Brazilian Journal of Microbiology (2011) 42: 1119-1127

ISSN 1517-8382

PRODUCTION OF CELLULASE BY ASPERGILLUS NIGER UNDER SUBMERGED AND SOLID STATE

FERMENTATION USING COIR WASTE AS A SUBSTRATE
Soma Mrudula*, Rangasamy Murugammal
Department of Microbiology, M.G.R. College, Dr. M.G.R. Nagar, Hosur, T.N - 635 109, India.
Submitted: February 28, 2010; Returned to authors for corrections: November 17, 2010; Approved: March 14, 2011.

ABSTRACT
Aspergillus niger was used for cellulase production in submerged (SmF) and solid state fermentation (SSF).
The maximum production of cellulase was obtained after 72 h of incubation in SSF and 96 h in Smf. The
CMCase and FPase activities recorded in SSF were 8.89 and 3.56 U per g of dry mycelial bran (DBM),
respectively. Where as in Smf the CMase & FPase activities were found to be 3.29 and 2.3 U per ml culture
broth, respectively. The productivity of extracellular cellulase in SSF was 14.6 fold higher than in SmF. The
physical and nutritional parameters of fermentation like pH, temperature, substrate, carbon and nitrogen
sources were optimized. The optimal conditions for maximum biosynthesis of cellulase by A. niger were
shown to be at pH 6, temperature 30 C. The additives like lactose, peptone and coir waste as substrate
increased the productivity both in SmF and SSF. The moisture ratio of 1:2 (w/v) was observed for optimum
production of cellulase in SSF.
Key words: Aspergillus niger, coir waste, cellulase, submerged fermentation, solid-state fermentation.
INTRODUCTION

cleave cellobiose and other soluble oligosaccharides to

glucose (10). These enzymes find potential applications in the

Complete cellulose hydrolysis to glucose demands the

action of exoglucanases, endoglucanases and -glucosidases.
Exoglucanases

(1,4- -D-glucancellobiohydrolase,

production of food, animal feed, textile, fuel, chemical,

pharmaceutical industries and in waste management (9, 40).

EC

The production of cellulase has been reported from a wide

3.2.1.91) are usually active on crystalline cellulose and cleave

variety of bacteria (22) and fungi (4, 34). However,

disaccharide units either from non-reducing or reducing end.

filamentous fungi are preferred for commercial enzyme

Endoglucanases

EC

production, because the level of the enzymes produced by these

3.2.1.4) are more active against the amorphous regions of

cultures is higher than those obtained from yeast and bacteria

cellulose and they can also hydrolyze substituted celluloses,

(8). Almost all fungi of genus Aspergillus synthesize cellulase,

such as carboxymethyl cellulose (CMC) and hydroxyethyl

therefore this genus has the potential to dominate the enzyme

cellulose (HEC) internally.

industry. Aspergillus and Trichoderma spp. are well known

(1,4- -D-glucan-4-glucanohydrolase,

-Glucosidases (EC 3.2.1.21)

*Corresponding Author. Mailing address: Department of Microbiology, M.G.R. College, Dr. M.G.R. Nagar, Hosur, T.N - 635 109, India.; Tel.: + 91 (04344)
261004 Fax: +91 (04344) 260573.; E-mail: somamrudula@hotmail.com

1119

Mrudula, S. et al.

efficient production of cellulases (33).

Industrially important enzymes have traditionally been

Production of cellulase by A. niger

subcultured and maintained on Czapek-Dox-agar slants and

stored at 4 C in a refrigerator, until needed.

obtained from submerged fermentation (SmF) because of the

ease of handling and greater control of environmental factors

Pre-treatment of substrates

such as temperature and pH. However, solid state fermentation

The procured cellulosic substrates such as rice husk, rice

(SSF) technique can improve the yield and reduces the cost of

bran, coir waste, wheat bran and saw dust were ground to fine

enzyme production (16, 21). Filamentous fungi are the most

powder and the substrates were individually treated with 1%

commonly used microorganisms in SSF because they are able

(w/v) NaOH solution in the ratio of 1:10 (substrate: solution)

to grow on solid materials with low water contents (30). There

for 1h and was brought to neutral pH by washing thoroughly

are several reports describing use of agro industrial residues for

with distilled water and dried at room temperature. The treated

the production of cellulose such as wheat straw, wheat bran and

substrates were autoclaved at 121C for 1 h (18).

