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This article reviews the varied sources of oligosaccharides available in nature as silent health promoting, integral ingredients of plants as well as animal products like honey and milk. The article focuses on exotic and unfamiliar oligosaccharides like Galactooligosaccharides, Lactulose derived Galactooligosaccharides, Xylooligosaccharides, Arabinooligosaccharides and algae derived Marine oligosaccharides along with the most acknowledged prebiotic fructooligosaccharides. The oligosaccharides are named as on the grounds of the monomeric units forming oligom- ers with functional properties. The chemical structures, natural sources, microbial enzyme mediated synthesis and physiological effects are discussed. An elaborate account of the different types of oligosaccharides with special refer - ence to fructooligosaccharides are presented. Finally, the profound health benefits of oligosaccharides are rigourously discussed limelighting its positive physiological sequel. Keywords: Oligosaccharides, Prebiotics, Functional food and applications Since past three decades there has been constant evalu- Introduction ation of market trend of western countries witnessing Food industry is presently witnessing an upcoming increased demand of functional foods. Even in develop- market for edible products having health benefits apart ing country like India, where the dairy industry is one of from nutrition, now well recognized as functional foods. the main industries supporting economy, there has been The market of functional foods is facing an increas - a significant rise in demand of value added dairy prod - ing demand also because of consumer awareness about ucts encompassing health benefits to the consumers health. According to the Global Industry Analyst (GIA) (Gour 2013). report on the demand of prebiotics, based on studies Prebiotics and probiotics have raised as best option for in market trends in countries like US, Canada, Japan, quench of the increasing need of functional food. Rober- Europe (France, Germany, Italy, UK, Spain, Russia and froid (2000) studied probiotics and prebiotics food and rest of Europe), Asia–Pacific (China, India and Rest of reviewed their properties to be rightly labeled as func- Asia–Pacific) and rest of World, the industry is likely to tional foods. He explained that prebiotics are non-digest- flourish to a tune of US $4.8 billion by 2018 from US $1.0 ible food ingredients that benefit the host by selectively billion in 2011 (Spinner 2013). stimulating the growth or activity of one or limited num- Japan is one of the leading countries giving importance ber of bacteria in colon. to functional food market focusing on “Food of Specified Food ingredients which naturally offer resistance to Health Use” (FOSHU). Many European countries like digestion, when reach the intestine exhibit a favoring Germany, France, United Kingdom and Netherlands have effect on normal flora of the colon are called as prebi - also showed an extended demand for functional foods otics. Prebiotics encompass several health benefits like (Katapodis et al. 2004; Menrad 2003). the calorie-free nature, act as artificial sweeteners, have non-carcinogenic nature and stimulate the growth of Bifidobacterium and probiotic Lactobacilli in the colon *Correspondence: firstname.lastname@example.org Department of Microbiology and Bioinformatics, Bilaspur University, (Saminathan et al. 2011). They possess preventive effect 206, Budhiya complex, Sarkanda, Bilaspur, Chhattisgarh 495004, India against colon cancer (Moore et al. 2003). They have Full list of author information is available at the end of the article © 2016 The Author(s). This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Belorkar and Gupta AMB Expr (2016) 6:82 Page 2 of 11 ability to decrease cholesterol levels in the serum (Fer- have great industrial applications (Crittenden and Playne nandez et al. 2003). Phospholipids and triglyceride levels 1996; Prapulla et al. 2000). The chemical structure of are also found to be regulated in the serum by prebiotic some important oligosaccharides are given in Fig. 1. food (Katapodis et al. 2004). Fructooligosaccharides (FOS) are gaining wide acceptance as prebiotics (Belorkar Structure of fructooligosaccharides et al. 2013). This mini review presents an overview of the FOS consist of a fructose units polymerized to different types of oligosaccharides existing in nature, their sources extent. Oligomers with two fructose units are called as and major thrust applications. 