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Solid-state fermentation — are there any biotechnological advantages? Udo Ho¨ lker 1 and Ju¨ rgen
Solid-state fermentation — are there any biotechnological advantages? Udo Ho¨ lker 1 and Ju¨ rgen

Solid-state fermentation — are there any biotechnological advantages?

Udo Ho¨ lker 1 and Ju¨ rgen Lenz 2

Solid-state fermentation (SSF) has developed in eastern countries over many centuries, and has enjoyed broad application in these regions to date. By contrast, in western countries the technique had to compete with classical submerged fermentation and, because of the increasing pressure of rationalisation and standardisation, it has been widely superseded by classical submerged fermentation since the 1940s. This is mainly because of problems in engineering that appear when scaling up this technique. However, there are several advantages of SSF, for example high productivities, extended stability of products and low production costs, which say much about such an intensive biotechnological application. With increasing progress and application of rational methods in engineering, SSF will achieve higher levels in standardisation and reproducibility in the future. This can make SSF the preferred technique for special fields of application such as the production of enzymes and food.

Addresses 1 Institute of Cellular and Molecular Botany, University of Bonn, Kirschallee 1, 53115 Bonn, Germany 2 Bioreact GmbH, Mu¨ lheimer Str. 26, 53840 Troisdorf, Germany

Corresponding author: Ho¨ lker, Udo (

Current Opinion in Microbiology 2005, 8:301–306

This review comes from a themed issue on Ecology and industrial microbiology Edited by Sergio Sa´ nchez and Betty Olson

Available online 5th May 2005

1369-5274/$ – see front matter # 2005 Elsevier Ltd. All rights reserved.

DOI 10.1016/j.mib.2005.04.006


The term solid-state fermentation (SSF) denotes cultiva- tion of microorganisms on solid, moist substrates in the absence of a free aqueous phase; that is, at average water activities (defined as the relative humidity of the gaseous phase in equilibrium with the moist solid) significantly below 1 [1 ]. In a broader definition, SSF can be seen as including processes during which microorganisms are cultivated in the presence of a liquid phase at maximal substrate concentrations [2] or on inert carriers [3].

To assess the possible advantages of the biotechnological uses of SSF relative to those of classical submerged fermentations (SmFs), different aspects of the two, including historical, biological, ecological and engineer-

ing aspects, will first be considered separately, and then compared from the economical viewpoint.

Historical aspects

When compared with the SSF processes that have long been established in Asian countries, SSF in western countries appears to still be in its infancy. SSF is the state-of-the-art technology that is used in many applica- tions in the food industry in Asia. For example, SSF is used in the production of enzyme-rich Koji (which is made of rice and uses Aspergillus oryzae as an enzymatic starter for different hydrolytic processes), in the sacchar- ification of rice used for the production of alcoholic beverages such as sake, in the production of cellulase (which uses Trichoderma strains), and in the production of angkak, which is rice that is coloured red by Monascus purpurea metabolites. In Asia, enzymes and metabolites are commonly produced on a large scale by SSF processes that may be thousands of years old. These processes vary widely depending on locality, and are transmitted as secret formulas from father to son.

In western countries, SSF had to compete with the technically easier SmF, which was more strongly devel- oped from the 1940s onwards because of the necessity to produce antibiotics on a large scale. Today, SSF has been nearly completely abandoned because of the pressure of increasing industrial rationalisation and standardisation. In the east, by contrast, SSF survived in many areas in the form of labour-convenient, hand-made processes; how- ever, some of the traditional SSF processes have also been further developed to achieve higher industrial rationalisa- tion through stronger automation.

Biological features

SSF processes simulate the living conditions of many higher filamentous fungi. Ascomycetes, basidiomycetes and deuteromycetes developed in terrestrial habitats on wet substrates. For this reason, SSF is the cultivation method of choice for instance in cases in which the biotechnological process is required to include morpho- logical and metabolic differentiation into substrate-pene- trating and aerial hyphae, such as in the production of conidiospores. Hence, SSF is used in the food industry in the production of conidiospores for the production of mould cheese (for which conidiospores of Penicillium roquefortii or P. camemberti are needed) or salami improve- ment (for which conidiospores of P. nalgiovensis can be used) and also in the field of production of plant protec- tion agents (which uses Coniothyrium minitans).