rice straw as substrates (11, 35, 2, 19). The other advantages of

SSF include superior productivity, simple technique; low

Preparation of inoculum

capital investment, low energy requirement and less water

The inoculum was prepared by growing the organism in

output (42, 38), better product recovery and lack of foam build

250 ml Erlenmeyer flask with 100 ml of Czapek-Dox broth

up and reported to be most appropriate process for developing

containing g/l of sucrose, 30; sodium nitrate, 3; K2HPO4, 1;

countries (39).

MgSO4, 0.5; KCl, 0.5; FeSO4, trace; agar, 15. The medium

The cellulase production by filamentous fungi in SmF and

was inoculated from the Czapek-Dox agar slants and

SSF has been studied extensively (15, 1, 38). However, there is

incubated at 30 C for 3 days in a shaker (200 rpm) before it

no report on comparison of the cellulase production by SmF

was used for the fermentation process.

and SSF conditions. From this point of view, the organism was
isolated and demonstrated for their improved efficiency in SmF

Submerged fermentation (SmF)

and SSF for the production of cellulase using agro-industrial

Submerged fermentation was carried out in 250 ml

waste as raw material. The influence of various parameters was

Erlenmeyer flasks containing 100 ml of fermentation medium.

evaluated under two different fermentation conditions. In the

The composition of the medium contained the following g/l of

present study, we describe the comparison of cellulase

distilled water. L-Glutamic acid, 0.3; NH4NO2, 1.4; K2HPO4,

production by Aspergillus niger in SmF and SSF systems.

2.0; CaCl2, 2.0; MgSO4, 0.3; protease peptone, 7.5; FeSO4,

5.0; MnSO4, 1.6; ZnSO4, 1.4; tween 80, 20 % (v/v) ; coir

MATERIALS AND METHODS

waste, 30. The medium was sterilized by autoclaving at 121C

for 15 min. Each flask was inoculated with 1ml of the above

Microorganism
The fungus was isolated from coir retting ground area of

said inoculum. The cultures were incubated on a rotary shaker

(120 rpm) at 30C for 72 h.

Karimangalam, Dharmapuri Dist, Tamil Nadu, India. The

isolate was grown on CMC agar medium and screened

Solid state fermentation (SSF)

according to Wood and Bhat (41). The culture was identified

Solid state fermentation was carried out in 250 ml

based on colony morphology and microscopic examination

Erlenmeyer flasks that contained 10 g of coir waste and 15 ml

(Devenathan et al. 2007). The isolated fungal colony was

of distilled water (moistening agent). The flasks were sterilized

1120

Mrudula, S. et al.

at 121C for 15 min and cooled to room temperature. About

1ml of inoculum was added, mixed well and incubated at 30C

Production of cellulase by A. niger

Effect of supplements (carbon & nitrogen sources)

Smf:

Various

carbon

and

nitrogen

sources

at

in a humidified incubator for 96 h. The flasks were periodically

concentration of 5% w/v were supplemented as individual

mixed by gentle shaking.

components to the fermentation medium containing coir waste

as substrate. The medium was inoculated and incubated at 30

Enzyme extraction

C for 3 days in an orbital shaker incubator (120 rpm). The

At the end of the fermentation the culture broth from

broth was centrifuged and the enzyme assay was carried out.

submerged fermentation was centrifuged at 6000 rpm for 15

Care was taken to see that monosaccharides and disaccharides

min and the supernatant was used as a source of extracellular

were prepared as 10 X solution and sterilized separately by

enzyme. In solid state fermentation (SSF) the enzyme was

autoclaving at 10 lbs for 10 min. and required concentration

extracted from the coir waste by mixing homogenously the

was added to the medium before inoculation.

entire waste with (1:10 w/v) distilled water and agitated on a

SSF:

Various

carbon

and

nitrogen

sources

at

rotary shaker (120 rpm) at 30 C with a contact time of 1h.

concentration of 4% w/v were supplemented as individual

Dampened cheese cloth was used to filter the extract and

components to the flasks that contain 10 g of coir waste and

pooled extracts were centrifuged at 6000 rpm for 15min and

moistened with 15 ml of distilled water (moistening agent).

the clear supernatant was used as a source of extracellular

The flasks were sterilized and cooled to room temperature.

enzyme.

