate lyase from a thermophilic Bacillus sp. Enzyme and
Microbial Technology, 25, 420–425.
Smail, T., Salhi, O., & Knapp, J. S. (1995). Solid state fermentation of
carob pods by Aspergillus niger for protein production-effect of
particle size. World Journal of Microbiology and Biotechnology, 11,
171–173.
Soares, M., Christen, P., Pandey, A., & Soccol, C. R. (2000). Fruity
flavour production by Ceratocystis fimbriata grown on coffee
husk in solid state fermentation. Process Biochemistry, 35, 857–
861.
Soccol, C. R., Iloki, I., Marin, B., & Raimbault, M. (1994).
Comparative production of alpha-amylase, glucoamylase and
protein enrichment of raw and cooked cassava by Rhizopus strains
in submerged and solid state fermentations. Journal of Food Science
and Technology, 31, 320–323.
Soccol, C. R., Marin, B., Rimbault, M., & Labeault, J. M. (1994).
Potential of solid state fermentation for production of
L
(+) lactic
acid by Rhizopus oryzae. Applied Microbiology and Biotechnology,
41, 286–290.
Soccol, R. S., & Vandenberghe, L. P. S. (2003). Overview of applied
solid-state fermentation in Brazil. Biochemical Engineering Journal,
13, 205–218.
Sodhi, H. K., Sharma, K., Gupta, J. K., & Soni, S. K. (2005).
Production of a thermostable a-amylase from Bacillus sp. PS-7 by
solid state fermentation and its synergistic use in the hydrolysis of
malt starch for alcohol production. Process Biochemistry, 40,
525–534.
Stamford, T. L., Stamford, N. P., Coelho, L. C., & Araujo, J. M.
(2001). Production and characterization of a thermostable alpha-
amylase from Nocardiopsis sp. endophyte of yam bean. Bioresource
Technology, 76, 137–141.
Stredansky, M., & Conti, E. (1999). Xanthan production by solid state
fermentation. Process Biochemistry, 34, 581–587.
Stredansky, M., Conti, E., Navarini, L., & Bertocchi, C. (1999).
Production of bacterial exopolysaccharides by solid substrate
fermentation. Process Biochemistry, 34, 11–16.
Stuart, D. M., Mitchell, D. A., Johns, M. R., & Litster, J. D. (1999).
Solid-state fermentation in rotating drum bioreactors: operat-
ing variables affect performance through their effects on trans-
port phenomena. Biotechnology and Bioengineering, 63, 383–
391.
Sugawara, E., Hashimoto, S., Sakurai, Y., & Kobayashi, A. (1994).
Formation by yeast of the HEMF (4-hydroxy-2 (or 5)-ethyl-5 (or
2)-methyl-3 (2H)-furanone) aroma components in Miso with
aging. Bioscience Biotechnology and Biochemistry, 58, 1134–
1135.
Suryanarayan, S. (2003). Current industrial practice in solid state
fermentations for secondary metabolite production: the Biocon
India experience. Biochemical Engineering Journal, 13, 189–
195.
Sutherland, I. W. (1998). Novel and established applications of
microbial
polysaccharides.
Trends
Biotechnology,
16,
41–
46.
Tsuchiya, K., Nagashima, T., Yamamoto, Y., Gomi, K., Kitamoto,
K., & Kumagai, C. (1994). High level secretion of calf chymosin
using a glucoamylase-prochymosin fusiongene in Aspergillus
oryzae. Bioscience Biotechnology and Biochemistry, 58, 895–
899.
Vandenberghe, L. P. S., Soccol, C. R., Pandey, A., & Lebeault, J. M.
(2000). Solid-state fermentation for the synthesis of citric acid by
Aspergillus niger. Bioresource Technology, 74, 175–178.
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
301
Vickroy, T. B. (1985). Lactic acid. In H. W. Blanch & S. Drew (Eds.).
Comprehensive biotechnology—the principles, applications and
regulation of biotechnology in industry, agriculture and medicine
(Vol. 3). Toronto: Dic Pergamon Press.
Yamguchi, N., Toda, T., Teramoto, T., Okuhira, T., Sugawara, E., &
Ito, T. (1993). Studies on pyrazine formation by Bacillus natto. V.
Effect of sugars on microbiological pyrazine formation by Bacillus
natto in synthetic liquid medium. Nippon Shokuhin Kogyo Gakkai-
shi, 40, 841–848.
Zadrazil, F., & Punia, A. K. (1995). Studies on the effect of particle
size on solid state fermentation of sugarcane bagasse into animal
feed using white-rot fungi. Bioresource Technology, 54, 85–
87.
Zheng, Z., & Shetty, K. (2000). Solid state production of polygalac-
turonase by Lentinus edodes using fruit processing wastes. Process
Biochemistry, 35, 825–830.
302
S.R. Couto, M
a
.
A. Sanroma´n / Journal of Food Engineering 76 (2006) 291–302
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