1-kestose. Oligomers with three fructose units are called as 1-nystose. Oligomers with four fructose units are Oligosaccharides: types, sources and applications called as 1-fructofuranosyl-nystose. The sugars are linked Extensive research has been done on various types of oli- by β-2, 1 position of sucrose (Sangeetha et al. 2005). gosaccharides. They differ in their nature of monomeric sugars and are named so. They have varied sources of ori - Occurrence of FOS gin and differ in their benefits imparted to the consumer. Varieties of sources cater fructooligosaccharides in vary- The most popular oligosaccharides are FOS, Galac - ing concentrations as its natural component like wheat, tooligosaccharides (GOS), Lactulose derived galactoo- honey, onion, garlic and banana (Roberfroid and Slavin ligosaccharides (LDGOS), Xylooligosaccharides (XOS), 2000). Barley and tomato contains 0.15 % of fructoo- Arabinooligosaccharides (AOS), algae derived marine ligosaccharides. Banana and brown sugar has 0.30 % oligosaccharides (ADMO). Other oligosaccharides occur- fructooligosaccharides. Honey has 0.75 % of fructooligo- ring in nature are Pectin-derived acidic oligosaccharides saccharides (Flamm et al. 2001). (pAOS), Maltooligosaccharides (MOS), Cyclodextrins Bornet et al. (2002) recorded the occurrence of short (CD) and human milk oligosaccharides (HMO) with chain FOS in many edible plants. Fructooligosaccha- specific acknowledged benefits. The oligosaccharides rides expresses degree of polymerization from 1 to 5 Fig. 1 Overview of structure of some common oligosaccharides Belorkar and Gupta AMB Expr (2016) 6:82 Page 3 of 11 units of fructose. Short chain oligosaccharides are simi- Fructosyltransferase enzyme lar to dietary fibers in resisting digestion in intestine and Some plants and microorganisms express fructosyl- getting converted to acetate, propionate, butyrate and transferase enzyme naturally. The activity of this enzyme gas in colon. Fructooligosaccharides also add up to the empowers these organisms to synthesize fructooligosac- fecal matter and gives improved bowel movement. In charides (Sanchez et al. 2008). Fructosyltransferase enzyme the digestive tract they promote Bifidobacterium and on from different sources exhibit different mechanisms of other hand have an inhibitory effect on Clostridium per - action and produce different mixtures of oligosaccharides. fringes in colon. FOS are found abundantly in nature as a component Beneficial health effects of FOS on consumers of cereals, fruits and vegetables next to starch specified FOS are receiving attention and importance not merely in Fig. 2 (Sangeetha et al. 2005). These exhibit resistance because of their application as alternative sweeteners to basic enzymes involved in digestion like alpha amyl- but rather for the accompanied desirable characteristics. ase, saccharase and maltase when investigated in humans The earlier known health benefits of FOS were inhibi - (Losada and Olleros 2002). tory effect on pathogens and stimulatory effect on Bifi - Johnson et al. (2013) reported that lentils are rich in dobacterium. The FOS was analyzed further to highlight prebiotics. There is a significant variation in prebiotic its detailed interaction with Bifidobacterium (probiotics) carbohydrate composition of different types of lentils. which paved a pathway for the concept of synbiotics (Per- They analyzed Raffinose-family oligosaccharides, sugar rin et al. 2001; Vander et al. 2004). The health benefits of alcohols, fructooligo-saccharide and resistant starch FOS have been reviewed by many workers (Antosova and carbohydrates. They recorded the occurrence of Raffi - Polakovic 2001; Hernandez et al. 2009; Patel and Goyal nose-family oligosaccharides, sugar alcohols, fructooli- 2011; Ganaje et al. 2014). gosaccharides and resistant starch as 4071, 1423, 62 and Some of the evident health benefits observed by con - 7500 mg per 100 g dry matter, respectively. sumption of fructooligosaccharides include the following: Fig. 2 Distribution of FOS in various natural products Belorkar and Gupta AMB Expr (2016) 6:82 Page 4 of 11 Promotes growth of the gut micro flora Promotes preferential growth of Bifidus Studies on Bifidobacterium species revealed that fruc - A statistical model was used by Shuhaimi et al. (2009) for tooligosaccharides preferred those carbohydrates which the study of growth of Bifidobacterium pseudocatenu - allow maximum growth and metabolic activities of this latum G4 under the influence of prebiotic. The physi - beneficial flora in human intestine (Palframan