Unravelling the conundrum of tannins in animal nutrition and health

This paper examines the nutritional and veterinary effects of tannins on ruminants and makes some comparisons with non-ruminants. Tannin chemistry per se is not covered and readers are referred to several excellent reviews instead: (a) Okuda T et al. Heterocycles 30:1195-1218 (1990); (b) Ferreira D and Slade D. Nat Prod Rep 19:517-541 (2002); (c) Yoshida T et al. In Studies in Natural Product Chemistry. Elsevier Science, Amsterdam, pp. 395-453 (2000); (d) Khanbabaee K and van Ree T. Nat Prod Rep 18:641-649 (2001); (e) Okuda et al. Phytochemistvy 55:513-529 (2000). The effects of tannins on rumen micro-organisms are also not reviewed, as these have been addressed by others: (a) McSweeney CS et al. Anim Feed Sci Technol 91:83-93 (2001); (b) Smith AH and Mackie RI. Appl Environ Microbiol 70:1104-1115 (2004). This paper deals first with the nutritional effects of tannins in animal feeds, their qualitative and quantitative diversity, and the implications of tannin-protein complexation. It then summarises the known physiological and harmful effects and discusses the equivocal evidence of the bioavailability of tannins. Issues concerning tannin metabolism and systemic effects are also considered. Opportunities are presented on how to treat feeds with high tannin contents, and some lesser-known but successful feeding strategies are highlighted. Recent research has explored the use of tannins for preventing animal deaths from bloat, for reducing intestinal parasites and for lowering gaseous ammonia and methane emissions. Finally, several tannin assays and a hypothesis are discussed that merit further investigation in order to assess their suitability for predicting animal responses. The aim is to provoke discussion and spur readers into new approaches. An attempt is made to synthesise the emerging information for relating tannin structures with their activities. Although many plants with high levels of tannins produce negative effects and require treatments, others are very useful animal feeds. Our ability to predict whether tannin-containing feeds confer positive or negative effects will depend on interdisciplinary research between animal nutritionists and plant chemists. The elucidation of tannin structure-activity relationships presents exciting opportunities for future feeding strategies that will benefit ruminants and the environment within the contexts of extensive, semi-intensive and some intensive agricultural systems. (c) 2006 Society of Chemical Industry

Feeding rumen-inert fats differing in their degree of saturation decreases intake and increases plasma concentrations of gut peptides in lactating dairy cows

Our objective was to determine the effect of feeding rumen-inert fats differing in their degree of saturation on dry matter intake (DMI), milk production, and plasma concentrations of insulin, glucagon-like peptide 1 (7-36) amide (GLP-1), glucose-dependent insulinotropic polypeptide (GIP), and cholecystokinin (CCK) in lactating dairy cows. Four midlactation, primiparous Holstein cows were used in a 4 x 4 Latin square experiment with 2-wk periods. Cows were fed a control mixed ration ad libitum, and treatments were the dietary addition (3.5% of ration dry matter) of 3 rumen-inert fats as sources of mostly saturated fatty acids (SFA), monounsaturated fatty acids (MUFA), or polyunsaturated fatty acids (PUFA). Daily DMI, milk yield, and composition were measured on the last 4 d of each period. Jugular vein blood was collected every 30 min over a 7-h period on d 12 and 14 of each period for analysis of plasma concentrations of hormones, glucose, and nonesterified fatty acids. Feeding fat decreased DMI, and the decrease tended to be greater for MUFA and PUFA compared with SFA. Plasma concentration of GLP-1 increased when fat was fed and was greater for MUFA and PUFA. Feeding fat increased plasma glucose-dependent insulinotropic polypeptide and CCK concentrations and decreased plasma insulin concentration. Plasma CCK concentration was greater for MUFA and PUFA than for SFA and was greater for MUFA than PUFA. Decreases in DMI in cows fed fat were associated with increased plasma concentrations of GLP-1 and CCK and a decreased insulin concentration. The role of these peptides in regulating DMI in cattle fed fat requires further investigation.

