IJPCBS 2013, 3(4), 1070-1090 Sharada et al. ISSN: 2249-9504

INTERNATIONAL JOURNAL OF PHARMACEUTICAL, CHEMICAL AND BIOLOGICAL SCIENCES

Available online at www.ijpcbs.com Review Article

PRODUCTION OF CELLULASE – A REVIEW

R. Sharada1 , G. Venkateswarlu2, S. Venkateshwar3 and M. Anand Rao4 1Government Junior College (G), Sangareddy, Dist.Medak, Andhra Pradesh, India. 2Bharat Heavy Electronics Limited, R.C.Puram,Hyderabad, Andhra Pradesh, India. 3University College of Technology, O.U,Hyderabad, Andhra Pradesh, India. 4University College of Science, O.U, Hyderabad, Andhra Pradesh, India.

ABSTRACT Cellulases are the enzymes which hydrolysis cellulosic biomass and are being produced by the microorganisms grown over cellulosic matters. Cellulase is an important enzyme which can be obtained from cheap agrowastes,as well as cellulose as substrates by using submerged fermentation and solid state fermentation. Cellulose protein can be degraded by cellulase enzyme produced by cellulolytic bacteria and fungi. This enzyme has various unique industrial applications and it has been considered as major group of industrial enzyme. In this present review paper, discussion attempted on “cellulase production” by submerged and solid state fermentation using various types of bacteria and fungi with different types of agro wastes.

INTRODUCTION cellulosic biomass (Berry et al., 1990), and an Enzymes are among the most important economic process for its production is thought products obtained for human needs through to be critical for the successful utilization of microbial sources. A large number of industrial cellulosic materials (Soloman et al., 1999, Wu processes in the areas of industrial, and Lee, 1997, Nwodo-Chinedu, S. et al., 2007). environmental and food biotechnology utilize The cellulase complex used in simultaneous enzymes at some stage or other. Current saccharification and fermentation systems developments in biotechnology are yielding new (SSFS) generally includes C1 [(EC 3.2.1.9), (exo- applications for enzymes (Ashok Pandey et al., 1, 4-β-D- glucanase, cotton lyase), Cx (EC 1999). 3.2.1.4), (endo- 1, 4 - β-D- glucanase, In the present techno- economic era, procurance carboxymethyl cellulase, or CMC ase), Cb (EC of energy is one of the major problems which 3.2.1.21), (β- glucosidase)], and pectinase. humanity is facing. All the waste cellulose is a (Debing Jing et al., 2007). source of food and is also a potential source of Cellulases have been used and studied for most energy (Elder et al., 1986). of the 20th century and are the most Cellulose present in renewable lignocellulosic commercially important of all the enzyme material is considered to be the most abundant families. The enzyme activities were increased organic substrate on earth as chemical feed about 30-80% when produced by SSF in stock (Krishna, 1999). comparison with conventional SmF enzyme Cellulose is a branched glucose polymer production. Cost of cellulase production may be composed of an -1,4 glucose units linked by a – brought down by multifaceted approaches 1, 4- D- glycosidic bond (Gielkens et al., 1999; which include the use of cheap lignocellulosic Han et al., 1995, Acharya. P.B et al., 2008). The substrates and the use cost efficient breakdown of cellulose into sugar can be fermentation strategies like solid state achieved by acid hydrolysis as well as by fermentation (Rajeev et al., 2008) The enzyme enzymatic hydrolysis. But enzymatic hydrolysis production is good by most of the fungi like is mostly preferred because it produces fewer Aspergillus and Trichoderma Sp. by-products and proceeds under milder Enzymolysis of native cellulose is carried out by condition reported by Mandels et al., (1974). three components of cellulase as: Cellulase, a group of enzymes which catalyze the a. Exo- β-1-4, glucanase: It acts on the non- hydrolysis of cellulose and related reducing end of the cellulose chain and oligosaccharide derivatives, is considered a successively removes single glucose units. potential tool for industrial saccharification if

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b. Endo-β-1-4, glucanase: It randomly attacks cellobiase, which eventually breaks down the internal β-1-4, linkages. cellobiose, the building unit of cellulose, to 3. β-glucosidases or Cellobiases: In addition to glucose. C1 and Cx, the cellulose system also contains

Enzyme filtrate Cellulase C1

C1+Cx Cellulase Cellobiase Native

Native Cellulose especially crystalline Cx region made responsive to C1 probably by

disaggregation of Micelles by braking H bonds

Exo-β-1-4-Glucanase Removes successively single glucose units from non-reducing end of cellulose chain

Random attack especially on internal β-1-4 Endo- β-1-4-Glucanase Linkages

Cellobiase Cellobiose

Glucose

Fig. Enzymolysis of native Cellulose

Microbial Sources of Cellulases and biosynthetic regulation of cellulases in Cellulase is one of the most useful enzymes in C.hutchinsonii were demonstrated on different industry. Cellulase can be produced by fungi, substrates (Clifford Louime et al., 2006) bacteria or actinomycetes, but the most Acetivibrrio cellulolytieus has been studied by common producer is fungi (Arriffin. H., et al., many workers investigated that both endo- 2006) glucanase and exo-glucanase can be regulated by induction and catabolite repression. It differs Bacteria from most of the cellulolytic organisms in its The bacterial systems have also been ability to utilize only cellulose, cellobiose or investigated for saccharification of the biomass salicin for growth (Saddler et al., 1980). This and might have advantage because of the fast organism has attracted special interest because growth rate of bacteria. It has also been of high specific activities for endo- and exo - reported that the enzyme preparation from glucanase (Mackenzie et al., 1985).The cellulolytic bacteria can effectively saccharify endocellulase activity of the culture broth was different cellulosic substrates (Choundry et al., determined during growth of Acinetobacter 1980, 1981; Bynd et al., 1987; Rajoka et al., anitratus and Branhamella Species by 1984; Waldron et al., 1986). measuring release of reducing sugars from CMC Cytophaga hutchinsonii revealed an unusual (Ekperigin.M.M., et al., 2006). The enzyme collection of genes for an organism that can preparations from A.cellulolyticus have the attack crystalline cellulose. Location, formation ability to saccharify cellulose (Mackenzie et al.,