302 Ecology and industrial microbiology

Higher fungi and their enzymes, spores or metabolites are well adjusted to growth on solid wet substrates. For instance, fungal spores produced by SSF show higher stability, are more resistant to drying and exhibit higher germination rates for extended periods of time after freeze-drying than do spores produced by SmF. This can be attributed to the fact that the conidiospores produced in SSF have a higher hydrophobicity, as was demonstrated with Penicillium oxalicum [4], or also pos- sess, apart from a higher hydrophobicity caused by a hydrophobin-like protein, a stronger cell wall and a smal- ler volume (this has been documented with Trichoderma harzianum [5]).

A direct comparison between the SSF and SmF cultiva-

tion modes of fungi is difcult to make because the two processes differ. Studies of the production of fungal

enzymes in SSF have repeatedly shown that SSF, in comparison with SmF, provides higher volumetric pro- ductivities, is less prone to problems with substrate inhi- bition and yields enzymes with a higher temperature or pH stability. Also, the fermentation time may be shorter and the degradation of the enzymes by undesirable proteases is minimised [6]. Interesting studies in this eld have explored the physiological differences that arise during the growth of microbial cells in the two types

of processes. In the well-studied fungus Aspergillus oryzae,

water activity seems to play a decisive role in this context [7]. This is apparent from the accumulation of glycerol, erythritol and arabinitol [8 ], and from the induction of glucoamylase genes, the promoters of which have been shown to contain motifs that suggest induction by tem- perature or water activity [9], under SSF conditions.

Similar results were also obtained in studies of the pro- duction of secondary metabolites [10]. When grown on solid substrates, numerous fungi have shown a markedly altered spectrum of secondary metabolites. As described

in streptomycetes, the production of secondary metabo-

lites often seems to be associated with the formation of aerial hyphae and the onset of spore production in the stationary growth phase. Various secondary metabolites have even been shown to be produced solely during growth on solid substrates, although the fungi in question can be readily cultivated in submerged processes. For this reason, SSF could play an interesting role in the screening for new metabolites and also in the production of these metabolites by wild-type fungi.

neous saccharication and fermentation that can be regu- lated by the joint action of aerobic and facultative anaerobic processes via the availability of oxygen. In addition, SSF offers the possibility to explore and to use interactions between fungi by properly regulating the water activity as a selection parameter depending on the different water demands of the individual co-cultivated fungi. Many SSF processes used in the food industry make use of mixed cultures that are important for the avour of the foods produced. For example, bamboo sprouts can be processed by undened mixed cultures that, during the fermentation process, produce a variety of metabolites including 29 volatile substances with aroma-active properties [16].

Despite the situation when using lamentous fungi, SSF has only found restricted applications in processes using unicellular organisms. SSF has, for instance, been used in the production of endospores (Bacillus thuringiensis) [17], secondary metabolites by means of aerial mycelium of streptomycetes [18] or alcohol by yeast on solid sugar-rich substrates, which can be conveniently combined with simultaneous or previous hydrolysis of the polysacchar- ides by extracellular fungal enzymes. Another example is the production of organic acids (such as vinegar produc- tion performed in Asia for thousands of years) or the traditional fermentation of foods [19] by microbial mixed cultures that involve yeasts and bacteria.

Ecological features

Ecological (environment-protective) advantages of SSF reect the fact that the processes are conducted in the absence of a free aqueous phase. This results in minimum water consumption and thus a low production of efuent water by the process, obviation of the necessity to use antifoam agents, and the possibility of conducting SSF processes in certain applications under semi-sterile con- ditions. Because SSF processes are performed at water activities below 1, the growth of contaminating bacteria and yeasts is minimised. This means that, in certain cases, energy- and instrumentation-demanding sterilisation pro- cesses can be eliminated.

Another additional environmentally friendly feature of SSF is that, in many cases, it can use agricultural wastes as carbon and energy sources. The production of enzymes or organic acids makes frequent use of plant remnants as carbon sources and inducers or mediators.