About 1 ml of inoculum was added, mixed well and incubated

at 30C in a humidifying incubator for 4 days. At the end of

Enzyme assay
Cellulase [(filter paperase (FPase) and carboxymethyl
cellulase (CMCase)] activities were assayed according to the
method described by Ghose (17).One unit of enzyme activity
(CMCase and FPase) is defined as the amount of enzyme

fermentation the contents from each flask were extracted with

distilled water and the supernatant was used for enzyme assay.
The effect of moisture content in SSF was studied by
varying the ratio of coir waste to distilled water (2.5 to 12.5).
The

optimum

conditions

for

enhanced

cellulase

which releases 1 mole of reducing sugars per min with

production were verified by performing the experiments using

glucose as standard, under the assay condition described above.

the selected parameters at their respective optimum conditions.

The values of enzymatic activity were expressed as U/ml for

SmF and U/g of dry mycelial bran (DMB) for SSF.

Comparative evaluation of SmF and SSF systems for

enzyme production

Optimization of process parameters in SmF and SSF

Evaluation of optimized culture conditions in SSF and
SmF using Aspergillus niger: The protocol adopted for the

The enzyme yield produced in SmF was compared with

the yield obtained in SSF using coir waste as substrate
according to Solis Pereira et al (36).

standardization of fermentation parameters was to evaluate the

effect of an individual parameter. The parameters optimized

RESULTS AND DISCUSSION

were: substrates (rice husk, rice bran, coir waste, wheat bran,
saw dust), temperature (20 to 40 C), pH (4.5 to 8),
fermentation period [(24 to 192 h in SmF) and (24 to 120 h in
SSF)].

Isolation and screening of cellulolytic organism

The soil sample collected from coir retting zone area was
serially diluted in saline and was spread on to the

1121

Mrudula, S. et al.

Production of cellulase by A. niger

carboxymethyl cellulose (CMC) agar plates. The plates were

systems. In a study conducted by Ojumu et al (29) on

incubated at 30C for 3-5 days. Cellulase producing fungal

Aspergillus flavus reported that the saw dust, corncobs and

colonies were selected after flooding the plates with congo red

bagasse were found to be the best substrates for the production

(0.1% w/v), followed by destaining with NaCl (0.1 M). The

of cellulase. Oberoi et al (28) reported kinnow pulp as the best

colony that showed largest halo forming zone was selected for

substrate for the production of cellulase in SSF. Sugarcane

the present study.

bagasse, tea production waste, coconut coir pith, rice husk,

wheat bran, rice bran etc., have been employed for production

Identification of fungal isolates

Colony morphology: The selected strain was subcultured
on CMC agar medium. Initially the colonies were white and

of cellulase using a variety of microorganisms such as

Trichoderma, Aspergillus, Penicillium, Botrytis, Neurospora
etc., (31).

changed to black, as culture matured. When immature the

colonies were covered with white fluffy aerial mycelia, while

Temperature

mature colonies showed salt and pepper effect which was

Incubation temperature plays an important role in the

covered with black spores and reverse of the colony was buff

metabolic activities of a microorganism. In the present study

colored.

the optimum temperature for maximum enzyme production

was recorded at 30C under SmF and SSF (Table 1). About 83

Microscopic examination of the isolates

% of cellulase production was observed at 35C. Whereas more

The matured colonies were subjected to lacto phenol

than 50 % of cellulase was produced when grown at 20, 25 and

cotton blue staining and microscopic examination was made.

