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ORAFTI, B3300 Tienen, Belgium
* To whom correspondence should be addressed. E-mail: jan.vanloo{at}orafti.com.
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ABSTRACT |
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 TOP ABSTRACT IntroductionLITERATURE CITED |
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Introduction |
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TOPABSTRACTIntroductionLITERATURE CITED |
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Inulin is composed of a set of molecules of sucrose of which the fructose moiety is substituted with a linear chain of ß(2–1) fructans ranging in length between 1 and 
65 fructose moieties. Oligofructose is a partial enzymatic hydrolysate of inulin. Typically, chicory inulin-type fructan chains with degree of polymerization (DP) < 10 are highly soluble in water (>80%), rapidly fermented, and interact significantly in a selective way with the intestinal flora. Chains that are longer than DP 10 are fermented slower, arrive in more distal parts of the intestine, and hence have a less important impact on the composition of the intestinal flora. Chicory inulin as extracted from the chicory roots contains 30–50% chains with DP < 10; the rest are longer chains. Oligofructose is entirely composed of chains with DP < 10. This distinction is important in animal nutrition; according to the intestinal architecture of the host (which is characterized by the volume of the different compartments, the transit of the feed bolus, and the bacterial density of each of the compartments) or the organ that is specifically targeted (small intestine, cecum, colon), either a short chain oligofructose or inulin that also contains an important fraction longer chains can be used.
It was observed that a particular combination of short and long inulin chains (Synergy) had more pronounced systemic effects. This justifies the use of this product in premium animal nutrition applications (pets) (4).
In the following section, a concise overview of nutritional research data in animals is presented. For a more detailed overview of literature data, the reader is referred to Flickinger (5). It is intended to link the zootechnical and well-being effects to factors that are directly related to prebiotic chicory inulin feeding.
Porcine nutrition
Although the pig is reportedly a good model for the human digestive system, there are some important differences in the intestinal architecture. There is a rather fast transit through the upper gastrointestinal (GI) tract (<8 h). Contrary to humans, there is a nonnegligible bacterial colonization in the pig ileum (density of up to 109 CFU compared with 106–7 log CFU in humans). As a consequence, fermentation of inulin and oligofructose already takes place to some extent in the small intestine. The rapidly fermented oligofructose is 30–40% fermented, whereas the inulin is 20–30% fermented in the upper GI tract. In humans with a lower bacterial overgrowth, only 10% of either oligofructose or inulin is fermented in the upper GI tract (6). Once either inulin or oligofructose arrives in the pig cecum, it is rapidly metabolized by the resident flora.
A positive dose effect on SCFA production and distal small intestinal villus height and crypt depth was demonstrated. In a separate experiment with both inulin and oligofructose, a significant increase in villus height vs. crypt depth ratio in the distal small intestine was observed (7). The pool of SCFA was also increased, which is indicative for stimulated bacterial activity. Correlating these findings with other experiments where an improved zootechnical performance with oligofructose or inulin was observed (7–11) seems to confirm the hypothesis that through bacterial fermentation and the concomitant increased production of butyrate (preferred energy source of the epithelial cells) of the nondigestible inulin or oligofructose, the intestinal architecture is improved (12). The increased absorptive capacity of the intestinal tract results in improved feed conversion and faster growth (7).
Another interesting application of prebiotic inulin in porcine nutrition is suppression of boar taint. Feeding male pigs an inulin preparation of up to 60 kg/metric ton suppresses proteolytic conversion of tryptophan into skatol, which within 3–5 d reduces the typical odor that makes the meat of intact pigs unsuitable for human consumption (13,14).
Also, prebiotic inulin-type fructans further strengthen the colonization resistance against invading pathogens. Moreover, feeding inulin suppresses parasites such as Ascaris (15) and Trichuris (16). The lowering of the intestinal pH, which is not favorable for the development of the parasite embryo, has been suggested as a possible mechanism.
Bovine nutrition
The rumen of a fully grown ruminant represents a huge fermentation organ in which the dietary inulin would be completely fermented and would not reach more distal areas of the GI tract where it can exert its beneficial activities. In calves, however, that are fed mainly a milk replacer, the rumen does not develop and from a digestive point of view the animal behaves as a monogastric. In these animals, inulin and oligofructose were shown to significantly increase growth performance (17). Intestinal infection in the young calf is a major problem. A significant improvement of the fecal consistency was observed in oligofructose- and inulin-fed groups compared with control animals. The inclusion of inulin in milk replacers also resulted in overall better growth and feed conversion (17).