Engineering features


the employment of microbial mixed cultures, SSF offers

The most serious reasons why SSF has not yet found a


option that cannot be achieved by SmF. This makes use

broad use in western countries are engineering problems,


the fact that, during growth on natural substrates, fungal

the low amenability of the processes to standardisation

consortia secrete a broad spectrum of enzymes [1113]. During co-cultivation of different fungi, some of the indi- vidual enzyme activities show synergistic increase [14], whereas others remain unchanged [15]. Filamentous fungi and yeasts can be broadly applied in processes of simulta-

and the limited reproducibility of the results. Scale-up represents a particular bottleneck because several differ- ent gradients (temperature, humidity, substrate concen- tration, and so on), which can arise in the course of the process, can have an adverse effect especially in static

solid bed processes but also in processes involving sub- strate agitation, such as those performed in rotating drums. Inter-relationships among environmental factors such as oxygen content, moisture level and temperature contribute to the difcult regulation of these parameters. The microbial growth under aerobic conditions in the bioreactor results in a considerable production of heat that causes fast increase in temperature. This effect, which is desirable in composting, is often fatal for biotechnological processes because a large part of the enzymes produced during the fermentation can be heat-denatured at the end of the process [20 ]. In the absence of a free aqueous phase, the produced heat is difcult to remove, for example, via the bioreactor double walls. Instead, the cooling of the process takes place through evaporation. However, this requires very high aeration rates because the rising metabolic activity and the associated increased heat production have to be overcompensated by high aeration intensity. The water escaping by evaporation has to be in many cases replenished, and this can lead to

Figure 1

Solid-state fermentation Ho¨ lker and Lenz


undesirable local increase in water activity during static processing. In semi-sterile processes, this increased water activity can in turn have adverse effects by facil- itating the growth of bacterial contaminants, whereas in sterile fermentations, it may bring about local insufcient oxygen supply to the microorganisms. Substrate mixing may help but it should be noted that many microorgan- isms respond very sensitively to the shear stress caused by it. Another factor that is difcult to account for is the production of metabolic water by aerobic microorgan- isms, which can cause problems especially in the forma- tion of conidiospores.

Despite all these problems, the development of rational, at least partly computer-controlled methods of process development within the past decade brought advances in the eld, leading to a better understanding of biological and physical processes that take place during the SSF (Figure 1), and paving the way to new starting points for further industrialisation of the process [21]. These

O 2 g g Aerial hyphae Mycelial mat l Substrate surface s s s CO
O 2
Aerial hyphae
Mycelial mat
Substrate surface
CO 2
Substrate pores
NH 3
2 O
Penetrative hyphae
Current Opinion in Microbiology

Schematic of some of the microscale processes that occur during solid-state fermentation (SSF). After sporulation, fungal hyphae develop into a mycelial mat (black), which spreads over the surface of the particles that comprise the solid substrate (brown). From the mycelial mat, aerial hyphae protrude into the gaseous space, whereas others penetrate the substrate by growing into liquid-filled pores. At normal moisture levels, the void spaces between the aerial hyphae are most likely to be filled with gas (g), whereas the void spaces within the mycelial mat and within the substrate are filled with liquid (l). The metabolic activities shown mainly occur near the substrate surface and within the pores; however, exposed regions of the mycelium (for instance the aerial hyphae) also show metabolism and there can be a transport of substances from the penetrative to the aerial hyphae. Hydrolytic enzymes (light blue), which are produced by the mycelium, diffuse to the solid matrix and catalyse the degradation of macromolecules into smaller units (green). The latter are taken up by the fungus to serve as nutrients. O 2 is consumed and CO 2 , H 2 0, heat and interesting biochemical products are produced during fermentation. Hence, gradients develop within the biofilm that, for instance, force O 2 to diffuse from the gaseous phase into deeper regions of the biofilm (lilac) and CO 2 to diffuse from these regions to the gaseous phase (red). Heat development (Q; orange) leads to a fast increase in temperature (T), which is a serious problem during SSF. Heat is therefore removed from the substrate not only via conduction but also by evaporation, which is part of the complex balance of water in the system (dark blue). Beside evaporation, water balance includes water uptake by the mycelium in the course of growth, water consumption during hydrolysis reactions and water production through respiration. As another important factor, local pH, might be changed owing to the release of carbon acids and the exchange of ammonia (grey). The biochemical products of interest (magenta) that are released into the solid matrix and the liquid-filled spaces during fermentation might absorb to the solid and might have to be extracted for further use at the end of the process. All these and many other phenomena can strongly influence process performance during SSF.