40C in SmF. 55 and 68% of cellulase production was recorded

From the microscopic observation it was clear that the colony

when the organism was grown at 40 and 20C, respectively in

showed the hyaline septate hyphae. The conidial head was

SSF. The results of SmF process in the present study confirm

large and appeared black to brownish black. The conidiophores

the findings of Devanathan et al (12) and Kathiresan and

were hyaline or brownish

near the vesicle. Each vesicle

Manivannan (23) for Aspergillus sp. Asquieri and Park (6)

appeared globose in shape and cover with brownish sterigmata

found that the optimum temperature for production of CMCase

on the entire surface in two series. Based on the colony

from thermostable Aspergillus sp. was 37C, whereas the

morphology and microscopic observation the strain was

maximum cellulase production was observed at 40C for

confirmed as Aspergillus niger (12).

Aspergillus terreus QTC 828 (3). In general the temperature

maintained in SSF system is in the range of 25 to 35C and

Optimization of fermentation parameters in SmF and SSF

depends on the growth kinetics of the microorganism employed

Substrates: Among the 5 substrates screened, coir waste

rather than on the enzyme produced (25). Ali et al. (3) reported

gave the maximum cellulase production when fermented with

maximum yield of cellulase from Aspergillus niger Z10 strain

Aspergillus niger under SmF and SSF (Table 1). Considerable

and A. terreus at 40C, respectively in SSF.

amount of enzyme production was observed on wheat bran and

rice bran. Comparatively less enzyme production was observed

with the rest of the substrates in both SmF and SSF. Since coir

Among physical parameters, pH of the growth medium

waste showed maximum production of cellulase, it was

plays an important role by inducing morphological changes in

selected for further optimization studies for SmF and SSF

microbes and in enzyme secretion. The pH change observed

1122

Mrudula, S. et al.

Production of cellulase by A. niger

during the growth of microbes also affects product stability in

maximum at 96 h of incubation when grown in SmF. These

the medium (20). Optimum pH for maximum production of

results are in agreement with the reports made by Devanathan

cellulase was 6.0 when grown in SmF and SSF (Table 1).

et al (12) and Acharya et al (1). However, further increase in

Similar observation was made for cellulase production by A.

the incubation time reduced the enzyme production. It might be

terreus QTC 828 in SmF by Ali et al (3) and Trichoderma

due to the depletion of macro and micronutrients in the

reesei in SSF by Doppelbauer et al (13), whereas pH 7 was

fermentation medium with the lapse in time, which stressed the

reported by Krishna (24) for the production of bacterial

fungal physiology resulting in the inactivation of secreting

cellulases by using banana wastes in SSF.

machinery of the enzymes (27). Incubation time necessary for

optimal production was observed at 72 h in SSF. Short

Fermentation period

incubation period for enzyme production offers the potential for

Table 1 shows the effect of fermentation period on the

inexpensive production of enzyme (37). Muniswaran and

production of the cellulase in SmF and SSF. In the present

Charyulu (26) have also reported similar trend in cellulase

study cellulase activity increased steadily and reached

production using Trichoderma viride.

Table 1. Effect of physical parameters on cellulase production in SmF and SSF by A. niger

Parameter
Substrates
Rice husk
Rice bran
Coir waste
Wheat bran
Saw dust
Temperature (OC)
20
25
30
35
37
40
pH
4.5
5.0
5.5
6.0
6.5
7.0
7.5
8.0
Fermentation period (h)
24
48
72
96
120
144
168
192

SmF (U/ml)
FPase
CMCase

SSF (U/g DMB)

CMCase
FPase

0.31
0.51
0.8
0.72
0.18

0.25
0.22
0.51
0.43
0.12

0.49
1.1
3.42
1.77
1.2

0.47
0.5
1.77
0.8
0.52

1.5
1.5
3.4
3.1
1.9
1.7

0.8
1.3
1.7
1.3
1.2
1.1

3
3.6
4.44
3.57
3.51
2.5

1.51
2
2.51
2.45
1.46
1.3

0.6
0.81
1.5
2
1.81
1.71
1.3
0.9

0.3
0.61
0.82
0.91
0.82
0.81
1.26
0.5

8.2
9.2
9.6
9.8
9
7.3
6.8
6.6

3.3
4.1
4.3
4.6
2.9
2.5
2.2
2.1

2.46
2.51
3
3.46
2.2
1.5
0.52
0.5

1.37
1.46
1.53
0.88
1
0.51
0.49
0.47

4.5
5.3
9
7.1
6.9

2.4
2.5
3.6
2.3
2.1

1123

Mrudula, S. et al.