et al. ological effect of inulin and fructooligosaccharides were 2003). The diet and its composition have an impact on investigated with sorbitol, arabinan and inoculum rate. gut and its microflora. It has been observed that any Fractional factorial design was used to determine their kind of change in the diet affects the metabolism of the effect on growth of selected bacterium in skimmed milk. inhabitants. The dietary fibres like oligosaccharides exert They optimized their growth conditions and concluded a combined effect on both the pH environment of the gut that in 1 L fermentor, the yield increase and Central and the metabolism of bacterial community (Chen et al. Composite Design was very effective in optimization of 2000; Flint et al. 2007). medium for growth of Bifidus. In a similar study, Ketabi and Dieleman (2011) investigated the effect of isomalto- Prebiotics have multidimensional effect on host‑bacteria oligosaccharides on intestinal microflora of rats and interaction inferred that it specifically stimulated the growth of It is now well established fact that host bacteria inter- Lactobacilli. actions are highly specific with varied dimensions. The digestion resistant carbohydrates in the gut are fer- Removal of cholesterol mented in the colon which causes increase in the serum Cholesterol was found to be evidently removed by Lacto- lactate levels. The study was conducted on horses by bacillus acidophilus ATCC 4962 in the presence of prebi- injecting fructooligosaccharides directly in caecum and otics in a study conducted by Liong and Shah (2005). The acidotic state was maintained. Its effect on caecum bacte - effect of six prebiotics including fructooligosaccharides ria and metabolites were analyzed. Streptococcal species was used to investigate the best combination for effec - (EHSS) showed positive relation with caecum lactate and tive removal of cholesterol. The first-order model, the negative response with serum lactate; however, serum second-order polynomial regression model and quadratic lactate has a positive influence on Enterobacteriaceae models were used in their study. (Rudi 2010). Artificial sweetness Genetic features direct the probiotic effect of bacteria Apart from all the above stated prime health benefits Excellent studies on genomics of lactic acid bacteria in fructooligosaccharides also has artificial sweetness and relation to their role in functional foods have been done low caloric value. Artificial sweeteners are constantly in by Klaenhammer et al. (2005). Their findings discovered demand due to need of diabetics and health conscious that many genetic features exert control over the bacte- consumers. Initially the demand was satisfied by aspar - rial metabolic and probiotic process. tame agent or natural sweeteners like palatinose. Due to their popular use all types of oligosaccharides remained Development of resistance to ill effects of bile salts poorly exploited despite their functional properties Fructooligosaccharides and their monomeric derivatives (Mussatto et al. 2009). offer resistance against the ill effects of bile salts on Bifi - dus group of intestinal inhabitants. Perrin et al. (2001) Role in osteoporosis studied the inhibitory effect of bile salts on three strains The most recent trial of fructooligosaccharides supple - of Bifidobacterium in medium containing any carbohy - mented with calcium in post menopausal women have drate source. In presence of fructooligosaccharides in registered beneficial effects in bone mineral density which the medium the Bifidobacterium improved their resist - is highly significant in osteoporosis (Slevin et al. 2014). ance and demonstrated better growth in presence of bile salts. Macfarlane et al. (2008) studied the effect of Galactooligosaccharides (GOS) and Lactulose inulo, galacto and fructooligosaccharides was extremely derived galactooligosaccharides (LDGOS) favorable for Bifidobacterium and also Lactobacilli but to Mammalian milk is the natural source of GOS. Indus- a lesser extent. Their health benefits encompass features trially trans galactosylation of lactose present in whey like putative anti-cancer properties, mineral absorption, catalysed by β-galactosidases is gaining momentum as an lipid metabolism, anti-inflammatory and other immune promising alternative for synthesis of GOS (Affertsholt- effects such as atopic disease. Allen 2009). Belorkar and Gupta AMB Expr (2016) 6:82 Page 5 of 11 β-Galactosidase is a hydrolase that attacks the o-gluco- Xylooligosaccharides (XOS) syl group of lactose. The general mechanism