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1985); the condition for maintaining high Effect of some nutritional and environmental specific activity have also been determined. factors on production of cellulases was studied Cellulase system of Pseudomonas Sp. has been using Bacillus licheniformis MVS1 and Bacillus fairly well studied. Yamane et al., (1970); Sp. MNS3 isolated from an Indian hot spring Yoshikawa et al., (1974) studied the biogenesis (Somen Acharya et al., 2012). Deepmoni Deka et of multiple cellulose components of P. al., 2011 attempted to optimize the medium fluorescens var. cellulose with special emphasis components for enhanced production from on effects of culture conditions on the Bacillus subtilis AS3 and significant variables for multiplicity of cellulose titres produced by the enhancing alkaline cellulase production were organism. Tewari et al., (1977) reported screened and selected using the Plackett – production, purification and properties of Burman design. cellulase (extracellular) from Pseudomonas Sp. Clostridium thermocellum, a thermophilic Localization of cellulase components in anaerobic bacterium, has attracted increased Pseudomanas Sp., isolated from activated sludge interest for conversion of LC biomass. Being have been investigated (Ramasamy et al.,1980). thermophilic, it offers several advantages over Endo-glucanase are the major components of mesophilic organisms because; the former cellulase complexes with three being from produces enzymes with increased P.fluorescens var. cellulose (Yashikawa et al., thermostability and significantly higher specific 1974) and four or more by Pseudomomas Sp. activity (Johnson et al., 1982; Ng and Zeikus, (Ramasamy et al., 1980). Gene for D-galactose 1981, 1988). Moreover, the cellulases produced dehydrogenase has been cloned in E.coli (Buckle by Clostridia are not inhibited by moderate et al., 1988). Pseudomonas fluorescens, Bacillus concentrations of glucose or cellobiose. The subtilis, E.coli and Serratia marcescens, cellulase organism produces exo and endo- glucanase producing bacreia was isolated from soil and having multiforms (Bisaria et al., 1981) which optimization of the fermentation medium for act together during saccharification. The maximum cellulase production was studied and organism forms high molecular weight found Pseudomonas fluorescens as best cellulase aggregated (Ait et al., 1979; Lamed et.al., 1983). producer (Sonia Sethi et al., 2013). Somen Acharya et al., 2012 isolated a The cellulolytic enzymes of Bacillus Species have thermophilic bacterium Aneurini bacillus been the focus of many due to their potential use thermoaerophilus WBS2 which produces in the conversion of agricultural wastes into extracellular thermophilic cellulases from hot useful products (Ozaki et al., 1990). It was spring in India in order to enhance cellulase reported that Bacillus subtilis CBTK 106 can production various fermentation parameters produce a considerable amount of cellulase was also optimized. Thermophilic bacteria activity (Chundakkadu Krishna, 1999). Femi- strain Geobacillus pallidus was successfully Ola. T.O et al., 2008 studied that aqueous extract isolated from empty fruit bunch (EFB) and palm of the woods inhibited the growth and oil mill effluent compost and characterized by production of cellulase in the strains of B.subtilis Azhari Samsu Baharuddin et al., 2010. significantly. Muhammad Salem Akhtarif et al., Rumen cellulolytic bacteria have been studied 2001 studied the usage of B.subtilis in extensively (Varel, 1984; Ohmiya et al., 1985). saccharification of wheat straw, rice straw and Cellulase produced by Ruminococcus albus can bagasse. Production of cellulase using carboxy hydrolyse CMC and acid swollen cellulose methyl cellulose as substrate by B.pumilus EB3 causing a rapid fall in degree of polymerization was studied by H.Ariffin et al., 2006.Saraswati while affecting only a small degree of hydrolysis Bai et al., 2012 studied production of cellulose (Wood et al., 1984). Groleu et al., 1981 have using Bacillus subtilis CEL PTK 1 from cow dung. shown that Bacterium Succinogens, another Mohammed S.A.Shabab et al., 2010 studied anaerobic rumen cellulolytic bacterium cellulose productivity by Bacillus subtilis KO producing high CMC-ase and cellobiase activity. strain using CMC zone and dinitro salicyclic acid.

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Fungi Deswal D. et al., 2011 in solid state fermentation These organisms grow under suitable condition using brown rot fungus, Fomitopsis sp. RCK 2010. to produce cellulase; e.g. Trichoderma viride and Eggin et al., (1968); Malik et al., (1970) studied Trichoderma reesei produce active cellulase the cellulase production by certain fungi and when grown in solid submerged culture found that microorganisms have the ability to Aspergillus niger produces highly active cellulase degrade native cellulose. Since the first when grown in liquid media by both surface and observation of the action of conversion of submerged culture methods and recently by cellulosic biomass to fermentable sugar needs solid state fermentation (Ikram-ul-Haq et al., economical process for the production of 2005). The detailed study was made on cellulases. Soil is inhibitated by a large number production of cellulase using Trichoderma reesei of microorganisms like bacteria, actinomycetes, (Muthuvelayudham et al., 2003). Production of algae and fungi; which form a major component cellulase also reported by using substrates like of soil. Although these microorganisms form a cellulose, xylose and lactose using T.reesei very small part of the soil, they are responsible (Muthuvelayudham et al., 2005). Saravanan et for many of the chemical transformations and al.,2013 studied production of cellulose using even for some of the physical changes that take Trichoderma reesei in solid state place in soil. Fungi especially make a unique fermentation.Some species of Penicillium i.e. contribution in these changes which result in Penicillium iriensis and P. citriviride produce the proper maintenance of soil fertility. significant quantities of cellulase, when grown The first attempt to isolate fungi from soil was under different conditions. Penicillium made by Adametz (1866) after which various funiculosum is capable of dissolving cotton investigators studied the soil fungus flora by completely by cellulase production. Fungi like various methods. Stachy, Botsysatra, Pesalotia, Palmarum, Among these investigators the names of Lender Merulius, Lnerymans, Polyspores, Neuraspora (1908), Hagem (1910), Bachwith (1911), Jensen have been reported to produce cellulase (1912), Goddard (1913), Dale (1914), (Mandels et al., 1976; Shoemaker et al., 1978; Werkenthin(1916), Paine (1927), Gilman et Metha et al., 1975; Kassim, 1983; Takao et al., al.,(1932), Wajid (1985) are more prominent. 1985; Brown et al., 1987; Hyashida et al., 1980; But the work of Waksman (1922, 1927) gave the Adikanae et al., 1983).Chaetomium sp.NIOCC 36 soil dilution plate methods to isolate soil fungi, was found better for production of which is still being extensively used. Both cellulase(Chinnarjan Ravindran et al., 2010). qualitative and quantitative studies were made Barros R R. et al., 2010 study evaluated by these investigators. They found the data on cellulases, xylanases and beta – glucosidases kind of soil, soil reaction (pH), depth of soil, produced by two fungi, the thermotolerant moisture, and season of year, tillage and Acrophialophora nainiana and Ceratocystis manuring practices. They also found that the parodoxa using submerged fermentation. result of isolation of fungi is also influenced by Production of cellulase was also reported by