Equine nutrition
Inulin-type fructans are often incorporated in commercial horse feed. There is, however, a paucity of scientific literature on this subject.
In preliminary studies where fistulated horses were monitored nutritionally sound intake levels of up to 2% on feed, oligofructose or inulin improved cecal fermentation (18) and the intake of the product was not associated with adverse effects of any kind. Further investigations should focus on reduction of incidence of colic.
Most available articles on inulin-type fructans in horses examining experimental reproduction of hoof laminitis by feeding massive amounts of up to 10 g fructans per kilograms bodyweight (19). Hoof laminitis is a condition resulting in lameness, which often leads to euthanasia of the animal. The doses that are used in these experiments are orders of magnitude higher than the doses having favorable effects on the digestive tract.
Poultry nutrition
Chickens typically have a very rapid intestinal transit with an orocloacal transit time <6 h. The ceca are perpendicular on the GI tract and the ileocecal valve evacuates insoluble material leaving the soluble material into the ceca (20). Through this mechanism, inulin and oligofructose enter the ceca. Chickens are used for meat production (broilers) as well as for egg production (laying hens).
Broilers. Feeding inulin or oligofructose to broilers resulted in significantly improved zootechnical performance, especially in female broilers. The improved performance again could be associated with a significantly increased absorptive capacity of the chicken GI tract due to increased length of both small intestine and colon (21–24). Rapidly fermented oligofructose had the most pronounced effects in this case. Feeding the chicory inulin-type fructans also had systemic effects. Serum cholesterol levels and deposit of fat tissue were significantly reduced, with the slower fermented inulin having the most pronounced effects. The observations were correlated with modified intestinal fermentation (25). Furthermore, in broilers artificially challenged with Salmonella or Campylobacter, oligofructose or inulin feeding suppressed infections and the positive impact on zootechnical performance was even more pronounced than in unchallenged animals (23,26–29).
Laying hens.
Laying hens produce 0.95 eggs/d for 55 wk, but afterwards the productivity goes down and eventually the hens are culled. When old laying hens were fed inulin or oligofructose, increased egg productivity was observed, but the effect with oligofructose was numerically more pronounced than with inulin (30). It is possible that the more rapidly fermented oligofructose is better adapted to the rapid intestinal transit times of these birds. Increased mineralization of the tibia suggests that in poultry, too, increased mineral (calcium) absorption is induced by the inulin-type fructans. In addition, the cholesterol content of the yolk was reduced in the prebiotic-fed animals (31).
Fish nutrition
Fish represent a diverse group of animals that are carnivorous, omnivorous, or herbivorous. Fish are ectothermic poikilotherms (cold-blooded) animals and they have a relatively simple intestinal tract. The bacterial population colonizing the fish intestine hardly overlap with the populations that are found in endothermic homeoterms (warm-blooded) animals.
Recent research on the effect of inulin on fish nutrition indicates that the prebiotic effect results in improved zootechnical performance, including increased growth and reduced feed conversion ratio of young carnivorous turbot (32). In herbivorous sturgeon and omnivorous catfish, improved zootechnical performance (specific growth rate and feed conversion ratio) was associated with longer intestine and modified intestinal fermentation, as shown by analysis of intestinal SCFA and bacteria vs. control group (32).
Feline and canine nutrition
In cats and dogs, where the prebiotic effect of the inulin-type fructans was confirmed (33), the impact of the prebiotics on systemic function related to prevention of chronic disease becomes important. It is a well-known phenomenon that similar to the human population, companion animals suffer from obesity, cardiovascular disease, and cancer. Inulin or oligofructose feeding of hyperlipidemic dogs decreased circulating cholesterol and triglyceride levels (34,35), which is a risk factor for cardiovascular disease. In cats, due to the increased intestinal fermentation, more nitrogen is fixed in the bacterial biomass, thus relieving the nitrogen burden on their kidneys (36). Similar effects were observed in dogs (37), but these omnivores suffer less from kidney insufficiency than the carnivorous cats. At present there is a research gap concerning the impact of prebiotic inulin-type fructans on the immune system and cancer risk suppression in pets.