of enzymatic Xylooligosaccharides or feruloyl oligosaccharides are lactose hydrolysis has a transgalactosylic nature, involv- known to be produced by Aspergillus, Trichoderma, Peni- ing a multitude of sequential reactions with disaccharides cillium, Bacillus and Streptomyces. It is found in plant (other than lactose) and higher saccharides, collectively sources like Bengalgram husk, wheat bran and straw, named galacto-oligosaccharides (GOS), as intermediate spentwood, barley hulls, brewery spent grains, almond products (Wallenfels and Malhotra 1960; Goulas et al. shells, bamboo and corn cob. XOS mainly exerts prebi- 2007). Non digestible oligosaccharides have wider appli- otic effect in consumers. cations (Sako et al. 1999). These unusual oligosaccharides are composed by The GOS are complex mixtures of oligosaccharides chains of xylose moieties linked by β-(1,4) bonds, ranging from two to eight moieties, and different glyco - with a polymerization degree ranging from two to ten sidic linkages: β-(1,1), β-(1,2), β-(1,3), β-(1,4) and β-(1,6) monosaccharides. (Playne and Crittenden 2009). The hydrolytic enzymes It is also known to act as a plant growth regulator. It preferentially expressed by Bifidobacterium species spe - has multidimensional applications as antioxidant and cifically target β-glycosidic linkages of GOS in the intes - gelling agent in food products, beneficial for diabetes, in tine (Macfarlane et al. 2008). treatment of arteriosclerosis, reduces total cholesterol Microbes are exuberant sources of the enzymes pro- and LDL in patients with type 2 diabetes mellitus and in ducing Lactulose and GOS (Nguyen et al. 2009; Splechtna colon cancer (Chung et al. 2007; Sheu et al. 2008; Lecerf et al. 2006, 2007; Maischberger et al. 2008; Placier et al. et al. 2012; Moure et al. 2006; Katapodis and Chistako- 2009). The operation conditions are to be properly moni - poulos 2008; Madhukumar and Muralikrishna 2010). tored for optimal ratio of lactulose and GOS for potential Figure 4 is a diagrammatic representation of applications synthesis of prebiotics (Guerrero et al. 2013; 2015). of XOS. The main physiological effects of GOS are related with their composition and activities of the intestinal microbiota Arabinooligosaccharides (AOS) (Algieri et al. 2014). The human intestinal tract harbors a Arabinooligosaccharides are yet another class of oligo- complex community of bacteria, eukaryotic microorganisms, saccharides which hold the potential of being labelled archaea, viruses, and bacteriophages, collectively referred to as prebiotics. The exuberant source of AOS is arabinan as the intestinal microbiota. Bacteria account for the major- polysaccharide a branched pectic polysaccaharide ity of these microorganisms: their total number in the human exhibiting linkage of 1,3 and 1,5 α l -arabinofurano- gut is estimated at 1014 cells mainly present in the colon syl residues (Vogel 1991). Arabinose occurs naturally (Backhed et al. 2005; Lupp and Finlay 2005). The wide appli - in arabinans, arabinogalactans or arabino xylans pre- cations of GOS and LDGOS are represented in Fig. 3. sent in plants cell wall components. The nature of linkages differ depending upon the sources. The brush border epithelial cells of the intestine are inefficient to degrade the polysaccharides present in plant cell wall. This resistance of cell wall polysaccharides towards intestinal hydrolysis confer them the potential to be used as prebiotics (Yoo et al. 2012; Rastall and Hotch- kiss 2003). The efficacy of the prebiotic effect of AOS is structure dependent (Casci et al. 2006; Gullón et al. 2011). Initially the extraction was practiced by hot alkali treat- ment (Cibe 2003) of sugar beet dried pulp (5.5 million tons) a major coproduct of beet sugar industries residue in European countries. AOS can also obtained by enzymatic hydrolysis of Arabinose containing polymers. Beldman et al. (1997) classified the Arabinan degrading enzymes in six classes- (i) α-l-Arabinofuranosidase (EC 184.108.40.206), which is not active with polymers (Komae et al. 1982; Weinstein Fig. 3 Functions of GOS and LDGOS and Alber sheim 1979). Belorkar and Gupta AMB Expr (2016) 6:82 Page 6 of 11 Fig. 4 Functions of XOS (ii) α-l -Arabinofuranosidase, which is active with support the intestinal bifidus flora nearly equal to FOS and polymers (Kaji and Tagawa, 1970; Rombouts et al. Inulooligosaccharides (Gómez et al. 2015; Palframan et al. 1988). 