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methods of investigation, nature of medium and al.,1986), A.aculeatus (Murao et al., 1988), material employed. A.fumigatus (Heptinstall et al., 1986), Aspergillus The work has also been carried out on the heteromorphus (Anitha Singh et al., 2009, Rajesh isolation and identification of Trichoderma Singh et al., 2009), Alternaria alternate (Bailey et Species from soil sample which determine the al., 1988; Macris, 1984), A.japonicus (Sharma et antagonistic properties of the mold cultures al., 1985; Sanyal et al., 1988), A.niger (Gokhale et (Dennis, 1970; Dennis et al., 1971; Rifai, 1964, al., 1984; MeCleary et al., 1988; Okada,1989; 1969) Poutanen et al., 1984; Jun-ichi-Abe et al.,1999; Among Trichoderma Species, T. harzianum Milala. M.A et al., 2005; Narasimha,G., et al., (Deschamp et al., 1985; Macris et al., 1985; Mes- 2006; Sharada,R et al.,2012), A.ustus (Macris et Hartree et al., 1988; Saddler et al., 1982; Saddler al., 1985), A.wentii (Panda et al., 1987; et al., 1987; Md.Zahinger Alam et al., 2005; Srivastava et al., 1984), A.flavus (Ojumu et al., Ikram-ul-Haq et al.,2006, Shazia Shafique et al., 2003), Aspergillus candidus (Milala,M.A., et al., 2009), T.aureoviride (Mercedes Zaldivar et al., 2009),Tricothecium roseum (Prashanth 2001), T.reesei (Muthuvelayudham,R., et al., shanmugam et al., 2008),Scopulariopsis 2006,2007; Maryam Latifian et al., 2007; (Bharathi Kodali et al., 2006),Penicillium Wang.J.Sh et al., 2006, Shazia Shafique et al., chrysogenum (Nwodo-chinedu et al., 2009), T.viride (Benkun Qi et al., 2007, Shazia 2007),A.oryzae (Adebace Johnson Adeleke et al., Shafique et al., 2009) and T.konningii (Wood, 2012),A.flavus NRRL5521 (Hussein Azzaz Murad 1988; Wood et al.,1982, 1986) has been studied. et al., 2013). Other well studied fungi are A.terreus (Araujo et

Table 2: Cellulase producing fungal strains Fungal strain Reference Pesalotia Metha et al., 1975 Stachy Mandels et al., 1976 Botsysatra Shoemaker et al., 1978 Polyspores Hyashida et al.,1980 Palmarum Kassim, 1983 Neuraspora Adikanae et al.,1983 A.ustus Macris et al., 1985 Merulius Takao et al., 1985 A.fumigatus Heptinstall et al., 1986 A. heteromorphus Anitha Singh et al., 2009, Rajesh Singh et al., 2009. A.terreus Araujo et al.,1986 Lnerymans Brown et al., 1987 A.aculeatus Murao et al., 1988 T.aureoviride Mercedes Zaldivar et al., 2001 A.flavus Ojumu et al., 2003 Scopulariopsis Bharathi Kodali et al., 2006 P. chrysogenum Nwodo-chinedu et al., 2007 T.viride Benkun Qi et al., 2007, Shazia Shafique et al., 2009. Tricothecium roseum Prashanth shanmugam et al., 2008 A.wentii Panda et al., 1987; Srivastava et al., 1984 Deschamp et al., 1985; Macris et al., 1985; Mes-Hartree et al., 1988; T. harzianum Saddler et al., 1982; Saddler et al., 1987; Md.Zahinger Alam et al., 2005; Ikram-ul-Haq et al., 2006, Shazia Shafique et al., 2009. Alternaria alternate Bailey et al., 1988; Macris, 1984 A.japonicus Sharma et al., 1985; Sanyal et al., 1988 Gokhale et al., 1984; MeClearyet et al., 1988; Okada,1989; A.niger Poutanenet et al., 1984; Jun-ichi-Abe et al.,1999; Milala,M.A., et al., 2005; Narasimha.G et al., 2006; Sharada,R et al., 2012. A. candidus Milala,M.A., et al., 2009 T.konningii Wood, 1988; Wood et al., 1982, 1986. Muthuvelayudham,R., et al., 2006,2007; Maryam Latifian et al., 2007; T.reesei Wang.J.Sh et al., 2006, Shazia Shafique et al., 2009, Saravanan et al.,2013 Acrophialophora nainiana and Barros R R. et al., 2010 Ceratocystis parodoxa Chaetomium sp.NIOCC 36 Chinnarjan Ravindran et al., 2010 Fomitopsis sp. RCK 2010 Deswal D. et al., 2011 A.oryzae (Adebace Johnson Adeleke et al., 2012) A.flavus NRRL5521 Hussein Azzaz Murad et al., 2013

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Production of cellulase Agricultural and industrial wastes are among 1995 when a mutant of Trichoderma reesei was the main causes of environmental pollution. co-cultured with a strain of Pleurotus sajor caju Their conversion into useful products may with wheat straw as the substrate. reduce the intensity of the problems caused by Fungi was isolated and identified from soil and them. These wastes include green gram husk, decomposing orange peels. Out of thirteen black gram husk, rice bran, wheat bran etc. are isolates, three highest producers of enzymes underutilized in India especially in Andhra were isolated and identified as Penicillium Pradesh. In most parts of A.P these materials are atrovenetum, Aspergillus flavus and Aspergillus mainly used as animal feeds. A large quantity is oryzae, these isolates were used to ferment left in farm lands to be decomposed by orange peels in solid state fermentation. microorganisms such as bacteria and fungi Maximum production was given by A.oryzae (Okafor et al., 1987).Economically, the most (Adebace Johnson Adeleke et al., 2012). important industrial material other than food Stoilova. I.S et al., 2005 studied the enzyme stuffs affected by microorganisms are cellulose production of two novel mixed cultures of and wood products (Wainright, 1992; Debing mycelia fungi Thermoascus aurantiacus and Jing et al., 2007). Proper utilization of these Aspergillus niger in solid state fermentation. wastes in the environment will eliminate Enzymatic hydrolysis of palm oil effluent solid pollution and convert them into useful by- using mixture of locally isolated fungi products (Milala et al., 2005). Aspergillus niger EB5 and Trichoderma sp.EB6 Cellulose is commonly degraded by an enzyme was reported by Wong Kok Mun et al., called Cellulase. This enzyme is produced by 2008.They also observed effect of substrate several microorganisms, commonly by bacteria pretreatment, different ratio of cellulase mixture and fungi (Bahkali et al., 1996; Shin et al., 2000; and incubation pH on the enzymatic hydrolysis Immanuel et al., 2006).Filamentous fungi are of POME solids. Ikram-ul-Haq et al., 2005 preferred for commercially important enzymes reported cotton saccharifying activity of production, because the level of the enzymes cellulases obtained from mono and co-culture produced by these cultures is higher than those fermentation of A.niger and Trichoderma viride. obtained from bacteria (Bakri et al., 2003). Four carbon sources i.e. CMC, cellulose powder, Fungal genera like Aspergillus and wheat bran and rice bran were used as Trichoderma are taught to be cellulase substrates, among them cellulose powder was producers and crude enzymes produced by having highest cotton degrading ability (963 these microorganisms are commercially U/h/L) after 72 hrs, compared to wheat bran available for agricultural use (Peig et al., (657 U/h/L). 1998).The microorganism which appear to be Local isolated fungal cultures including A.niger, most promising at present are Trichoderma Fusarium oxysporum, Fusarium avenaceum and mutant (Jecu et al., 2000), Shazia Shafique et al., Cephalosporium acremonium were employed for 2009 studied evaluation of cellulase production cellulase production. Wheat straw was used as potential by Trichoderma strains, in particular carbon source and Aspergillus niger was chosen T.viride, T.reesei and T.harzianum available in on the basis of best mean cellulase activity First Fungal Culture Bank of Pakistan (FCBP). (H.H.Azzaz et al., 2011), Hussein Azzaz Murad et However, the cellulase systems of al., 2013 studied production of celulase under hypercellulase producing mutants of optimum conditions using Aspergillus flavus Trichoderma reesei are deficient in β-glucosidase NRRl 5221 utilizing agricultural waste as carbon (Duff et al., 1986).One of the problems related to source. the economic viability of the enzymatic Omojasola.P.F et al., 2008 studied cellulase hydrolysis of cellulose is due to low β - production from cellulosic pineapple waste glucosidase levels (Umikalsom et al., using Trichoderma longibrachiatum, Aspergillus 1997).Unlike Trichoderma Species Aspergillus niger and Saccharomyces cerevisiae. The amount Species exhibited high β -glucosidase activity of glucose produced was optimized by varying (Wong Kok Mun et al., 2008). the fermentation parameters: time, pH, There are several reports describing co- substrate concentration, inoculum size and culturing of two cultures for enhanced cellulase temperature. Among three cultures, production. Gupte,A., et al., 1997 cultivated two Trichoderma longibrachiatum found to highest strains of Aspergillus ellipticus and A.fumigatus producer of glucose (0.92 mg/0.5 ml) at pH 4.5 and reported improved hydrolytic and β- and temperature of 45 ºC on day 7 of glucosidase activities compared to when they fermentation. were used separately using SSF, improved Optimization of the media components for enzyme titres were achieved by Kanotra,S., et al., cellulase production using Trichoderma reesei

