Process Biochemistry 27 (1992) 109-I 17

Recent Process Developments in Solid-State

Fermentation

Ashok Pandey
Regional Research Laboratory, Council of Scientific & Industrial Research, Trivandrum-695019, India

(Received 26 June 1991; accepted 12 October 1991)

Solid-state fermentation has gained renewed interest and fresh attention

from researchers owing to its importance in recent developments in
biomass energy conservation, in solid waste treatment and in its
application to produce secondary metabolites. Solid-state fermentation
must be seen as a technique for producing higher yields of desired
products; in addition, there are various other advantages. Solid-state
fermentation processes are also of considerable promise on the pollution
front.
This paper reviews the recent process developments in solid-state
fermentation, especially in the past decade. The importance of
microorganism and water activity are dealt with, and the substrates and
end-products of the process are outlined. Advances in growth
parameters and kinetic studies are also discussed.

INTRODUCTION Confucius book of Chau-Lai). Miso, a fermented

Solid-state (substrate) fermentation (SSF) is gen-
erally defined as the growth of microorganisms on
solid materials in the absence of near-absence of
free water. The substrate, however, must contain
enough moisture, which exists in the absorbed form
within the solid matrix.
Solid-state fermentations have been used since
antiquity. The use of soy sauce koji in China, Japan
and south-east Asia goes back as far as 1000 years
BP and probably 3000 years BP in China (see
Corresponding author : Dr A. Pandey. Telephone : 0471
76774; Telex: 0435 232; Fax: 0471 75186.
109
ProcessBiochemistry 0032-9592/92/$5.00 Cl 1992 Elsevier Science Publishers
food in Japan, China, Taiwan, the Philippines,
Indonesia and in the Orient has also traditionally
been prepared by SSF. Tempeh is yet another
important solid-state fermented food used in Indo-
nesia, New Guinea and Surinam.
The history and developments in SSF have been
reviewed by several authors.lmg Aidoo et al. made
an attempt to trace the history of growth of
microorganisms on solid substrates, a process that
has led to the development of the term solid-state
fermentation. Table 1 gives a brief account of the
historical developments in SSF. Hesseltine2v3 re-
viewed and discussed SSF for the production of
secondary metabolites and fermentation of animal
wastes. Cannel and Moo-Young4*5 also reviewed
Ltd, England.
110 Ashok Pandey

Table 1. History and development of solid-state fermentation Table 2. Enzymes produced by Aspergillus niger

Period Development I Alpha amylase 11. Lipase

2. Beta amylase 12. Cellulase
2600 B.C. Bread making by Egyptians 3. Maltase 13. Amidase
Before birth of Cheese making by Penicillium 4. Lactase 14. Glucose oxidase
Christ in Asia roqueforti 5. Catalase 15. Glucose dihydrogenase
(Recorded history 6. Proteinase 16. Urease
1ooo BP) 7. Trehalase 17. Insulase
2500 BP Fish fermentation/preservation 8. Tannase 18. Melibase
with sugar, starch, salts, etc. 9. Dipetalase 19. Zymase (trace)
2500 BP Koji process 10. Polypetalase
7th Century Koji process from China to Japan
by Buddhist priests
18th Century Vinegar from pomace
18th Century Gallic acid used in tanning,
Microorganisms for SSF
printing, etc.
1860-1900 Sewage treatment Based upon the type of microorganism involved,
190&1920 Fungal enzymes, kojic acid SSF processes can be categorized into two main
192&1940 Fungal enzymes, gluconic acid, groups, viz. natural (indigenous) SSF and pure
rotary drum fermenter, citric acid
19*1950 Fantastic development in culture SSF (individual or mixed). Ensiling and
fermentation industry. Penicillin composting are the two best SSF processes which
production by SSF and submerged utilize natural microflora. SSF with pure cultures is
fermentation
Steroid transformation by fungal
known since antiquity, viz. the koji process with
195&1960
cultures Aspergillus oryzae. Pure cultures are generally used
196&1980 Production of mycotoxins, protein in industrial SSF processes to improve the control
enriched feed
of the substrate utilization and end-product forma-
198cpresent Various other products like alcohol,
gibberllic acid tion. Bioconversion of agricultural residues (straw,
etc.) to fungal biomass (protein) using two different
a Based upon Ref. 1. microorganisms, viz. Chaetomium cellulolyticum
and Candida utilis, is a typical example of mixed-
culture SSF systems. In nature, SSF is oftenly
the development of SSF as it was initiated and carried out by mixed cultures in which several
improved in the oriental food industry and com- microorganisms show development and symbiotic
posting. They also compared SSF with liquid cooperation.
(submerged) fermentation, SmF and described Selection of a suitable microorganism is one of
advantages and disadvantages of SSF. Knapp and the most important criteria in SSF. The vast
Howell7 reviewed the literature on SSF from the majority of wild type microorganisms are incapable
fundamental point of view of the microorganisms of producing commercially acceptable yields of the
and microbial enzymes involved. The advantages product. The importance of the microorganism can
and disadvantages of SSF have been discussed by be seen from the fact that a culture of A. niger can
several other authors.*O SSF processes are used on produce as many as 19 types of enzymes (Table 2)
a commercial scale for the production of different while alpha amylase alone can be produced by some
types of fermented foods, fungal metabolites and 28 different types of culture (Table 3).26
for bioconversion of organic wastes into useful Several groups of the microorganisms can grow
products.11-24 on solid substrates.2 Filamentous fungi, however,
This article does not intend to go deeply into the have the best capability to grow in the absence of
historical developments, advantages and dis- free water. The cultivation of the filamentous fungi
advantages and production of any specific products on solid substrates has been widely used for different
by SSF; rather an attempt is made to review the purposes at laboratory scale, e.g. for koji fer-
more recent developments in microbial types, mentation,28 for lignocellulose fermentation, for
growth kinetics and process parameters for SSF. fungal spores,30.31 and for mycotoxin production.8x
The paper also does not provide information on Among the filamentous fungi, three classes, viz.
design of bioreactor systems for SSF, which has Phycomycetes (Mucor and Rhizopus), Ascomycetes
been separately discussed. (Aspergillus and Penicillium) and Basidiomycetes
(white rot fungi, Polyporus) have most commonly
been used. Table 4 gives a comprehensive view of
 Recent process developments in solid-state fermentation 111