Another aspect of prebiotic feeding to pets is the improvement of bowel habit and fermentation, resulting in improved stool quality, measured in terms of consistency and odor (38,39).
Well-being
Dietary inulin-type fructans contribute to animal well-being at several levels. Improvement of bowel habit is a general way in which inulin and oligofructose contribute to well-being. Relief of constipation (40) and relief of diarrhea (41), as observed in human studies, have also been confirmed in animal trials. Moreover, it was shown by means of short-term ethological studies in rats that cognitive performance, muscular tonus, and learning ability were significantly improved (42). These results were confirmed in a lifetime ethological rat trial in which the improved general well-being resulted in less obese animals with a whiter and softer fir than control animals. Ultimately, the rats fed chicory inulin-type fructans showed increased life span and the mortality curve at the end of the lifetime study was steeper in the inulin-fed animals, indicating that the period of morbidity prior to death was shortened (43).
The inulin-type fructans are prebiotic (nondigestible and specific interaction with intestinal bacteria) in various intestinal environments of diverse animal genera (mammal, fish, bird, etc.). The specific intestinal fermentation results in a bacterial ecosystem that is less prone to pathogen invasion and in an increased production of SCFA characterized by a higher proportion of butyrate. As a result, the increased absorptive capacity, reflected in longer intestinal length and increased villus height or crypt depth of the intestine, results in improved feed conversion and better growth in the young animal. The systemic effects on lipid metabolism result in reduced fat deposition in production animals such as broilers. In pets, the intestinal fermentation correlates with better bowel habit, resulting in improved stool quality. Earlier described systemic effects in experimental models and human studies seem to be valid for pets too and offer the potential of suppressing the emergence of chronic disease. Inulin and oligofructose feeding contribute to animal well-being, which is especially relevant in the short term for the young animal and in the long term for pets.
Future research needs to explore these properties in other animal species (e.g. crustaceans) and dose-effect relationships in practical production conditions (inexpensive and dense antibiotic-free feed and highly crowded environments).
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FOOTNOTES |
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2 Author disclosure: J. Van Loo, is an ORAFTI employee.
3 In these proceedings, the term inulin-type fructan shall be used as a generic term to cover all ß-(2
1) linear fructans. In any other circumstances that justify the identification of the oligomers vs. the polymers, the terms oligofructose and/or inulin or eventually long-chain/or high molecular weight inulin will be used, respectively. Even though the oligomers obtained by partial hydrolysis of inulin or by enzymatic synthesis have a slightly different DPav (4 and 3.6, respectively), the term oligofructose shall be used to identify both. Synergy will be used to identify the mixture of oligofructose and inulin HP otherwise named oligofructose-enriched inulin.
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LITERATURE CITED |
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TOPABSTRACTIntroductionLITERATURE CITED |
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1. Gibson GR, Probert HM, Van Loo J, Rastall RA, Roberfroid MB. Dietary modulation of the human colonic microbiota: updating the concept of prebiotics. Nutr Res Rev. 2004;17:259–75.
2. Van Loo J, Coussement P, de Leenheer L, Hoebregs H, Smits G. On the presence of inulin and oligofructose as natural ingredients in the western diet. Crit Rev Food Sci Nutr. 1995;35:525–52.[Medline]
3. Gibson G, Willis CL, Van Loo J. Non-digestible oligosaccharides and bifidobacteria: implications for health. Int Sugar J. 1994;96:381–7.
4. Van Loo J. The specificity of the interaction with intestinal fermentation by prebiotics determines their physiological efficacy. Nutr Res Rev. 2004;17:89–98.
5. Flickinger EA, Van Loo J, Fahey GC Jr. Nutritional responses to the presence of inulin and oligofructose in the diets of domesticated animals: a review. Crit Rev Food Sci Nutr. 2003;43:19–60.[Medline]
6. Ellegard L, Andersson H, Bosaeus I. Inulin and oligofructose do not influence the absorption of cholesterol, or the excretion of cholesterol, Ca, Mg, Zn, Fe, or bile acids but increases energy excretion in ileostomy subjects. Eur J Clin Nutr. 1997;51:1–5.[Medline]
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8. Spreeuwenberg M, Meijer J, Van Loo J. Inulin increases performance, but does not affect diarrhoea and faecal microbial profiles in E. coli challenged piglets. Poster presented at the Dietary Fibre 2003 symposium, Bio-active carbohydrates for food and feed, 18-21 May 2003, Noordwijkerhout, Netherlands.