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was carried out. Optimization of cellulase Trichoderma reesei and Aspergillus niger using production using pine apple waste as substrate black gram husk and green gram husk and was performed with statistical methodology reported maximum cellulose production using (P.Saravanan et al., 2013). green gram husk as substrate in solid state Omajasola. P.F et al., 2008 studied production of fermentation by Aspergillus niger. cellulase from the orange peel using Ikram-ul-Haq el al., 2006 described the Trichoderma longibrachiatum, Aspergillus niger production of cellulolytic enzymes (CMC-ase, and Saccharomyces cerevisiae after treatment FP-ase, β-glucosidase) and hemicellulolytic with alkali and steam. The activity of the test enzyme (xylanase) along with total extracellular organisms cellulase against CMC on the orange protein by Aspergillus niger and Trichoderma wastes was also reported. T.longibrachiatum viride using submerged fermentation. (3.86 mg ml-1), A.niger (2.94 mg ml-1) and Debing Jing et al., 2007 studied optimization of S.cerevisiae (2.30 mg ml-1) glucose amounts cellulase inoculent mixture, to improve woody from orange pulp. lignocellulosic hydrolysis using pea shrub Muhannad I. Massadeh et al., 2001 used woody biomass with cellulase inoculent mixture sugarcane bagasse as substrate with consisting of Trichoderma koningii, Aspergillus Trichoderma reesei QM9414 and Aspergillus niger, Lactobacillus and observed that terreus SUK-1 to produce cellulase and reducing Lactobacillus inoculent resulted in lactic acid sugars. The percentage of substrate degradation inhibitory effect on further improvement of achieved employing mixed culture was 26% cellulase hydrolysis and crude protein content compared to 50% using separate cultures of two in SSF fermentation. moulds. Bokhary, H.A et al., 1994 reported that, Haque et al., (1989, 1990, 1991) have reported Aspergillus was a predominant genus, among 61 the production of cellulase by Aspergillus and fungal species isolated as cellulose degraders, Penicillium by solid substrate fermentation from rhizosphere soil mycoflora of Alfalfa, Date using different cellulosic biomass. Of all the palm and Grape wine. The production of substrates however, enzymes production by all cellulase was examined by cleared zone the cultures were maximum with wheat bran. technique. The parameters such as depth of wheat bran, Ajayi. A.A et al., 2007 described the effects of partial replacement of wheat bran by different some chemicals and cations on the activity of agricultural by products selection of diluents partially purified cellulase from tomato fruits and extraction of the enzymes from the deteriorated by Aspergillus flavus Linn. The fermented substrate, were optimized. enzyme was partially purified by a combination Rajesh Singh et al., 2009 reported cellulase of ammonium sulphate precipitation, molecular production using rice straw as substrate by exclusion chromatography and ion exchange fungal strain Aspergillus heteromorphus. The chromatography. They observed that uninfected RSM methodology was used for optimization of tomato fruits did not possess detectable cellulase production. cellulase whereas tomato fruits infected with Immanuel, G et al., 2007 studied the cellulase Aspergillus flavus have appreciable quantity of enzyme production ability of fungal strains cellulase activity. The production of cellulase by Aspergillus niger and Aspergillus fumigatus A.niger on three different carbon sources was against the bio wastes, coir waste and saw dust compared by Gautam. S.P. et al., 2010 as carbohydrate source at varying Umar Dahot. M et al., 1996 made tests to utilize environmental parameters of pH and wheat straw as a carbon and energy source for temperature. Partial purification of the cellulase the growth of Aspergillus fumigatus and enzyme was reported using Poly Acrylamide Gel production of cellulases. They observed that the Electrophoresis (PAGE) showed that A.niger maximum production of β- glucosidase and CM- grown in coir waste supplemented media had cellulase was achieved by Aspergillus fumigatus two protein bands with the molecular weight of grown on H2SO4 and HCl pretreated wheat straw 36 and 23 kDa respectively and A.fumigatus substrate in comparison to HNO3 and HClO4 grown in the same medium had two protein pretreated wheat straw. bands with molecular weight of 32 and 21 kDa Aspergillus flavus can be grown on different respectively. substrates such as bagasse, corncob and saw Cellulase enzyme production was studied by dust, but saw dust pretreated with caustic soda Charitha Devi.M et al., 2012 using fungal strain gave the best result with an enzyme activity Aspergillus niger against the lignocellulosic bio value of 0.0743 IU/ml as reported by Ojumu et wastes like sawdust, paper cellulose at varying al., 2003. Acharya, P.B., et al., 2008 studied environmental parameters. Sharada.R et al., factors relevant for the improvement of 2012 studied production of cellulase using enzymatic hydrolysis of saw dust pretreated