Table 3. Different cultures from which alpha amylase may be produced

1. Bacillus acidocaldrius 15. Thermonospora viridis

2. B. amyloliquefaciens 16. Pseudomonas saccharophilus
3. B. caldolyticus 17. AspergilIus awamori
4. B. coagulans 18. A. batatae
5. B. licheniformis 19. A. candidus
6. B. mesentericus 20. A. niger
7. B. stearothermophilus 21. A. oryzae
8. B. subtilis 22. A. terricola
9. Other Bacillus spp. 23. A. usmii
10. Eacteroides amylophilus 24. Mucor meihei
11. Closiridium acetobutylicum 25. M. pusillus
12. Thermoactinomyles vulgaricus 26. Neurospora crassa
13. Thermonospora curvata 27. Peniciliium expansum
14. T. vulgaricus 28. Rhizopur javanicus

different microorganisms used in recent years in trichum pulverulentum. Saccharomyces cerevisiae

various SSF processes.
A. niger has most commonly been used for
protein enrichment33*34*42 of the substrate as well
as for enzyme production (cellulase,40s41 amylase,43
glucoamylase,44T45 beta glucosidase*l and acid
protease*). It0 et aZ.48 reported production of
alcohols, ketones and aldehydes in rice fermentation
using A. oryzae. A. flaws has been used for
protease enzyme production.50 Rhizopus oligosporus
has widely been used for protein enrichment and
kinetic studies related with SSF process.52m55
Karanthl reported the production of fungal rennet
by R. oligosporus and Mucor meihei. Cultures of
Trichoderma spp. have been used for protein
enrichment and enzyme production in pure or
mixed cultures.*~ Munoz et a1.6g reported six
species of Penicillium, two species of Rhizopus and
Geotrichum candidum and M. meihei for lipase
enzyme production. Maximum enzyme activity was
obtained with P. candidum, P. camembertii and M.
meihei. Several other species of Penicillium have
been used for the production of various enzymes,
e.g. hydrolaseT4 and pectic enzymes. 75 Gonzalez
et a1.73 reported production of penicillin by a non-
sterile SSF process on bagasse impregnated with
culture medium using P. chrysogenum. Polyporus
spp. have been used for protein enrichment of the
substrate, for lignin degradation7 and for cellu-
lase and ligninase enzymes.77 Kumar and Lon-
sane*, reported the production of gibberllic acid
by Fusarium monoliforme and Gibberella jiijikuroi.
Ramesh and Lonsanes2* g3 reported the production
of bacterial alpha amylase with Bacillus licheni-
formis.
Several yeasts have been used for protein en-
richment or ethanol fermentation in SSF. Smith
et a1.99 reported protein production with Sporo-
has most commonly been used for ethanol pro-
duction.00-103 Hanlo reported protein enrichment
of straw with Candida utiiis.
Water activity factor
In general, the types of the microorganisms that can
grow in SSF systems are determined by the water
activity factor, a,. Water activity is defined as the
relative humidity of the gaseous atmosphere in
equilibrium with the substrate. The a, of the
substrate quantitatively expresses the water re-
quirement for microbial activity.27
- Vm+
a, = ~
55.5
where
V= number of ions formed
m = molar concentration of the solute,
#J = molar osmotic coefficient, and
55.5 = molar concentration of an aqueous
solution of pure water.
Pure water has an a, = 1.00 and a, will decrease
with the addition of solutes. Bacteria mainly grow
at higher a, values while filamentous fungi and
some yeasts can grow at lower a, values (06-0~7).
The microorganisms, which can grow and are
capable of carrying out their metabolic activities at
lower a, values are suitable for SSF processes.
Water activity of the substrate has been proposed
as the condition of growth and viability of the
microorganisms. I The importance of a, in SSF
has widely been studied.36*109-116It has been shown
that in the course of fungal growth in SSF, high
water activities favour sporulation while low water
activities favour spore germination or mycelial
growth.11,2 Oriol et aZ.38 made estimations of
112 Ashok Pandey