304 Ecology and industrial microbiology

developments include progress in the modelling of micro- bial growth on solid substrates and energy and mass transfer in different types of bioreactors [2226,27 , 28,29 ,30], formulation of scale-up rules and recommen- dations for optimal design and scale-up of industrial bioreactors [3133], new methods for measuring the spatial distribution of process parameters, evaluation of parameters that are difcult to measure on-line such as humidity and biomass [3436], the use of statistical meth- ods for parameter optimisation [3742], and the develop- ment of new strategies for process control in SSF [4345]. Interestingly, the practical development of commercially employed bioreactors proceeded largely independently of the results of the above studies. This was in part because of the difculties in converting the general statements of rational studies pertaining to bioreactor and process design into concrete guidelines for specic applications. The reason again is the lack of sufcient bodies of process data needed for validating the mathematical models. Hence, the commercially employed industrial bioreactors often still represent empirical developments of traditional processes, or they are large but technically relatively low- level facilities restricted correspondingly to non-sterile and robust processes suitable for manufacture of bulk products (e.g. bioreactors for Koji fermentation). The engineering shortcomings are then compensated by high space and time gains and low production costs so that these processes can be operated with satisfactory economy. Although a variety of different SSF bioreactors are avail- able on the laboratory and pilot scale [46], only some of these variants have the potential for scale-up to the industrial level because large-volume operations may give rise to effects, such as the inuence of compaction, which are not perceptible on a smaller scale.

Economic viewpoint

The described biological and environmental benets of SSF can bring direct economical advantages that have much to say for an intensive biotechnological application of SSF. For instance, calculation of the costs of production of crude enzymes by SSF showed that, for example in the case of cellulase production, the economical efciency is higher by a factor of 100 than in the case of SmF [47]. This striking difference has several reasons. In addition to the much cheaper growth substrates, the minimised require- ments for sterility and low requirements for instrumenta- tion and equipment, this approach can make downstream processes in the production of enzymes unnecessary. In general, this holds for those cases when the fermented substrate can be used directly in a follow-up process such as the saccarication of renewable raw materials employed in ensiling, production of bio-ethanol or biogas, in the pulp- and-paper industry, textiles industry or in the production of animal fodder supplements. This is usually made possible thanks to the high concentrations of enzymes in the solid substrates, which makes further concentrating processes unnecessary. The enzyme activities that have been mea-

sured in solid substrates can reach levels common in commercial enzyme preparation [48].


Yet, despite the multitude of the above advantages, a broad application of SSF is, at least at present, hard to envisage. The main obstacles are the low amenability of the process to regulation, the strongly heterogeneous fermentation conditions and the ensuing frequently unsa- tisfactory reproducibility of the results, difcult scale-up, the often unfeasible biomass determination and compli- cated product purication by downstream processes resulting from the use of heterogeneous organic growth substrates. For these reasons, SSF will never replace or supersede the established processes for the production of enzymes and metabolites by bacteria or yeasts that have for decades been successfully cultivated in submerged bioreactors and also optimised as heterologous expression systems. Yet only in special cases, SSF can be applicable also with unicellular organisms.

A different situation exists in the cultivation of differ- entiating microorganisms, in which SSF simulates the natural growth conditions and its biological and environ- mental advantages can outweigh the engineering pro- blems. In principle, SSF processes can be economically operated on a large scale despite their engineering dif- culties when the technical demands (for example, in non- sterile processes or in those that do not require further expensive product processing) and hence also the pro- duction costs are low. In these cases, SSF can provide substantial advantages over SmF.

At any rate, even though further progress in the eld of SSF can be expected to bring improved biomonitoring and thereby optimisation and higher standardisation of industrial processes, the future applications of SSF for the above purposes can be expected to be mainly limited to those that even today exhibit substantial advantages above all the production of spores, screening for second- ary metabolites and enzymes, protein enrichment of bre-rich natural materials, degradation and bioconver- sion of plant material [49 ], and co-cultivation of several fungi for the production of enzymes [50 ] and foods.