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with NaOH using Aspergillus niger. Cellulase straw, rice husks and maize straw as suitable activity obtained was around 0.0925 IU/ml at substrates for the production of cellulase by 120 rpm after 96 hrs incubation period. Aspergillus niger. Maize straw moistened with Fade, M., 2000 studied the production diluents containing mineral salts plus basal salt physiology of endoglucanase, exoglucanase and solution gave the highest cellulase activity of β-glucosidase on radicle waste medium by 102 (IU/ml) at about 72 hrs of fermentation. Aspergillus niger F-119 under condition of SSF. Narasimha, G., et al., 2005 reported effects of Cellulase production from radicle waste was nutrient on cellulase production by Aspergillus markedly affected by different parameters such niger in submerged fermentation. Cellulase as moisture level, incubation period, initial pH, production by Aspergillus niger on three media incubation temperature and inoculum size. (minimal, basal and Czapek-Dox) in liquid shake Rita Pye et al., 2003 studied conditions of the were compared. Czapek-Dox medium was found biosynthesis of pectinolytes, cellulolytes and to be superior for the growth and cellulase hemicellulases by the filamentous fungi production by A.niger. They also studied effects Aspergillus niger IBT-90 by a mathematical of different carbon and nitrogen sources to activity of factorial planning and gradient Czapek-Dox medium on cellulase production. On optimization. They found that the process defined media carboxymethyl cellulose and optimization led to a three fold increase in the NaNO3 was found to best carbon and nitrogen activity of pectinolytic enzymes and the double sources respectively. the activity of cellulolytic enzymes and xylanase. Paul, L., et al., 1976 studied mechanism of action King.K.W et al., 1963 studied distinctive of cellulase and purification, physico-chemical properties like ultraviolet spectra, pH-activity and enzymatic properties of 1,4-β-glucan responses, substrate specificities, thermal glucanohydrolase (3.2.1.4) from A.niger. They stabilities, and kinetic changes in the viscosity of observed that the enzyme was rich in acidic and substrate, adsorption characteristics on aromatic amino acids and kinetic studies gave cellulose and exclusion characteristics on pK values between 4.2 and 5.3 for groups dextran gels of β-glucosidases and related involved in the enzyme-substrate complex. The enzymes from Aspergillus niger. Effect of pH on molecular weight of the enzyme calculated was hydrolysis of carboxymethyl cellulose and 26,000, and cellulase was found to be stable to cellohexaose was compared and suggested that heat treatment at pH 8.0. The mode of action a negative charge center on the substrate has and substrate specificity of a cellulase purified pronounced inhibitory effect on the enzymes. from A.niger was reported by Paul. L. Hurst et Gokhan Coral et al., 2002 studied some al., 1977. The specificity region of cellulase was properties such as molecular weight, optimum five glucose units in length, indicating cellulase pH, temperature and heat stability of is an endoglucanase. carboxymethyl cellulase of new mutant strain of Gretty K. Villena et al., 2006 studied the Aspergillus niger Z10 a cellulase producer. The production of lignocellulolytic enzymes crude enzyme preparation was subjected SDS- (cellulase, endoglucanase and xylanase) of PAGE to determine the homogeneity and A.niger in submerged fermentation (SmF) and molecular weight of the enzyme. During the biofilm fermentation (BF) cultures. Maximal electrophoresis two bands were observed filter paper activity, endoglucanase and xylanase showing cellulolytic activity, the molecular activities was higher in BF (2.96, 4.7, and 4.61 weights of these proteins was calculated to be IU/ml respectively) but biomass yields was 83,000 and 50,000.These enzymes were not higher in SmF (0.431 g g-1). Sharada.R et al.,2009 having ability to hydrolyse insoluble studied optimization of cultural conditions for microcrystalline cellulose (Avicel) but active cellulase production by Aspergillus niger (MTCC towards carboxy methyl cellulose. 2196) using submerged fermentation. Jun-ichi Abe et al., 1999 reported purification and characterization of endo-β-1,4-glucanase Solid state fermentation and β-glucosidase from cellulolytic fungus The term solid state fermentation (SSF) denotes Aspergillus sp. K-27 by ion exchange cultivation of microorganisms on solid, moist chromatography and affinity chromatography. substrates in the absence of free aqueous phase. The molecular weight of endo-β-1,4-glucanase The possible advantages of the biotechnological was calculated to be 21,000; the molecular uses of SSF relative to those of classical weight of β-glucosidase was calculated to be submerged fermentation (SmF), different 1,30,000 and 1,05,000 suggesting that the β- aspects of the two, include: a) historical, b) glucosidase was a hetero dimer enzyme. biological, c) ecological, d) engineering and e) 1`Milala, M.A., et al., 2005 evaluated cellulosic economical differences. agricultural wastes such as millet, guinea corn

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A. Historical aspects E. Economic aspects When compared with the SSF processes that Calculation of the costs of production of have long been established in Asian countries, cellulase, the economical efficiency is higher by SSF in Western countries appears to still be in a factor of 100 than in the case of SmF its infancy. SSF is the state of the art technology (Tengerdy,R.P et al., 2003). This striking that is used in many applications in the food difference has several reasons: much cheaper industry in Asia, i.e. SSF is used in the growth substrates, minimized requirements for production of enzyme rich Koji (which is made sterility and low requirements for of rice and uses Aspergillus sp. as an enzymatic instrumentation and equipment. starter for different hydrolytic processes), in the SSF hence offers advantages over fermentation saccharification of rice used for the production in liquid broth (SmF) like higher product yield, of alcoholic beverages such as sake, in the better product quality, cheaper product production of cellulase, which uses Trichoderma recovery and cheaper technology (Oguntimein and Aspergillus strains. In Asia, enzymes and et al., 1992). Microbial degradation of the metabolites are commonly produced on a large biological wastes is a natural process that has scale by SSF processes that may be thousands of occurred since the on set on earth. In years old. Today, SSF has been nearly fermentation processes microorganisms utilize completely abandoned because of the pressure the wastes as potential energy source for of increasing industrial rationalization and synthesis of very useful products such as standardization. enzymes (Kishwar Hayat et al., 2001). Solid state fermentation processes are distinct B. Biological aspects from submerged fermentation (SmF) culturing, SSF processes simulate the living conditions of since microbial growth and product formation many higher filamentous fungi: Ascomycetes, occurs at or near the surface of the solid basidiomycetes and deuteromycetes developed substrate particle having low moisture contents. in terrestrial habitats on wet substrates. Studies Solid state fermentation (SSF) holds tremendous of the production of fungal enzymes in SSF have potential for the production of enzymes. It can shown that SSF, in comparision with SmF, be of special interest in those processes where provides higher volumetric productivities, is the crude fermented products may be used less prone to problems with substrate inhibition directly as enzyme sources. and yields enzymes with a higher temperature Tengerdy, R. P., et al., 1996 compared cellulase or pH stability. Also, the fermentation time may production in SmF and SSF techniques. While be shorter and the degradation of the enzymes the production cost in crude fermentation by by undesirable proteases is minimized SmF was about $ 20/kg, by SSF it was only $ (Holker.U et al., 2004). 0.2/kg. Nigam, P., et al., 1994 have reviewed processing of agricultural wastes in SSF systems C. Ecological aspects for cellulolytic enzyme production. They also Ecological advantages of SSF reflect the fact that enumerated advantages of cellulase production the processes are conducted in the absence of a together with the factors affecting the cellulase free aqueous phase. Because the SSF processes production in SSF systems. are performed at water activities below 1, the Elisashvili, V.I., et al., 1998 studied on the growth of contaminating bacteria and yeasts is lignolytic system of Cerrena unicolor 062 a minimized. Which means that, in certain cases, higher basidiomycete upon supplementation of energy and instrumentation demanding the medium with carbon sources and phenolic sterilization processes can be eliminated. compounds in SSF system; it was observed that Another additional environmentally friendly the growth of C.unicolor 062 could be regulated feature of SSF is, in many cases it can use by the exogenous addition of these compounds. agricultural wastes as carbon and energy The efficiencies of the degradation of cellulose sources. The production of enzymes or organic and lignin were dependent on the nature and acids makes frequent use of plant remnants as concentration of the compounds added. carbon sources and inducers or mediators. Solid state fermentation of palm kernel cake has been carried out by Mohd Firdaus Othman et al., D. Engineering aspects 2013 using locally isolated Rhizopus oryzae ME The most serious reasons why SSF has not yet 01. found a broad use in western countries are Sun, T., et al., 1997 developed a novel fed batch engineering problems, the low amenability of SSF process for cellulase production which the processes to standardization and the limited could overcome the problems associated with reproducibility of the results. high initial nutrients concentration while