Table 4. Microorganisms, substrates and products in SSF processes

Microorganism Process/product Substrate Reference

Aspergillus spp. Protein enrichment Starchy raw materials Czajkowska & Ilnickaa3
Aspergillus niger Protein Banana waste Baldensperger er aLa4
Koji for ethanol Corn Han & SteinbergB5
Growth and kinetic Cassava Raimbault & Alazard,3B
studies Castaneda et al.,a
Oriol et aZ.38
Aflatoxins Cassava Gonzalez et al.38
Protein, cellulase Corn cob Singh et al.40
Cellulase, beta Bagasse, corn cobs, Madamwar et a1.41
glucosidase sawdust, computer
cards
Protein enrichment and Citrus peel Rodriguez ef al.42
growth studies
Amylase Wheat bran Malik & Khan43
Glucoamylase Wheat bran Pandeya4
Glucoamylase Wheat bran Ramakrishna et a1.45
Aspergillus phoenicis Beta glucosidase Sugar beet pulp Deschamps & Huet*
Aspergillus oryzae Alcohol, aldehydes, Rice It0 ef al.R
ketones
Aspergillus terreus Protein Sugarcane by-products Blanc0 er a1.48
Aspergillus flaws Protease Wheat bran Malathi & Chakraborty
Aspergillus carbonarius Pectinase Wheat bran Karantl?
Aspergillus niveus Catalase Wheat bran Karanthsl
Rhizopus oligosporus Kinetic studies Cassava Mitchell et al.52
Kinetic studies Cassava Mitchell et ~1.~~
Protein food Chickpea Lopez et al.5
Growth studies Soyabean Rathbun & Shule?&
Rhizopos oryzae Protein Cassava Daubresse er a1.b6
Rhizopus oligosporus and Rennet Wheat bran Karanth51
Mucor meihei
Trichoderma viride Cheese aroma Agar Gervais et aZ.5
Trichoderma viride Protein Sugar beet pulp Durand & Chereau5s
Hydrolase enzymes Wheat bran Atev et al.==
Trichoderma reesei Hydrolase enzymes Wheat bran Pauaiotov et al.so
T. reesei and yeast Protein Cassava Opoku & Adogael
T. viride and A. niger Growth and kinetics, Sugar beet pulp Desgranges & Durengsa
cellulase and amylase
T. reesei and Endomycopsis Protein Wheat straw Laukevics et a1.63
jibuleger
T. reesei, Chaetomium Protein Wheat straw Abdullah et al.@
cellulolyticum, Candida utilis
Trichoderma spp., A. ustus, Cellulase, beta Wheat bran and rice Shamala & Sreekantiaha5
Botritis spp. and Sporotriehum glucosidase, xylanase straw
pulverulentum
T. viride, S. thermophile and Growth studies Sugar beet pulp Grajekse
Thermoascus auranticus
T. reesei and Fusarium Protein Sugar beet pulp Nigam & VogeP
oxysporum
T. reesei, A. terreus, Protein Wheat Pollard and bran Rhodes & Broderickas
Chaetomium virescens and
Shizophyllum commune
Penicillium spp., Geotrichum Lipases Wheat bran Munoz et al.ss
candidum, Mucor meihei,
R. arrhizus and R. delemer
Penicillium roqueforti Growth studies Buckwheat seeds Desfarges et aZ.O
Growth studies and Inert porus Larroche et al.
spore production
Penicillium capsulatum Enzymes Sugar beet pulp Considine et al.*
Penicillium chrysogenum Penicillin Bagasse Gonzalez ef a1.3
PenicilIium resticulosum Proteins and hydrolase Wheat bran Atev et aLT4
enzymes
Penicillium charlesii, Pectic enzymes Citrus pulp pellets Siessere & Said5
Talaromyces flavus and
Tubercularia vulgaris
 Recent process developments in solid-state fermentation 113

Tubercularia vulgaris Pectic enzyme Citrus pulp pellets Vieira et al.