References and recommended reading

Papers of particular interest, published within the annual period of review, have been highlighted as:

of outstanding interest

of special interest

1. Pandey A: Solid-state fermentation. Biochem eng 2003,

The author gives a compact, yet very informative, overview of the present status and applications of SSF. He envisages SSF to become on a par with SmF technology, with respect to industrial application in the future, if the current trends in rational engineering are continued.

2. Mitchell DA, Berovic M, Krieger N: Biochemical engineering of solid state bioprocessing. Adv Biochem Eng Biotechnol 2000,



3. Ooijkaas LP, Weber F, Buitelaar RM, Tramper J, Rinzema A:

Defined media an inert supports: their potential as solid-state fermentation production system. Trends Biotechnol 2000,


4. Pascual S, de Cal A, Magan N, Melgarejo P: Surface hydrophobicity, viability and efficacy in biological control of Penicillium oxalicum spores produced in aerial and submerged culture. J Appl Microbiol 2000, 89:847-853.

5. Munoz GA, Agosin E, Cotoras M, San Martin R, Volpe D:

Comparison of aerial and submerged spore properties for Trichoderma harzianum. FEMS Microbiol Lett 1995, 125:63-70.

6. Ho¨ lker U, Ho¨ fer M, Lenz J: Biotechnological advantages of laboratory-scale solid-state fermentation with fungi. Appl Microbiol Biotechnol 2004, 64:175-186.

7. te Biesebeke R, Ruijter G, Rahardjo YSP, Hoogschagen MJ, Heerikhuisen M, Levin A, van Driel KGA, Schutyser MAI, Dijksterhuis J, Zhu Y et al.: Aspergillus oryzae in solid state fermentation. FEMS Yeast Research 2002, 2:245-248.

8. Ruijter GJ, Visser J, Rinzema A: Polyol accumulation by

Aspergillus oryzae at low water activity in solid-state fermentation. Microbiology 2004, 150:1095-1101. In this interesting paper, it was shown that Aspergillus oryzae accumu-

lates high concentrations of polyols at water activities between 0.96 and

0.97 during solid-state processing, which seems to be typical for this type

of processing. In addition, four different polyol dehydrogenases were detected, which might be induced by osmotic stress, as concluded by the authors.


Ishida H, Hata Y, Kawato A, Abe Y, Suginami K, Imayasu S:

Identification of functional elements that regulate the glucoamylase-encoding gene (glab) expressed in solid-state culture of Aspergillus oryzae. Curr Genet 2000, 37:373-379.


Robinson T, Singh D, Nigam P: Solid-state fermentation: a promising microbial technology for secondary metabolite production. Appl Microbiol Biotechnol 2001, 55:284-289.


Koroleva OV, Gavrilova VP, Stepanova EV, Lebedeva VI, Sverdlova NI, Landesman EO, Yavmetdinov IS, Yaropolov AI: Production of lignin modifying enzymes by co-cultivated white-rot fungi Cerrena maxima and Coriolus hirsutus and characterization of laccase from Cerrena maxima. Enz Microbial Technol 2002,



Stepanova EV, Koroleva OV, Vasilchenko LG, Karapetyan KN, Landesman EO, Yavmetdinov IS, Kozlov YP, Ranbinovich ML:

Fungal decomposition of oat straw during liquid and solid- state fermentation. Prikl Biokhim Mikrobiol 2003, 39:65-74.


Yang YH, Wang BC, Wang QH, Xiang LJ, Duan CR: Research on solid-state fermentation on rice chaff with a microbial consortium. Colloids Surf B Biointerfaces 2004, 34:1-6.


Gutierrez-Correa M, Portal L, Moreno P, Tengerdy RP: Mixed culture solid substrate fermentation of Trichoderma reesei with Aspergillus niger on sugar cane bagasse. Bioresour Technol 1999, 68:173-178.


Massadeh MI, Yusoff WMW, Omar O, Kader J: Synergism of cellulase enzymes in mixed culture solid substrate fermentation. Biotechnol Lett 2001, 23:1771-1774.