Evaluation of NRC, UC Davis and ADAS approaches to estimate the metabolizable energy values of feeds at maintenance energy intake from equations utilizing chemical assays and in vitro determinations

Feed samples received by commercial analytical laboratories are often undefined or mixed varieties of forages, originate from various agronomic or geographical areas of the world, are mixtures (e.g., total mixed rations) and are often described incompletely or not at all. Six unified single equation approaches to predict the metabolizable energy (ME) value of feeds determined in sheep fed at maintenance ME intake were evaluated utilizing 78 individual feeds representing 17 different forages, grains, protein meals and by-product feedstuffs. The predictive approaches evaluated were two each from National Research Council [National Research Council (NRC), Nutrient Requirements of Dairy Cattle, seventh revised ed. National Academy Press, Washington, DC, USA, 2001], University of California at Davis (UC Davis) and ADAS (Stratford, UK). Slopes and intercepts for the two ADAS approaches that utilized in vitro digestibility of organic matter and either measured gross energy (GE), or a prediction of GE from component assays, and one UC Davis approach, based upon in vitro gas production and some component assays, differed from both unity and zero, respectively, while this was not the case for the two NRC and one UC Davis approach. However, within these latter three approaches, the goodness of fit (r(2)) increased from the NRC approach utilizing lignin (0.61) to the NRC approach utilizing 48 h in vitro digestion of neutral detergent fibre (NDF:0.72) and to the UC Davis approach utilizing a 30 h in vitro digestion of NDF (0.84). The reason for the difference between the precision of the NRC procedures was the failure of assayed lignin values to accurately predict 48 h in vitro digestion of NDF. However, differences among the six predictive approaches in the number of supporting assays, and their costs, as well as that the NRC approach is actually three related equations requiring categorical description of feeds (making them unsuitable for mixed feeds) while the ADAS and UC Davis approaches are single equations, suggests that the procedure of choice will vary dependent Upon local conditions, specific objectives and the feedstuffs to be evaluated. In contrast to the evaluation of the procedures among feedstuffs, no procedure was able to consistently discriminate the ME values of individual feeds within feedstuffs determined in vivo, suggesting that the quest for an accurate and precise ME predictive approach among and within feeds, may remain to be identified. (C) 2004 Elsevier B.V. All rights reserved.

The effect of wilting and soaking Eupatorium adenophorum on its digestibility in vitro and voluntary intake by goats

An experiment was conducted to determine what effect simple treatments might have on the voluntary intake by goats in Nepal of Eupatorium adenophorum, an invasive weed that is usually only consumed by goats to a very limited extent. Samples of E. adenophorum were collected and either untreated, soaked for 2 h or wilted for 2 h before being oven dried (60 degrees C) and ground. Soaking and wilting had little effect on the chemical composition of E. adenophorum, but did increase (P=0.036) its in vitro organic matter degradability, by approximately 8%. The short-term intake rate (STIR) of treated and untreated E. adenophorum was then estimated with eight goats. Soaking time (from 2 to 24 h) was not related to STIR (r = -0.111, P=0.198), but the time E. adenophorum was left to wilt (from 2 to 48h), was positively related to STIR (r=0.521, P<0.001), with values of STIR (g dry matter/min kg goat liveweight(0.75)) being 0.405, 0.649,1.058, S.E.M. 0.088 for E. adenophorum, that had been wilted for 0, 24 and 48 h respectively (P<0.001). Liveweight change of goats and voluntary intake of E. adenophorum by goats was then estimated with 24 goats. E. adenophorum was fed either unwilted, or wilted for 24 or 48 h. It was fed as the sole forage or as a 3:1 mixture (dry matter basis) with Ficus cunia. There was a linear (P<0.001) and quadratic (P<0.01) increase in the intake of total forage and E. adenophorum with wilting time of E. adenophorum. Offering Ficus cunia increased total forage intake, but decreased E. adenophorum intake (P<0.05). After four weeks, there was virtually no change in goat liveweight and no significant difference between treatments. The results suggest that wilting E adenophorum for 24 h could increase its intake by goats, and thereby increase its usefulness, as a potential source of forage in the dry season of Nepal, when forage scarcity is a common constraint to livestock production. (C) 2007 Elsevier B.V. All rights reserved.