2002; Rycroft et al. 2001; Sanz et al. 2005). The extent of (iii) α-l-Arabinofuranohydrolase, which is specific response is directly proportional to the dp of the oligosac- for arabinoxylans (Kormelink et al. 1991; Van Laere charide (Sulek et al. 2014; Westphal et al. 2010). et al. 1997). Apart from the normal benefits, AOS is reported to (iv) exo-α-l-Arabinanase, which is not active reduce the inflammatory conditions in Ulcerative coli - with p-nitrophenyl-α-l-arabinofuranoside (Kaji and tis patients. Invitro experiments have proved about spe- Shimokawa1984; McKie et al. 1997). cific stimulation of Bifidobacterium and Lactobacillus (v) β-l-Arabinopyranosidase (Dey 1983; Kaji and Saheki accompanied by production of SCFA acetate which is well 1975). known stimulator of anti inflammatory response. AOS (vi) endo-1, 5-α-l-Arabinanase (EC 220.127.116.11) (Vora - can prove to be a boon for patients suffering from Ulcera - gen et al. 1987). tive colitis after in vivo confirmation (Vigsnæs et al. 2011). Algal-oligosaccharides lysate (AOL) and neoagarooli- gosaccharides (NAOS) occur in the algal polysaccha- The various degree of polymerization (dp) are obtained ride extracts (APEs) of Gracilaria and Monostromaand when subjected to ultrafiltration can produce Oligosaccha - inenzymatic hydrolysis of agarose. They have a prebiotic rides of uniform molecular weight (Matsubara et al. 1996; effect and also act as an antioxidant (Wu et al. 2005; Hu Jian et al. 2013). AOS derived from sugar beet pectin (Al- et al. 2006). Tamimi et al. 2006) and lemon peel (Hotchkiss et al. 2010) Belorkar and Gupta AMB Expr (2016) 6:82 Page 7 of 11 Algae‑derived marine oligosaccharides Other oligosaccharides Recently, algae are reported to contain bipolysaccha- Mannanoligosaccharides (MOS) are mainly isolated rides (Stengel et al. 2011; Barra et al. 2014). The bioac - from cell wall fragments of yeast. It was found to alter tive components mainly include glucose, starch and other the gut microflora in fishes. It has been used as an alter - polysaccharides (Hamed et al. 2015). Besides these, oli- native to antibiotics and added to improve the nutritive gosaccharides are another group of carbohydrates with value of broiler diets (Dimitroglou et al. 2010; Eseceli small dp containing 3–10 sugar units, ranging from et al. 2010). Chitosanoligosaccharides (COS) has been disaccharides and/or carbohydrates with up to 20 resi- recorded to be produced by depolymerisation of chi- dues with defined functions (Patel and Goyal 2010). tosan. They are mainly used as an antioxidative agent, The chemical structure and conformation decides the anti-tumor agent and anti-microbial agent. Chitosan classification of algae-derived marine oligosaccharides oligosaccharides have been recorded to protect normal namely chitosan-, laminarin-, alginate-, fucoidan-, carra- cells from apoptosis (Liu et al. 2010). Human milko- geenan- and ulvan-oligosaccharides. ligosaccharide (HMO) naturally occurs in human breast The note worthy bioactive compounds in Marine mac - milk. It signifies the preferential growth of Bifidobacte - roalgae or seaweeds is namely polysaccharides, tannins, rium and Lactobacilli in the colon of mother fed babies and diterpenes. (O’sullivan et al. 2010). These ingredients (Quigley 2010). Gentiooligosaccharides (GeOS) is pro- may lead a pivitol role in nutraceuticals (Milinki et al. duced by digestion of starch and mainly used as a prebi- 2011). The functions of ADMO are given in Fig. 5. otic (Cote 2009; Fujimoto et al. 2009). Fig. 5 Functions of AOS Belorkar and Gupta AMB Expr (2016) 6:82 Page 8 of 11 Pectin-derived acidic oligosaccharides (pAOS) occur cheaper alternatives for purification strategies of synthe - in higher plant products like fruits and vegetables. It sized oligosaccharides. mainly finds its applications in infant formulae to sub - side diarrhoea, increase absorption of minerals and Future prospects calcium ions and also has antioxidant effects (Liu et al. As stated in the introduction of the review the demand 2010). pAOS also successfully helped in the lung infec- of health promoting food is expected to rise up to US tions by modulating the intestinal microbiota and the $4.8 billion by 2018. The hike in the demand is indicative inflammatory and immune responses (Bernard et al. of the future directions towards which the food indus- 2015). Cyclodextrins (CDs) are produced by transfor- try is fastly marching. The so called health promoting mation of starch by Bacillus macerans. It is used as a food or pro and prebiotics under the unanimous label stabilizer for volatile compounds in food preparations of “Nutraceuticals” will be a focus of attraction