Polyporus spp. Cellulase and ligninase Bagasse Nigam et al.
Protein Bagasse Nigams
Lignin degradation Oat straw Bone & Muno2*
Pleurotus spp. Protein Saccharum munja Gujral et a/.*
residues
Coprinus spp. Protein Wheat straw Yadav*
Coprinus jimetarius Growth studies Wheat straw Singh et al.82
Coprinus fimetarius Protein Wheat straw Kumar & Singha
Phanaerochaeta chrysosporium Lignin/nonlignin Birch lignin Mudgett & Paradiss4
conversion
Chaetomium globusum and Protein Lignocellulosics Gruju et ds5
Chaetomium ceihdolyticum
Fusarium monobforme and Gibberllic acid Wheat bran Kumar & Lonsanes6
Gibberlla fujikuroi
Gibberlla fujikta-oi Gibberllic acid Wheat bran Kumar & Lonsanes7
Mucor meihei Rennet Wheat bran Thakur et aLB8
Glocophyllum trabeum Growth studies Pine sawdust Edurado et aLB9
Panus tignium Hydrogen peroxide Wheat straw Maltseva er aI.*O
Leniinula edodes Extracellular enzymes Lianocellulosics Mishra & Leathmana
BaciNus Iichenzformis Alpha amylase Wheat bran Ramesh & Lonsanea2
Alpha amylase Wheat bran Ramesh & Lonsanes3
Bacillus sub tilis Protease Wheat bran Hafiz et al.g4
Slreptomyces clavuligerus and Cephalosporin Barley Get-mini & Demaing5
Cephalosporium aermonium
Phubia tremellosa Delignitication Aspen wood ReidsB
Neurospora spp. Aroma Polished rice Yamauchi et al.g
Neurospora crasse CM Case and beta Straw Macris et 01.~~
glucosidase
Sporotrichum pulveruhwtum Protein Cassava Smith er al.@
Sarcharomyces cerevisiae Ethanol Sweet sorghum Kargi et aI.O
Ethanol Sugar beet pulp Cachet er a!.
Ethanol, protein Fodder beets Gibbons et al.oz
Ethanol Apple pomace Kazimierzlo3
Candida utilis Protein Straw Hanlo
Amylolytic yeasts (Pichia Protein Sweet potato YanglO
barroniz)
Thermophilic yeasts Ethanol Corn gits Sat0 et al.B
(S. cerevisiae)
Wine yeasts Ethanol Grape pomace Hang et ~1.~