Inter-laboratory variation of in vitro cumulative gas production profiles of feeds using manual and automated methods

A study was conducted to estimate variation among laboratories and between manual and automated techniques of measuring pressure on the resulting gas production profiles (GPP). Eight feeds (molassed sugarbeet feed, grass silage, maize silage, soyabean hulls, maize gluten feed, whole crop wheat silage, wheat, glucose) were milled to pass a I mm screen and sent to three laboratories (ADAS Nutritional Sciences Research Unit, UK; Institute of Grassland and Environmental Research (IGER), UK; Wageningen University, The Netherlands). Each laboratory measured GPP over 144 h using standardised procedures with manual pressure transducers (MPT) and automated pressure systems (APS). The APS at ADAS used a pressure transducer and bottles in a shaking water bath, while the APS at Wageningen and IGER used a pressure sensor and bottles held in a stationary rack. Apparent dry matter degradability (ADDM) was estimated at the end of the incubation. GPP were fitted to a modified Michaelis-Menten model assuming a single phase of gas production, and GPP were described in terms of the asymptotic volume of gas produced (A), the time to half A (B), the time of maximum gas production rate (t(RM) (gas)) and maximum gas production rate (R-M (gas)). There were effects (P<0.001) of substrate on all parameters. However, MPT produced more (P<0.001) gas, but with longer (P<0.001) B and t(RM gas) (P<0.05) and lower (P<0.001) R-M gas compared to APS. There was no difference between apparatus in ADDM estimates. Interactions occurred between substrate and apparatus, substrate and laboratory, and laboratory and apparatus. However, when mean values for MPT were regressed from the individual laboratories, relationships were good (i.e., adjusted R-2 = 0.827 or higher). Good relationships were also observed with APS, although they were weaker than for MPT (i.e., adjusted R-2 = 0.723 or higher). The relationships between mean MPT and mean APS data were also good (i.e., adjusted R 2 = 0. 844 or higher). Data suggest that, although laboratory and method of measuring pressure are sources of variation in GPP estimation, it should be possible using appropriate mathematical models to standardise data among laboratories so that data from one laboratory could be extrapolated to others. This would allow development of a database of GPP data from many diverse feeds. (c) 2005 Published by Elsevier B.V.

Dichrostachys cinerea and Acacia nilotica fruits as dry season feed supplements for goats in a semi-arid environment: Summary of a DFID funded project in Zimbabwe

Indehiscent fruits of six tree species, common in Matabeleland were examined in in vitro and in vivo trials. The results for two of them, Acacia nilotica and Dichrostachys cinerea are presented here. Acacia nilotica-contained more total phenolics than D. cinerea, but less nitrogen (N) and fibre (ADF and NDF). After 48 h incubation, in vitro OMD of both species was increased by PEG and NaOH or wood ash treatment, except when NaOH or wood ash were used in combination with PEG with D. cinerea fruits. DM intake, DMD were lowest and N-retention negative in goats fed A. nilotica as supplement. However when fed a supplement of D. cinerea, untreated or treated with PEG or NaOH, digestibility and N-retention were highest, and similar to a commercial goat meal, with the untreated fruit. In a trial in which milking does were supplemented with D. cinerea fruits, for 65 before and 65 days after kidding, kid birthweight and weaning weight were increased by supplementation. Deaths of twin-born kids were lowest in the supplemented but unmilked group. Supplementation with D. cinerea fruit resulted in improved goat performance. The only treatment applied of practical significance, wood ash, is currently being tested in an in vivo study. More research is required on detoxification of tannins, especially with A. nilotica. (c) 2005 Elsevier B.V. All rights reserved.