for every and chemicals. It acts as an antioxidant. It is used as such layman growing conscious about health in near taste enhancers in bitter medicines and food items future. The present scenario of the health market trend (Astray et al. 2009; Courtois 2009). is facing certain health issues pertaining to intake of Although all oligosaccharides are exhibiting prebi- the prebiotics viz. aggravation of intolerance to lactose, otic properties but fructo-oligosaccharides has gained increments in allergic responsiveness of sensitive indi- much attention as artificial sweeteners because they viduals as reported in several human based case studies. provide sweet taste to the consumer and do not increase Looking forward with this setback associated with the blood glucose level. Therefore, they find important probiotics, prebiotic are coming up as more promis- place in the food of diabetics. Thus, fructooligosaccha - ing option. Above all the prebiotic effect of oligosaccha - rides act as artificial sweeteners with functional prop - rides are now extended to their antidiarrheal, antiobesity erties apart from sweetness similar to that of natural and presently towards suppression of type 2 diabetes. sweeteners. The future would really depend on the synergistic effect Oligosaccharides from various sources have been con- developed by combinational use of prebiotics and pro- sidered as boon due to health benefits they encompass biotics. The incremental benefits of synbiotics would be along with property of being used as an artificial sweetner. auxiliary to the nature’s boon. Due to the diversified health benefits conferred by them, they have earned a prominent recognition as Nutraceu- Abbreviations ticals presently limelighted in the health market. The ADMO: algae derived marine oligosaccharides; AOS: arabinooligosaccharides; microbial production of enzymes catering the catalysis CD: cyclodextrins; dp: degree of polymerization; FOSHU: food of specified health use; FOS: fructooligosaccharides; GOS: galactooligosaccharides; GIA: of oligosaccharides are now targeted by the biotechnolo- Global Industry Analyst; HMO: human milk oligosaccharides; LDGOS: lactulose gists for their optimum synthesis. Microorganisms, espe- derived galactooligosaccharides; MOS: maltooligosaccharides; pAOS: pectin- cially molds have been the most prominent microbe for derived acidic oligosaccharides; XOS: xylooligosaccharides. enzymatic synthesis of the prebiotic oligosaccharides. Authors’ contributions Since 1980s teeming research work was focused towards The corresponding author has prepared script under the guidance of co isolation of potent microbes for oligosaccharide synthe- author. Both authors read and approved the final manuscript. sis. The oligosaccharide production has been successfully Author details attempted employing diverse approach viz. SmF, SSF, 1 Department of Microbiology and Bioinformatics, Bilaspur University, 206, immobilization of the intact microbial cells or derived Budhiya complex, Sarkanda, Bilaspur, Chhattisgarh 495004, India. Pt. Ravis- hankar Shukla University, Raipur, CG 492010, India. enzymes. The successful attempts have been made to improve the strain through mutations. Acknowledgements These laboratory processes have although recorded The author’s acknowledge the support of SOS in Life Sciences, Pt. Ravishankar Shukla University, Raipur and Department of Microbiology and Bioinformatics, successful production of oligosaccharides but scaling Bilaspur University, Bilaspur for supporting the work for which the topic was up introduces exuberant increase in the cost of produc- reviewed. tion of oligosaccharides. The bio process improvement Competing interests should be inculcated using cheaper agro-industrial The authors declare that they have no competing interests. wastes as substrates for oligosaccharide production. To decrease the cost of production following issues have to Funding The present review is not funded by any funding agency. be addressed: (i) a potent and stable microbial enzyme source is to be fetched (ii) scrutinizing agro-industrial Received: 4 April 2016 Accepted: 15 September 2016 wastes befitting the oligosaccharide production (iii) Belorkar and Gupta AMB Expr (2016) 6:82 Page 9 of 11 References Fernandez RC, Maresma BG, Juarez A, Martinez J. Production of fructooligo- Aer ff tsholt-Allen T. Market developments and industry challenges for lactose saccharides by β-fructofuranosidase from Aspergillus sp. 27 H. J Chem and lactose derivatives. 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