Fu SG, Yoon Y, Bazemore R: Aroma-active compounds in fermented bamboo shoots. J Agric Food Chem 2002,



Hongzhang C, Fujian X, Zhonghou T, Zuohu L: A novel industrial- level reactor with two dynamic changes of air for solid-state fermentation. J Biosci Bioeng 2002, 93:211-214.


Ellaiah P, Srinivasulu B, Adinarayana K: Optimisation studies on neomycin production by a mutant strain of Streptomyces marinensis in solid state fermentation. Process Biochem 2003,



Han B-Z, Rombouts FM, Nout MJR: A Chinese fermented soybean food. Int J Food Microbiol 2001, 65:1-10.


Muller dos Santos M, Souza das Rosa A, DalBoit S, Mitchell DA,

Krieger N: Thermal denaturation: is solid-state fermentation really a good technology for the production of enzymes? Bioresour Technol 2004, 93:261-268.

Solid-state fermentation Ho¨ lker and Lenz


This study investigates critically the effect of heat production on pro- ductivity during SSF. Overheating, which results from difculties in heat removal, is a severe problem when trying to scale up a process.

21. Lenz J, Ho¨ fer M, Krasenbrink JB, Ho¨ lker U: Computational and physical methods in solid-state fermentation. Appl Microbiol Biotechnol 2004, 65:9-17.

22. Ashley VM, Mitchell DA, Howes T: Evaluating strategies for overcoming overheating problems during solid-state fermentation in packed bed bioreactors. Biochem Eng J 1999,


23. Gervais P, Molin P: The role of water in solid-state fermentation. Biochem Eng J 2003, 13:85-101.

24. Ikasari L, Mitchell DA: Two-phase model of the kinetics of growth of Rhizopus oligosporus in membrane culture. Biotechnol Bioeng 2000, 68:619-627.

25. Mitchell DA, Tongta A, Stuart DM, Krieger N: The potential for establishment of axial temperature profiles during solid-state fermentation in rotating drum bioreactors. Biotechnol Bioeng 2002, 80:114-122.

26. Mitchell DA, von Meien OF, Krieger N: Recent developments in modelling of solid-state fermentation: heat and mass transfer in bioreactors. Biochem Eng J 2003, 13:137-147.

27. Mitchell DA, von Meien OF, Krieger N, Dalsenter FDH: A review of

recent developments in modelling of microbial growth kinetics and intraparticle phenomena in solid-state fermentation. Biochem Eng J 2004, 17:15-26. In an excellent review, the authors describe various approaches to the modelling of those processes that occur on the particle scale during SSF. Mechanistic modelling of, for instance, intra-particle diffusion of

enzymes, O 2 and hydrolysis products helps us to understand how these effects inuence fungal growth and could potentially limit bioreactor performance.

28. Oostra J, le Comte EP, van der Heuvel JC, Tramper J, Rinzema A:

Intra-particle oxygen diffusion limitation in solid-state fermentation. Biotechnol Bioeng 2001, 75:13-24.

29. Schutyser MAI, de Pagter P, Weber FJ, Briels WJ, Boom RM,

Rinzema A: Substrate aggregation due to aerial hyphae during discontinuously mixed solid-state fermentation with Aspergillus oryzae: experiments and modelling. Biotechnol Bioeng 2003, 83:503-513. This study is the rst attempt to quantify and to predict the effect of mycelial bonds on particle mixing and vice versa during SSF. The tensile strength of mycelial bonds was measured using a new device on the basis of two wheat-dough-plates and texture analysis. Using these measure-

ments, mixing experiments performed with Aspergillus oryzae that grows on wheat grain in a rotating drum bioreactor were compared with results obtained from computer simulations.

30. Schutyser MAI, Weber FJ, Briels WJ, Rinzema A, Boom RM:

Heat and water transfer in a rotating drum containing solid substrate particles. Biotechnol Bioeng 2003,


31. Hardin MT, Mitchell DA, Howes T: Approach to designing rotating drum bioreactors for solid-state fermentation on the basis of dimensionless design factors. Biotechnol Bioeng 2000,


32. Mitchell DA, Pandey A, Sangsurasak P, Krieger N: Scale-up strategies for packed-bed bioreactors for solid-state fermentation. Process Biochem 1999, 35:167-178.