Summer drought alters plant-mediated competition between foliar- and root-feeding insects

Summer droughts are predicted to increase in severity and frequency in the United Kingdom, due to climate change. Few studies have addressed the impacts of drought on interactions between species, and the majority have focussed on increases in CO2 concentration and changes in temperature. Here, the effect of experimental summer drought on the strength of the plant-mediated interaction between leaf-mining Stephensia brunnichella larvae and root-chewing Agriotes larvae was investigated. Agriotes larvae reduced the abundance and performance of S. brunnichella feeding on a mutual host plant, Clinopodium vulgare, as well as the rate of parasitism of the leaf-miner. The interaction did not, however, occur on plants subjected to a severe drought treatment, which were reduced in size. Changes to summer rainfall, due to climate change, may therefore reduce the occurrence of plant-mediated interactions between insect herbivores.

Responses of invertebrate trophic level, feeding guild and body size to the management of improved grassland field margins

P>1. Management of lowland mesotrophic grasslands in north-west Europe often makes use of inorganic fertilizers, high stocking densities and silage-based forage systems to maximize productivity. The impact of these practices has resulted in a simplification of the plant community combined with wide-scale declines in the species richness of grassland invertebrates. We aim to identify how field margin management can be used to promote invertebrate diversity across a suite of functionally diverse taxa (beetles, planthoppers, true bugs, butterflies, bumblebees and spiders). 2. Using an information theoretic approach we identify the impacts of management (cattle grazing, cutting and inorganic fertilizer) and plant community composition (forb species richness, grass species richness and sward architecture) on invertebrate species richness and body size. As many of these management practices are common to grassland systems throughout the world, understanding invertebrate responses to them is important for the maintenance of biodiversity. 3. Sward architecture was identified as the primary factor promoting increased species richness of both predatory and phytophagous trophic levels, as well as being positively correlated with mean body size. In all cases phytophagous invertebrate species richness was positively correlated with measures of plant species richness. 4. The direct effects of management practices appear to be comparatively weak, suggesting that their impacts are indirect and mediated though the continuous measures of plant community structure, such as sward architecture or plant species richness. 5. Synthesis and applications. By partitioning field margins from the remainder of the field, economically viable intensive grassland management can be combined with extensive management aimed at promoting native biodiversity. The absence of inorganic fertilizer, combined with a reduction in the intensity of both cutting and grazing regimes, promotes floral species richness and sward architectural complexity. By increasing sward architecture the total biomass of invertebrates also increased (by c. 60% across the range of sward architectural measures seen in this study), increasing food available for higher trophic levels, such as birds and mammals.

Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method

Similarities between the anatomies of living organisms are often used to draw conclusions regarding the ecology and behaviour of extinct animals. Several pterosaur taxa are postulated to have been skim-feeders based largely on supposed convergences of their jaw anatomy with that of the modern skimming bird, Rynchops spp. Using physical and mathematical models of Rynchops bills and pterosaur jaws, we show that skimming is considerably more energetically costly than previously thought for Rynchops and that pterosaurs weighing more than one kilogram would not have been able to skim at all. Furthermore, anatomical comparisons between the highly specialised skull of Rynchops and those of postulated skimming pterosaurs suggest that even smaller forms were poorly adapted for skim-feeding. Our results refute the hypothesis that some pterosaurs commonly used skimming as a foraging method and illustrate the pitfalls involved in extrapolating from limited morphological convergence.