Production of cellulase by A. niger

Moisture content

cellulase

production

with

an

increase

in

inoculum

Moisture content is a critical factor in SSF processes

concentration from 5 to 15 % and found to be maximum at

because this variable has influence on growth, biosynthesis and

15%. Higher or lower inoculum size resulted in a significant

secretion of enzyme. In the present study the maximum

decrease in enzyme production (Fig.2). Similar findings have

enzyme production was obtained at coir waste to distilled water

been reported for the production of bacterial cellulases by solid

ratio of 1:2. (Fig.1). Any further increase and decrease in the

state bioprocessing of banana wastes (24). In contrast 10% of

ratio resulted in decreased enzyme activity.

inoculum was reported to be optimum for the production of

Similar

observation was made by Anto et al (5) for production of -

cellulase by Aspergillus niger under SSF (15).

amylase by Bacillus cereus, whereas Babu and Satyanaryana

(7) reported maximum

-amylase enzyme production by

thermophilic Bacillus coagulans at a high level of substrate

moisture ratio of 1: 2.5. According to Lonsane et al (25), lower
moisture content causes reduction in solubility of nutrients of
the substrate, low degree of swelling and high water tension.
On the other hand, higher moisture levels can cause a reduction
in enzyme yield due to steric hindrance of the growth of the
producer strain by reduction in porosity (interparticle spaces)
of the solid matrix, thus interfering oxygen transfer.

Figure 2. Effect of inoculum size on cellulase production by

A. niger in solid state fermentation with coir waste as substrate

Carbon sources
Cellulase production was found to be dependent upon the
nature of the carbon source used in the culture media. To
evaluate the carbohydrates to cause induction or repression of
cellulase, the organism was grown on monosaccharide and
disaccharides. In general, enhanced production of enzyme was
Figure 1. Effect of moistening agent on cellulase production

observed with all the tested sugars when compared to the

by A. niger in solid state

control. Among the carbon sources examined, lactose was

fermentation with coir waste as

substrate

found to be the best inducer in SmF and SSF (Table 2). This
study substantiates the work of Kathiresan and Manivannan
(23) and Devanathan (12) who demonstrated lactose as best

Inoculum size
In the present study there is a significant increase in

inducer of Aspergillus sp.

Nochure et al (27) identified

fructose as the best inducer of cellulase in Clostridium

1124

Mrudula, S. et al.

Production of cellulase by A. niger

thermocellum. In another study dextrin was found to enhance the

nitrogen sources enhanced cellulase production when compared to

production of cellulase by Trichoderma sp. in SSF (32). Increase

control. Among them peptone supported maximum enzyme

in enzyme production with additional carbon sources have been

production followed by beef extract, groundnut oilcake, yeast

demonstrated by both the SmF and SSF systems as a result of

extract and casein. These results are in agreement with the reports

good growth (36).

of Kathiresan and Manivannan (23) and Devanathan et al. (12) for

production of cellulase by Penicillium fellutanum and Aspergillus

Nitrogen sources

niger, respectively in SmF. Enari et al. (14) reported that good

The effect of supplementation of nitrogen sources in SmF and

SSF system on the enzyme production is shown in Table 2. All the

cellulase production can be obtained with peptone as the organic

nitrogen source in SSF.