33. Weber FJ, Tramper J, Rinzema A: A simplified material and energy balance approach for process development and scale- up of Coniothyrium minitans conidia production by solid-state cultivation in a packed-bed reactor. Biotechnol Bioeng 1999,


34. Aldarf M, Amrane A, Prigent Y: Reconstruction of the biomass history from carbon and nitrogen substrate consumption, ammonia release and proton transfer during solid cultures of Geotrichum candidum and Penicillium camembertii. Appl Microbiol Biotechnol 2002, 58:823-829.

35. Bellon-Maurel V, Orliac O, Christen P: Sensors and measurements in solid state fermentation: a review. Process Biochem 2003, 38:881-896.

306 Ecology and industrial microbiology

36. Pena y Lillo M, Perez-Correa R, Agosin E, Latrille E: Indirect measurement of water content in an aseptic solid substrate cultivation pilot-scale bioreactor. Biotechnol Bioeng 2001,


37. Abdel-Fattah YR, Olama ZA: l-Asparaginase production by Pseudomonas aeruginosa in solid-state culture: evaluation and optimization of culture conditions using factorial designs. Process Biochem 2002, 38:115-122.

38. Banerjee R, Bhattacharyya BC: Evolutionary operation as a tool of optimization for solid-state fermentation. Biochem Eng J 2003, 13:149-155.

39. Francis F, Sabu A, Nampoothiri KM, Ramachandran S, Ghosh S, Szakacs G, Pandey A: Use of response surface methodology for optimizing process parameters for the production of a-amylase by Aspergillus oryzae. Biochem Eng J 2004, 15:107-115.

40. Hamidi-Esfahani Z, Shojaosadati SA, Rinzema A: Modelling of simultaneous effect of moisture and temperature on A. niger growth in solid-state fermentation. Biochem Eng J 2004,


41. Kar B, Banerjee R, Bhattacharyya BC: Optimization of physicochemical parameters for gallic acid production by evolutionary operation-factorial design technique. Process Biochem 2002, 37:1395-1401.

42. Park YS, Kang SW, Lee JS, Hong SI, Kim SW: Xylanase production in solid state fermentation by Aspergillus niger mutant using statistical experimental designs. Appl Microbiol Biotechnol 2002, 58:761-766.

43. Nagel FJI, Tramper J, Bakker MSN, Rinzema A: Temperature control in a continuously mixed bioreactor for solid-state fermentation. Biotechnol Bioeng 2001, 72:219-230.


Nagel FJI, Tramper J, Marjolein SN, Bakker MSN, Rinzema A: Model for on-line moisture-content control during solid-state fermentation. Biotechnol Bioeng 2001,



Thibault J, Acuna G, Perez-Corea R, Jorquera H, Molin P, Agosin E: A hybrid representation approach for modelling complex dynamic bioprocesses. Bioprocess Eng 2000,



Durand A: Bioreactor designs for solid state fermentation. Biochem Eng J 2003, 13:113-125.


Tengerdy RP: Cellulase production by solid substrate fermentation. J Sci Ind Res 1996, 55:313-316.


Ho¨ lker U, Janssen M, Lenz J: Directed selective solid-phase culturing of stable microbial mixed populations for the continuous preparation of dened enzyme and metabolite mixtures. PCT / EP2004 / 051234.


Tengerdy RP, Szakacs G: Bioconversion of lignocellulose

in solid substrate fermentation. Biochem Eng J 2003,


This review presents the state-of-the-art of lignocellulose bioconversion by solid-substrate fermentation. It also contains descriptions of novel solid-state bioreactors with on-line process control.

50. Viniegra-Gonzalez G, Favela-Torres E, Aguilar CN,

Romero-Gomez S, de J, Diaz-Godinez G, Augar C: Advantages of fungal enzyme production in solid state over liquid fermentation systems. Biochem Eng J 2003, 13:157-167. This paper attempts to explain why enzyme production in solid-state fermentation is higher than in SmF. The authors showed that solid-state culture seems to be working as a natural fed-batch culture with fast oxygenation but slow sugar supply.

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