total water, consumed water and residual water in
SSF of cassava starch. A theoretical calculation
based on the Ross equation showed that water
activity of the substrate decreased to O&5 towards
the end of the culture.
Regulation of the a, of the solid substrate can be
controlled by the relative humidity (RH) of the air.
Murrell and ScotP5 used a biological system with
static control of RH. Gervais et ~1.~ reported a SSF
process allowing the control of ~1,. Gervais et ~1.~~
studied the influence of a, on cheese aroma (2-
heptanone) production in SSF and found that
optimum aroma production occurred at a, = O-98.
Gervais and Sarrettel also studied the influence of
a, on aroma production by T. viride.
Numerous experiments have demonstrated the
influence of a, on the metabolism of micro-
organisms.LR-lnl Gervais et ~1.~~ reported the
influence of a, on a solid substrate on growth rate
and sporogenesis of filamentous fungi. Grajek and
GervaiP3 studied the influence of water activity on
enzyme biosynthesis and enzyme activities pro-
duced by fungi. Gervais 124 developed a new sensor
which allowed continuous a, measurement of
submerged and solid-state fermentations.
The a, of the medium is a fundamental parameter
for mass transfer of the water and solutes across the
cell membrane. The control of this parameter could
be used to modify the metabolic production or
excretion of a microorganism.125
A kinetic model which relates the rate constant of
the death of the microbial cells to water activity and
temperature has been proposed by Moser,12 using
the following equation
gd&
k = k,u,exp- RT
where constants k, and EA are calculated from the
experimental value of u,.
114 Ashok Pandey
Substrates for SSF
Natural as well as synthetic substances can be used
as substrates in SSF processes. The major organic
materials available in nature are polymeric in
structure, e.g. polysaccharides, proteins and lignins.
In general, all these can be used by microorganisms
as substrate (carbon source). Solid substrates used
in SSF are insoluble in water; in practice water is
absorbed onto substrate particles which can be used
by microorganisms for growth and metabolic
activity. Bacteria and yeast grow on the surface of
the substrate while fungal mycelia penetrate into
the particles of the substrate.
The utilization of solid substrates by the micro-
organisms is affected by several physical and
chemical factors. Among the physical factors,
assessibility of substrate to microbes, film effects
and mass effects are important.7 The physical
morphology, especially porosity and particle size of
the substrate, governs the accessible surface area to
the organism. Among the chemical factors, the
chemical nature of the substrate (degree of polymer-
ization and crystallinity) is an important criterion.
The heterogenous nature of the natural solid
substrates creates problems in the fermentation
kinetic studies. Use of synthetic polymers or
substances like agar and gelatin provide hom-
ogenous solid substrate and are good for such
studies.57.127 Georgiou and ShulerlZ8 used nutrient
agar as a model substrate. Gelatin and Kappa-
carrageenan have been used by Wei et aZ.12 and
Mitchell et af.lso to produce a model substrate to
mimic the growth of microorganisms.
Knapp and Howell reviewed the literature on
the effects of alteration of substrate particle size on
SSF. It was reported that substrates with finer
particles showed improved degradation due to an
increase in surface area.ll, Huang et a1.133
reported that greater growth of the fungal cultures
was stimulated by smaller particle size substrate.
Particles of 30 mesh were found to be the optimum
particle size. Recently, Pandey34 found that enzyme
productivities were higher with a substrate that
contained particles of mixed sizes varying from
smaller than 180 ,um to bigger than 1.4 mm.
Nowadays, in soya sauce koji preparation the
whole soyabeans have been replaced by defatted
soyabeans, and wheat bran has replaced wheat
(Xyu). 135 XyL? also reported that the present
process of Chinese soy sauce production is different
from the procedure as described by Young &
Wood. The koji and mash stages have been
shortened to 24 h and 30 days, or even 15 days,
respectively.
In general, several agricultural or agro-industrial
residues, which are cellulosic or starchy in nature,
have been used in SSF processes. Table 4 indicates
the substrates used for protein enrichment or
secondary metabolite production in recent years.
Kinetic studies/growth factors for SSF
Kinetic studies of SSF processes have not received
much attention in comparison to SmF. This is
probably because of the difficulties associated with
the measurement of various factors, due to the
heterogenecity of the medium which are structurally
and nutritionally complex. In addition, control of
fermentation parameters is also difficult. Use of the
synthetic model substrate which is homogenous in
nature has appeared as a way of dealing with this
problem. 12-130
Temperature, moisture, humidity, aeration, agi-
tation and pH are the most important factors which
govern monitoring and controlling of SSF. High
substrate moisture results in decreased substrate
porosity which in turn prevents oxygen penetration.
This may facilitate bacterial contamination. At the
same time, low moisture levels lead to poor
microbial growth and poor accessibility to
nutrients. Hang and WoodamsJ? studied the effect
of substrate moisture on citric acid production and
found that it critically affected mould growth and
activity. Ramesh and Lonsane also reported the
critical importance of moisture content of the
substrate in alpha amylase production in a SSF
system. They observed a large reduction in the
production of enzyme in standardized wheat bran
medium under SSF when the moisture content of
the medium was higher than the standardized value.
Similar results were noted by the author39 during
SSF of wheat bran for glucoamylase enzyme
production.
Gonzalez et al.ldo developed and discussed the
stoichiometric relation and yield parameters of
growth of fungi in SSF and illustrated the possibility
that water may be a limiting substrate. Kumar and
Lonsaneldl stressed the need for optimal physical
factors, including moisture content, for gibberellic
acid production in SSF.
Temperature is yet another critical parameter
that can affect SSF processes. Due to the production
of large amounts of metabolic heat, the fermenting
substrate temperature shoots up.44~a5~ 14~,143 Rath-
bun and Shulers5 studied the heat and mass transfer
effects in static SSF. The time-dependent values of
temperature, mol.% 0, and mol.% CO, were
measured. They observed that during a tempeh
fermentation, temperature gradients could be as
 Recent process developments in solid-state fermentation 115

high as 3 C cm-l. Lai et al.144 described a math- Department of Science, Technology and Environ-
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diffusivity in SSF process of Sorghum brewing.
Habib145 described a bioengineering model for
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