Table 2. Effect of carbon and nitrogen sources on cellulase production in SmF and SSF by A. niger
Supplement
Carbon sources
(5 % w/v in SmF and 4 %
w/w in SSF)
Control
Glucose
Xylose
Lactose
Maltose
Sucrose
Nitrogen sources
(5 % w/v in SmF and 4 %
w/w in SSF)
Control
Yeast extract
Beef extract
Peptone
Groundnut oil
Casein
Sodium nitrate

SmF (U/ml)
CMCase

SSF (U/g DMB)

FPase

CMCase

FPase

0.7
2.52
2.2
3
2.51
2.54

0.4
0.54
1.42
1.71
1.5
1.51

3.7
12.1
15.7
18
17.5
13.7

2
6.5
6.6
10.9
6.3
6.2

0.56
1.1
1.53
2.1
1.61
0.53

0.4
0.6
0.9
1.36
1.1
0.47

3.8
8.2
11
13.8
8.1
4.2
4

2
4.2
4.5
6.2
4.3
3.8
3.6

Enzyme yields

dividing the yield obtained from SSF in U/g DMB with the

Under the optimum conditions, the strain produced 2.04

yield from SmF in U/ml culture broth. With this method,

units of cellulase per ml of culture broth (Fig. 3) in SmF and

cellulase production by A. niger in SSF and SmF using coir

29.11units of cellulase per gram of dry mycelial bran in SSF.

waste as substrate, were compared in terms of their

extracellular enzyme production in U/g DMB and U/ml,

Comparative evaluation of SmF and SSF system for

respectively (Fig. 3). When comparison made between SmF

enzyme titres

and SSF, production of total cellulase by SSF was 14.6 fold

The method adopted for comparison of submerged and

higher than that of SmF.

solid state fermentation is the same as reported by Solis-Pereira

In conclusion the result of the present study clearly

et al. (36) for pectinase production from Aspergillus niger by

indicates the potential of Aspergillus niger can be successfully

1125

Mrudula, S. et al.

Production of cellulase by A. niger

Bacillus cereus MTCC 1305 using solid state fermentation. Food Tech.

cultivated under SSF conditions for the production of cellulase

using coir waste as substrate. The coir waste is abundantly
available in India as it is one of the major producers of
coconut. Having considered the means of reducing disposal

Biotechnol., 44 (2), 241-245.

6.

Asquieri, E.R.; Park, Y.K. (1992). Production of extracellular cellulases

7.

Babu, K.; Sathyanarayana, T. (1995).

from the thermostable Aspergillus sp. Rev. Microbiol., 23, 183-188.

-Amylase production by

problem, therefore it can be effectively utilized by a potential

thermophilic Bacillus coagulans in solid state fermentation. Process

strain like Aspergillus niger for production of cellulase which

Biochem., 30 (4), 305-309.

is commercially important.

8.

Bakri, Y.P.; Jacques, P.; Thonart. (2003). Xylanase production by

Penicillum canescens 10-10c in solid state fermentation. Appl. Biochem.
Biotechnol., 108 (1-3), 737-748.

9.

Beguin, P. (1990). Molecular biology of cellulose degradation. Ann. Rev.

Microbiol., 44 (2), 19-248.

10.

Bhat, M.K.; Bhat, S. (1997). Cellulase degrading enzymes and their

potential industrial applications. Biotechnol. Adv., 15 (3-4), 583-620.

11.

Biswas, S.R.; Jana, S.C.; Mishra, A.K.; Nanda, G. (1990). Production,

purification and

characterization of xylanase from a hyperxylanotic

mutant of Aspergillus ochraceus. Biotechnol. Bioeng., 35, 244-251.

12.

Devanathan, A.; Shanmugan, T.; Balasubramanian.; Manivannan, S.

(2007). Cellulase production by Aspergillus niger isolated from coastal
mangrove debris. Trends. Appl. Sci. Res., 2, 23-27.

13.

Doppelbauer, R.; Esterbauer, H.; Steiner, W.; Lafferty, R.M.;

Steinmuller, H. (1987). The use of lignocellulosic wastes for production
of cellulase by Trichoderma reesei. Appl. Environ. Microbiol., 26 (5),
485-494.

Figure 3. comparative evaluation of cellullase production in

SmF and SSF using Aspergillus niger.

14.

Enari, T.M.; Markenan, P.; Fiechter, A. (1977).

Production of

15.