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retaining advantages from the high total Mandels and Reese (1957) produced cellulose effective salt concentration. by growing Trichoderma viride and observed In significant finding, Smits, J.P., et al., 1996 that it was influenced by carbon source and reported that glucosamine level of the fungi in metals. Cellulase was an adaptive enzyme in liquid culture could not be used to estimate the cultures of T.viride. It was produced on cellulose, biomass contents in SSF. They studied the SSF of lactose, glucose and cellobiose, but not on a wheat bran by T.reesei and reported that using wide variety of other substrates. Trace elements glucosamine, correlation between the fungal such as iron, manganese and zinc or cobalt were growth and respiration kinetics could only best for cellulose production. partly be described with the linear growth Trichoderma viride can be grown on different model of Pirt. substrates such as avicel, rice straw, filter paper, Shi – hao Zhao et al., 2011 reported production wheat straw, but wheat bran moistened with of cellulose in solid state fermentation using dilutents containing mineral salts plus CMC gave Trichoderma reesei SEMCC – 3.217 strain with the best yield as reported by Haq et al., (1989, water hyacinth. Fractional factorial design 1991a, 1991b). showed that, the addition amount of wheat bran, Dissolving cellulose pulp, newspaper cellulose, ammonium sulphate, calcium chloride and avicel and cellophane are more reactive towards Tween 80 had significant effect on cellulase hydrolysis with cellulose of T.viride. Cellophane, production. coated on one side with nitrocellulose, would be Ikram ul Haq et al., 2006 studied the hydrolysed to glucose in high yields in a exploitation of agricultural by products like reasonable time. Toymao et al., (1978) reported wheat bran, wheat straw, rice bran, rice straw that in case of solid cultures, stronger enzyme and soybean for the production of industrially activities were produced than were produced in important enzyme Cellulases by SSF using the submerged culture. Chand et al., (1990) locally isolated Trichoderma harzianum. Of all evaluate the agro-residues and grass as carbon the substrates wheat bran was found to be best source for cellulose production. Agro cultural substrate for production of cellulase. Kishwar wastes were used as carbon source for the Hayat et al., 2001 reported kinetics of cellulase production of cellulose. These wastes were and Xylanase production by culturing subjected to NaOH treatment and fermentation Chaetomium thermophile on wheat straw by SSF. parameters with T.reesei were detected. They observed that the enzyme activity was Properly treated cellulosic waste promises to be affected by fermentation period, and found a reasonable substrate for cellulase production maximum at 72 hrs. by T.reesei. Sathyavarathan .P. et al., 2013 studied cellulase Tappa Mohammad Munawar et al., 2011 production using Trichoderma reesei NCIM – investigated cellulase production in solid state 1052 and Aspergillus niger NRRL – 322. production using T.reesei NCIM 1186 by Maximum cellulase activity reported was 217.17 Antigonum leptopus leaves as substrate gave U ml -1 using T.reesei. maximum cellulosic activity. Jian Liu et al., 2007 studied cellulase production Cellulase was produced from corn straw by solid in SSF using the waste from the vinegar industry state fermentation with the help of T.reesei. The as the substrate for Trichoderma koningii AS3 corn straw was supplemented with wheat bran 4262. The effects of water content, initial pH for the supply of nitrogen and carbon, by Wang, value in solid substrate and culture temperature J., et al., 2006. They also studied the optimal on cellulase synthesis was observed for optimal conditions like temperature, pH, and water production in flask fermentors. contents of substrate. Cellulase production Benkun Qi et al., 2007 reported production of studies was carried out using fungal strain cellulases from T.viride using SSF by different Trichoderma reesei NCIM 992 by using three ratios of rice straw and wheat bran as substrate. different lignocellulosic materials by solid state It was observed that activities of Filter paper fermentation and reported maximum enzyme, Endoglucanase and β-glucosidase were production using wheat bran yielding 2.63 U ml - significantly affected by the substrate mixture. 1 during incubation time of 6 days(Devendra p. Henkelekian et al., (1925) have shown that Maraya et al., 2012). Trichoderma and Penicillium possess the ability Muthuvelayudham, R., et al., 2007 used RSM to to decompose cellulose completely, with CO2 as evaluate the effects of the medium parameters the only waste product. They also observed a on cellulase production by using T.reesei Rut direct correlation between the amount of C30 and to evaluate the kinetic model for nitrogen transformed into ammonia by species attaining a higher cellulase yield in SSF. Maryam of Penicillium and Trichoderma and the amount Latifian et al., 2007 used RSM to evaluate the of cellulose decomposed. effects of fermentation parameters for cellulase