9. Shipp TE, Madsen FC. Synergistic effects: prebiotics and piglet performance. USA Feed Management, 2003. p. 18–22.
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15. Petkevicius S, Knudsen KE, Nansen P, Roepstorff A, Skjoth F, Jensen K. The impact of diets varying in carbohydrates resistant to endogenous enzymes and lignin on populations of Ascaris suum and Oesophagostomum dentatum in pigs. Parasitology. 1997;114:555–68.
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17. Van Leeuwen P, Verdonk JMAJ. The gastro-intestinal degradation of inulin preparations and their effects on production performance and gut microflora in calves. 2004. Animal Sciences Group Wagingen UR, Lelystad, Netherlands, Project nr 825.20552.02, Report No: 04/l00287:1-31.
18. Van Loo J, Bosscher D, Janssens GP. Prebiotic inulin interacts with the caecal fermentation in the horse. Horse health nutrition: Third European Equine Health and Nutrition Congress, 17 and 18 March 2006, Faculty of Veterinary Medicine of Ghent University, Merelbeke, Belgium: 75-82.
19. Thoefner MB, Pollitt CC, Van Eps AW, Milinovich GJ, Trott DJ, Wattle O, Andersen PH. Acute bovine laminitis: a new induction model using alimentary oligofructose overload. J Dairy Sci. 2004;87:2932–40.
20. Zhang Y, Joergensen H, Fernandez J, Knudsen K. Digestibility of carbohydrates in growing pigs: a comparison between the t-cannula and the steered ileo-caecal valve cannula. Arch Anim Nutr. 2004;58:219–31.[Medline]
21. Yusrizal Chen TC. Effect of adding chicory fructans in feed on broiler growth performance serum cholesterol and intestinal length. Int J Poult Sci. 2003;2:214–9.
22. Van Leeuwen P, Verdonk JMAJ, Wagenaars CMF, Kwakernaak C. Effects of fructooligosaccharide inclusion in diets on performance before and after inoculations with Eimeria acervulina and Clostridium perfringens in broilers. 2005. Animal Sciences Group Wagingen UR, Lelystad, Netherlands, Project number 220.13180.04, Report nr. 05/l01056: 1-27.
23. Van Leeuwen P, Verdonck JMAJ, Wagenaars CMF, Kwakernaak C. The effect of three inulin preparations on performance before and after an inoculation with Salmonella and Campylobacter in broilers. 2005. Animal Sciences Group Wagingen UR, Lelystad, Netherlands, Project nr 825.20552.03, Report No: 05/l00651: 1-40.
24. Ammerman EQ. Broilers response to the addition of dietary fructooligosaccharides. Poult Sci. 1988;67:46.
25. Yusrizal X, Chen TC. Effect of adding chicory fructans in feed on fecal and intestinal flora. Int J Poult Sci. 2003;2:188–94.
26. Chambers JR, Spencer JL, Modler HW. The influence of complex carbohydrates on Salmonella typhimurium colonization, pH, and density of broiler ceca. Poult Sci. 1997;76:445–51.
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31. Chen Y, Chen T. Effects if chicory fructans on egg cholesterol in commercial layers. Int J Poult Sci. 2005;4:109–14.
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40. Den Hond EM, Geypens B, Ghoos Y. Effect of high performance chicory inulin on constipation. Nutr. Res. 2000;20:731–736.
41. Cummings JH, Christie S, Cole TJ. A study of fructo oligosaccharides in the prevention of travellers' diarrhoea. Aliment Pharmacol Ther. 2001;15:1139–45.[Medline]
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43. Messaoudi M, Rozan P, Nejdi A, Bisson JF, Hidago S, Desor D. Assessment of the effects of inulin-type fructans, daily administered during 21 months on biological parameters, emotional state and cognitive function. In press. 2007.
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