Fadel, M. (2000). Production physiology of cellulases and -glucosidase

cellulolytic enzymes by fungi. Adv. Biochem. Eng., 5, 1-24.

enzymes of Aspergillus niger grown under solid state fermentation
conditions. Online Biol. Sci., 1, 401-411.
16.

REFERENCES
1.

Murthy, V. (1985). Economics of submerged and solid state fermentation

Acharya, P.B.; Acharya, D.K.; Modi, H.A. (2008). Optimization for

for the production of amyloglucosidase. J. Food Sci.Technol., 22, 171-

cellulase production by Aspergillus niger using saw dust as substrate.

Afr. J. Biotechnol., 7 (22), 4147-4152.
2.

176.
17.

Alegre, A.C.P.; Polizeli, M. L.T.M..; Terenzi, H.F.; Jorge, J.A.;

Guimaraes, L.H.S. (2009).Production of thermostable invertases by

Comparative studies on production of cell wall-degrading hydrolases by

using agroindustrial residues as carbon source. Braz. J. Microbiol., 40,

Trichoderma reesei and T. viride in submerged and solid state

cultivations. Bang. J. Microbiol., 23 (2), 149-155.

Ali, S.; Sayed, A.; Sarker, R.I.; Alam, R. (1991). Factors affecting

19.

Anita, S.; Namita, S.; Narsi, R.; Bishnoi.

Goyal, M.; Kalra, K.L.; Sareen, V.K.; Soni, G.

(2008). Xylanase

production with xylan rich lignocellulasic waste by a local soil isolate of

J. Microbiol. Biotechnol., 7 (1), 62-66.

5.

Gomes, I.; Mohammad, S.; Sabita, R. R.; Donald J. G. (2006).

Aspergillus caespitosus under submerged or solid state fermentation

cellulase production by Aspergillus terreus using water hyacinth. World

4.

Ghose, T.K. (1987). Measurement of cellulase activities. Pure.

Appl.Chem., 59 (2), 257-268.

18.

612-622.
3.

16. Ghildyal, N.P.; Lonsane, B.K.; Sreekantiah, K.R.; Sreenivasa

Trichoderma viride. Braz. J. Microbiol. 39, 535-541.

(2009). Production of

20.

Gupta, R.; Gigras, P.; Mohapatra, H.; Goswami, V.K.; Chauhan, B.

cellulases by Aspergillus heteromorphus from wheat straw under

(2003). Microbial -amylases. A Biotechnological Perspective. Process

submerged fermentation. Int. J. Environ. Sci. Eng., 1, 23-26.

Biochem., 38 (11),1599-1616.

Anto, H.; Ujjval, T.; Kamlesh, P. (2006).

amylase production by

21.

Hui,L.;Wan, C.; Hai-tao, D.; Xue-jiao, C.; Qi-fa, Z.; Yu-hua, Z. (2010).

1126

Mrudula, S. et al.

22.

Production of cellulase by A. niger

Direct microbial conversion of wheat straw into lipid by a cellulolytic

Production of cellulolytic enzyme by a newly isolated, Trichoderma sp.

fungus of Aspergillus oryzae A-4 in solid-state fermentation.

FETL C 3-2 via solid state fermentation grown on sugar cane baggase:

Bioresour.Technol 101, 7556-7562.

Palm kernel cake as substrates. Pak. J. Biol. Sci., 9, 1430-1437.

Immanual, G.; Dhanusha, R.; Prema, P.; Palavesan. (2006). Effect of

33.

The transcriptional activator X in R regulates both xylanolytic

bacteria isolated from coir retting effluents of estuarine environment.

endoglucanase gene expression in Aspergillus niger. Appl. Environ.

Int. J. Environ. Sci. Tech., 3 (1), 25-34.

23.

24.
25.

Microbiol., 64, 3615-3617.

Kathiresan, K.; Manivannan, S. (2006). Cellulase production by

34.

acremonium for the production of cellulose and xylanase through

Res. J. Microbiol., 1 (5), 438-442.

submerged fermentation. Afr. J. Microbiol. Res., 4 (14), 1506-1510.