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production by T.reesei QM 9414 and T.reesei Shahera, H., et al., 2002 used SSF to evaluate the MCG77 in SSF using rice bran as substrate. possibility of re-use of orange peel and pulp H.H. Azzaz et al., 2012 studied cellulase wastes as a source of enzymes production; production by locally isolated fungal cultures cellulase, amylase, pectinases, lipases, esterases including A.niger, Fusarium oxysporum, and peroxidases. Microorganisms were isolated Avenaceum and Cephalosporium acrenonium from fermented waste and tested for their using wheat straw as a sole of carbon source enzymes production. Maximal production of and reported that the highest cellulase cellulase and lipase was reported by the use of production obtained from A.niger. orange peel waste with yeast strain. Ben Faber et al., 2003 studied differences in the The effects of glucose, crystalline cellulose and production of cellulase from different mulching saw dust of Mitragyna cilata on the growth and materials in a field setting using SSF. Organic cellulase production, inferred from cellulase mulches are found to control root rot of activity of Penicillium chrysogenum PCL501 was avocado, caused by Phytophthora cinnamomi. studied (Nwodo-Chinedu et al., 2007). Saw dust The possible mechanism for this control was is reported as a good inducer of cellulase activity enzyme production specifically cellulase and in the organism. glucanase. There are several reports indicating production Solid state fermentation of lignocellulosic of cellulases in SSF using various substrates material oil palm biomass generated from palm with Aspergillus sp. (Debing Jing et al., 2007; oil industries as waste was used for cellulase Omajasola, P.F., et al., 2008; Acharya, P.B., et al., production through T.harzianum in lab scale by 2008; Milala, M.A., et al., 2005; Fadel, M., 2000; Md.Zahangir Alam et al., 2005. The parameters Umar Dahot, M., et al., 1996, Jahir Alam Khan et glucosamine and reducing sugar was observed al., 2011, Umbrin Ilyas et al., 2011,sharada.R et to evaluate the growth and substrate utilization. al., 2012).

Table 3: Various strains and substrates used for cellulase production Strain Substrate Reference T.koningii Saw dust, wheat bran Arima, K., et al., 1964 T.reesei Wheat bran Mudgett, R.E., et al., 1982 T.reesei QMY-1 Wheat straw Chahal, D.S., et al., 1985 Sporotrichum pulverulentum , Wheat bran Kim, J.H., et al., 1985 T.reesei T.harzianum Wheat straw, Wheat bran Deschamps, F., et al., 1985 Strains of Basidiomycetes Bagasse Nigam, P., et al., 1986 Trichoderma sp. Botritis sp. Wheat bran, rice straw Shamla, T.R., et al., 1987 Polyporus sp. Bagasse Nigam, P., et al., 1987 Neurospora crassa Wheat straw Macris, B.J., et al.,1987 A. heteromorphus Wheat straw Anitha Singh et al., 2009 A.niger Wheat bran Talukdar, S., et al., 1992 Wheat bran, bagasse, rice and wheat T.viride, A.niger Ikram ul Haq et al., 1992 straw, Cerrena unicolor Grape wine trimming dust Zakariasvili, N.G., et al., 1993 Penicillium citrinum Rice husk Kuhad, R.C., et al., 1993 Mesophilic fungal strain Rice husk Begum, A.A., et al., 1993 A.heteromorphus Rice Straw Rajesh Singh et al., 2009 Cerrena,unicolor, Coriolus hircutus, Tea production waste Kokhreidze, N.G., et al., 1993 Pleurotus ostreatus A.niger Coconut coir pith Muniswaran, P., et al., 1994 Aspergillus sp. Bagasse, Wheat bran, rice bran Gupte, A., et al., 1994 Lentinus edodus Wheat straw Giovannozzisermanni, G., et al., 1994 T.reesei, A.niger Sweet sorghum silage Castillo, M.R., et al., 1994 Trichoderma reesei+Aspergillus Bagasse Duenas, R., et al., 1995 phoenicis Phanerochaete chrysosporium Soyhull Jha, K., et al., 1995 T.reesei Paddy straw Kanotra, S., et al., 1995 Gliocladium sp. Trichoderma sp. Sweet sorghum silage Szakaca, G., et al., 1996 Penicillium sp. T.harzianum Cassava waste Onilude, A.A.,et al., 1996 T.reesei Steam pretreated willow Reczey, K., et al., 1996 A.fumigatus Wheat straw Umar Dahot, M., et al., 1996 A.niger Palm oil mill waste Prasertsan, P.,et al., 1997 P.sajor caju Sago hampas Kumaran, S., et al., 1997 Cerrena unicolor Grape wine cutting waste Elisashvili, V.I., et al., 1998 Bacillus subtilis Banana wastes Chundakkadu Krishna 1999

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Radicle waste (malt manufacture A.niger Fadel, M.,2000 residue) T.reesei QM9414, A.terreus SUK-1 Sugarcane bagasse Muhannad I. Mussadeh et al., 2001 A.flaves, Nigrospora sp. Orange waste Shahera H. Attyia et al., 2002 Phytophthora cinnamomi mulch Ben Faber et al., 2003 A.flavus Linn NSPR 101 Saw dust, bagasse and corn cob Ojumu et al., 2003 Bacillus subtilis Banana stalk Shafique, S.,et al., 2004 A.niger KK2 Rice straw, Wheat bran Kang, S.W., et al., 2004 T.harzianum Wheat straw, Wheat bran Deschamps, F., et al., 2004 T.harzianum Oil palm biomass Md.Zahangir Alam et al., 2005 A.niger MSK-7,T.viride MSK-10 Wheat bran Ikram ul Haq et al., 2005 Thermoascus aurantiacus, A.niger Oats straw, Wheat bran Stoilova, I.S., et al., 2005 Millet, guinea corn straw, rice husk A.niger Miala, M.A., et al.,2005 and maize straw Rice husk, millet straw, guinea corn A.candidus Milala, M.A., et al., 2009 stalk and saw dust Wheat bran, sugarcane bagasse, orange bagasse, corn cob, green Thermoascus aureantiacus miche Robutoda Silva et al., 2005 grass, dried grass, saw dust and corn straw A.niger, T.viride Rice bran, wheat bran, cotton Ikram ul Haq et al., 2005 T.reesei Sugarcane bagasse, rice straw Muthuvelayudham,R., et al., 2006 Wheat bran,wheat straw, rice bran, T.harzianum Ikram ul Haq et al., 2006 rice husk and soybean T.reesei LW1 Corn straw Wang.J.Sh et al., 2006 Scopulariopsis Rice bran Bharathi Kodali et al., 2006 A.niger,A.terreus and Rhizopus Cassava waste Pothiraj, C.,et al., 2006 stolonifer Penicillium echinalatum Bagasse, wheat bran Camassola, M., et al., 2007 T.koningiiAS3 4262 Wheat bran, vinegar waste Jian Liu et al., 2007 T.viride ZY-01 Rice straw, wheat bran Benkun Qi et al., 2007 T.reesei QM9414, T.reesei MCG77 Rice bran Maryam Latifian et al., 2007 Humicola insolens TAS-13 Sugarcane bagasse Muhammad Mohsin Javed et al., 2007 Penicillium chrysogenum PCL 501 Saw dust Nwado-Chinedu et al., 2007 T.koningii. A.niger, Lactobacillus Pea shrub biomass Debing Jing et al., 2007 A.niger Eb5, T.sp.EB6 Palm oil mill effluent Wong Kok Mun et al., 2008 T.longibrachiatum, A.niger, Orange waste Omajasola, P.F., et al., 2008 Saccharomyces cerevisae T.longibrachiatum, A.niger, Pine apple waste Omajasola, P.F., et al., 2008 Saccharomyces cerevisae A.niger Saw dust Acharya, P.B., et al., 2008 Penicillium roqueforti Pumpkin oil cake Draginja pericin et al., 2008 A.niger Green gram husk Sharada.R et al., 2012 A.niger Corn cob Jahir Alam Khan et al., 2011