Krishna, C. (1999). Production of bacterial cellulases by solid state

35.

(2010). Advancement and comparative profiles in the production

Lonsane, B.K.; Ghidyal, N.P.; Budiatman, S.; Ramakrishna, S.V.

technologies using solid-state and submerged fermentation for microbial

Muniswaran,

P.K.A.;

cellulases. Enzym. Microb. Tech., 46, 541-549.

36.

Charyulu,

N.C.L.N.

(1994).

Solid

state

Aspergillus niger in submerged and solid state fermentation.

Microb. Technol., 16 (5), 436-446.

Nochure, S.V.; Roberts, M.F.; Demain, A.L. (1993). True cellulase

Trichoderma reesei (RUT-C30) on municipal solid waste.

Biotech. Lett., 15 (6), 641-646.

Biochem. Biotechnol., 51-52(1),145-153.

substrate. Food Bioprocess. Technol., 3 (4), 528-536.

38.

39.

Szendefy, J.; Szakacs, G.; Christopher, L. (2006). Potential of solid-state

fermentation enzymes of Aspergillus oryzae in biobleaching of paper

(2003). Cellulase production by Aspergillus flavus linn isolate NSPR 101

pulp. Enzym. Microb. Tech., 39, 1354-1360.

40.

Tarek, A.A.M.; (2007). Optimization of cellulase and

-glucosidase

induction by sugarbeet pathogen Sclerotium rolfsii. Afr. J. Biotechnol., 6

Pandey, A. (1992). Recent developments in solid state fermentation.

Process Biochem., 27 (2), 109-117.

(8), 1048-1054.
41.

Pandey, A.; Selvakumar, P.; Soccol, C.R.; Nigam, P. (1999). Solid state
fermentation for the production of industrial enzymes. Cur. Sci., 77 (1),

32.

Souza, P. M.; Magalhaes, P.O. (2010). Application of microbial

amylase in industry-A review. Braz. J. Microbiol. 41, 850-861.

150-152.

31.

Appl.

Ojumu, T.V.; Solomon, B.O.; Betiku, E.; Layokun, S.K.; Amigun, B.

fermented in saw dust, bagasse and corncob. Afr. J. Biotechnol., 2 (6),

30.

Sonjoy, S.B.; Bex, K.; Honston, H. (1995). Cellulase activity of

production by C.thermocellum grown on different carbon sources.

Oberoi, H.S.; Yogita, C.; Sunil B.; Gurpreet, S. D. (2008). Production of

Appl.

Microbiol. Technol., 39 (1), 36-41.

37.

cellulose through solid state fermentation using kinnow pulp as a major

29.

Solis-Pereira, S.; Favela-Torres, E.; Viniegra-Gonzalez, G.; (1993).

Effects of different carbon sources on the synthesis of pectinase by

fermentation of coconut coir pith for cellulase production. Enzyme

28.

Singhania, R.R.; Sukumaran, R.K.; Patel, A.K.; Larroche, C.; Pandey, A.

bioprocessing of banana wastes. Bioresour. Technol., 69 (3), 231-239.

Technol., 7 (6), 258-265.

27.

Sarao, L.K.; Arora, M.; Sehgal, V.K. (2010). Use of scopulariopsis

Penicillium fellutanum isolated from coastal mangrove rhizosphere soil.

(1985). Engineering aspects of solid state fermentation. Enzyme Microb.

26.

Peij, N.; Gielkens, M.M.C.; Verles, R.P.; Visser, K.; Graff, L.H. (1998).

different growth parameters on endoglucanase enzyme activity by

Wood, T.M.; Bhat, K.M. (1988). Methods for measuring cellulose

activities. Methods Enzymol., 160, 87-112.

42.

Zeng, W.; Chen, H.Z. (2009). Air pressure pulsation solid state

149-162.

fermentation

Pang P.K.; Darah I.; Laszlo P.; George S.; Ibrahim C. O. (2006).

Bioresour.Technol., 100, 1371-1375.

of

feruloyl

esterase

by

Aspergillus

niger.

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