A.niger Vigna mungo Umbrin Ilyas et al., 2011,

T.reesei NCIM 1186 Antigonum leptopus leaves Tappa Mohammad Munawar et al., 2011

T.reesei NCIM 1052 Ground nut Satyavarathanetal.,2013 T.reesei SEMCC Water hyacinth Shi hao Zhao et al., 2011 Rhizopus oryzae ME01 Palm kerne cake Mohd.Firdaus Othman et al., 2013

CONCLUSION examine Aspergillus sp. to improve cellulase cellulases were produced by SmF and SSF using production which is a known good producer of various bacterial and fungal strains. cellulases (Jecu et al., 2000, Sharada.R. et al., Development of an economical process for 2012). cellulase production is hindered because of the Many researches have been conducted on high costs of substrate (pure cellulose) and of enzymatic hydrolysis of various lignocellulolytic some chemicals, such as proteose peptone, and substrates like Pumpkin oil cake, Saw dust, Pine also because of low yields of cellulases per unit apple waste, Orange waste, Palm oil mill of cellulose. To overcome these bottlenecks, effluent, pea shrub biomass, Sugarcane bagasse, cheap source of cellulose; lignocelluloses, Rice bran, Rice straw, wheat bran, vinegar agricultural wastes are used in SSF. waste, Cassava waste, Corn straw, wheat straw, The microorganisms which appear to be most rice husk, soybean, cotton, corn cob, green grass, promising at present are Aspergillus sp. and dried grass, Millet, Oats straw, Oil palm biomass, Trichoderma sp. However, it is of interest to Banana stalk, mulch, Radicle waste (malt

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manufacture residue), Sago hampas, Grape wine production of industrial enzymes. cutting waste, Steam pretreated willow, Sweet Curr. Sc. 77(1): 149-162. sorghum silage, soyhull, Paddy straw, Coconut 10. Azhari Samsu Baharuddin, Razak, coir pith, Tea production waste, Grape wine Mohamad Nafis Abd; Lim Siong Hock; trimming dust, green gram husk utilized Ahmad, Mohd Najib; Abd-Aziz, cellulases from bacterial and fungal strains. Suraini; Rahman, Nor'Aini Abdul; Md Shah, Umi Kalsom; Hassan, Mohd Ali; ACKNOWLEDGEMENTS Sakai, Kenji; Shirai, Yoshihito., 2010. The authors are thankful to research scholars Isolation and Characterization of K.Swathi, J.Swarnalatha, Anamika, Neeraja for Thermophilic Cellulase-Producing their friendly co –operation. Bacteria from Empty Fruit Bunches- Palm Oil Mill Effluent Compost. REFERENCES American Journal of Applied sciences, 1. Acharya,P.B., Acharya,D.K. and 7(1): 56 Modi,H.A., 2008. Optimization for 11. Bachwith,T.B., 1911. Effect of ethyl cellulase production by Aspergillus alcohol and CO2 sporulation of bakers niger using saw dust as substrate. Afr. yeask nature, London, 164, 544. Symp. J. Biotechnol., 7(22): 4147-4152 Proceeding 1st., 387-395. 2. Adametz,F.A., 1866. The soil and 12. Bahkali,A.H., 1996. Influence of microbe. various carbohydrates on xylanase 3. Adikanae,H.V.and Patil,M.B., 1983. production by V.tricorpus. Cellulase production by Fusarium Bioresource Technol. 33(3): 265-268. solani. Indian Bot. Rep., 2(1): 97-98. 13. Bailey,M.J. and Poutaneu,K., 1988. 4. Ait,N., Creuzet,N. and Caltaneo,J., 1979. Production of xylanolytic enzymes by Characterization and purification of strains of Aspergillus. Appl. Microbiol. thermostable β-glucosidase from Biotechnol., 3(1): 20-25. Clostridium thermocellum. Biochem. 14. Bakri,Y.P., Jacques,P., Thonart., 2003. Biophys. Res. Commun., 90: 537-546. Xylanase production by Penicillium 5. Ajayi,A.A., Adejuwon,A.O., canescens 10-10c in solid state Awojobi,O.K. and Olutiola,P.O., 2007. fermentation. Appl. Biochem. Effect of cations and chemicals on the Biotechnol., 108(1-3): 737-748. activity of partially purified cellulase 15. Barros RR, Oliveira RA, Gottschalk LM, from Tomato (Lycopersicon Bon EP, 2010. Production of esculentum mill) fruits deteriorated cellulolytic enzymes by fungi by Aspergillus flavus Linn. Pakistan J. Acrophilophora nainiana and Nutrition, 6(2): 198-200. cetatocystis paradoxa using different 6. Anitha Singh, Namita Singh and Narsi, carbon sources. Appl. Biochem. R. Bishnoi, 2009. Production of Biotechnol., 161 (1-8): 448-454. cellulases by Aspergillus 16. Ben Faber and Michael Spiers., 2003. heteromorphus from wheat straw Cellulase production by various under submerged fermentation. sources of mulch. Proceedings V Proceedings of International World Avocado Congress (Actas V Conference on Energy and Congreso Mundial del Aguacate), 561- Environment, March 19-21. 565. 7. Araugo,A. and Souza,J.D., 1986. 17. Benkun Qi, Risheng Yaoa, Ying Yua, Characterization of cellulolytic Yuan Chena., 2007. Influence of enzyme components from Aspergillus different ratios of rice straw to wheat resseus and its mutant derivatives. J. bran on production of cellulolytic Ferment. Technol., 64: 463-467. enzymes by Trichoderma viride ZY-01 8. Ariffin,H., Abdullah,N., Umi in solid state fermentation. EJEAF Kalson,M.S., Shirai,y. and Hassan,M.A., Che., 6(9): 2341-2349. 2006. Production and characterization 18. Berry,D.R., Paterson,A., 1990. Enzymes of cellulase by Bacillus pumilus EB3. in food industry: In enzyme chemistry, International Journal of Engineering impact and applications. 2nd edn. CJ and Technology, 3(1): 47-53. Suckling (Ed.)., 306-351. 9. Ashok Pandey,Selvakumar,P., 19. Bharathi Kodali and Ravindra Pogaku., Carlos,R.Soccol. and Poonam Nigam., 2006. Pretreatment studies of rice 1999. Solid state fermentation for the bran for the effective production of

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