Comparison of the euryarchaeal microbial community in guts and food-soil of the soil-feeding termite Cubitermes fungifaber across different soil types

Termites are an important component of tropical soil communities and have a significant affect on the structure and nutrient content of soil. Digestion in termites is related to gut structure, gut physico-chemical conditions and gut symbiotic microbiota. Here we describe the use of 16S rRNA gene sequencing and Terminal-restriction Fragment Length Polymorphism (T-RFLP) analysis to examine methanogenic Archaea (MA) in the guts and food-soil of the soil-feeder Cubitermes fungifaber Sjostedt across a range of soil types. If they are strictly vertically inherited, then MA in guts should be the same in all individuals even if the soils differ across sites. In contrast, gut MA should reflect what is present in soil if populations are merely a reflection of what is ingested as the insects forage. We show clear differences between the euryarchaeal communities in termite guts and in food-soils from five different sites. Analysis of 16S rRNA gene clones indicated little overlap between the gut and soil communities. Gut clones were related to a termite-derived Methanomicrobiales cluster, to Methanobrevibacter and, surprisingly, to the haloalkaliphile Natronococcus. Soil clones clustered with Methanosarcina, Methanomicrococcus or Rice Cluster I. T-RFLP analysis indicated that the archaeal communities in the soil samples differed from site to site, whereas those in termite guts were similar between sites. There was some overlap between the gut and soil communities but these may represent transient populations in either guts or soil. Our data does not support the hypothesis that termite gut MA are derived from their food soil but also does not support a purely vertical transmission of gut microflora.

Comparison of Euryarchaea strains in the guts and food-soil of the soil-feeding termite Cubitermes fungifaber across different soil types

Termites are an important component of tropical soil communities and have a significant effect on the structure and nutrient content of soil. Digestion in termites is related to gut structure, gut physicochemical conditions, and gut symbiotic microbiota. Here we describe the use of 16S rRNA gene sequencing and terminal-restriction fragment length polymorphism (T-RFLP) analysis to examine methanogenic archaea (MA) in the guts and food-soil of the soil-feeder Cubitermes fungifaber Sjostedt across a range of soil types. If these MA are strictly vertically inherited, then the MA in guts should be the same in all individuals even if the soils differ across sites. In contrast, gut MA should reflect what is present in soil if populations are merely a reflection of what is ingested as the insects forage. We show clear differences between the euryarchaeal communities in termite guts and in food-soils from five different sites. Analysis of 16S rRNA gene clones indicated little overlap between the gut and soil communities. Gut clones were related to a termite-derived Methanomicrobiales cluster, to Methanobrevibacter and, surprisingly, to the haloalkaliphile Natronococcus. Soil clones clustered with Methanosarcina, Methanomicrococcus, or rice cluster I. T-RFLP analysis indicated that the archaeal communities in the soil samples differed from site to site, whereas those in termite guts were similar between sites. There was some overlap between the gut and soil communities, but these may represent transient populations in either guts or soil. Our data do not support the hypothesis that termite gut MA are derived from their food-soil but also do not support a purely vertical transmission of gut microflora.

Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method

Similarities between the anatomies of living organisms are often used to draw conclusions regarding the ecology and behaviour of extinct animals. Several pterosaur taxa are postulated to have been skim-feeders based largely on supposed convergences of their jaw anatomy with that of the modern skimming bird, Rynchops spp. Using physical and mathematical models of Rynchops bills and pterosaur jaws, we show that skimming is considerably more energetically costly than previously thought for Rynchops and that pterosaurs weighing more than one kilogram would not have been able to skim at all. Furthermore, anatomical comparisons between the highly specialised skull of Rynchops and those of postulated skimming pterosaurs suggest that even smaller forms were poorly adapted for skim-feeding. Our results refute the hypothesis that some pterosaurs commonly used skimming as a foraging method and illustrate the pitfalls involved in extrapolating from limited morphological convergence.

Probiotics and prebiotics in infant nutrition

The human colonic microflora has a central role in health and disease, being unique ill its complexity and range of functions. As such, dietary modulation is important for improved gut health, especially during the highly-sensitive stage of infancy. Diet call affect the composition of the gut microflora through the availability of different substrates for bacterial fermentation. Differences in gut microflora composition and incidence of infection exist between breast-fed and formula-fed infants, with the former thought to have improved protection. Historically, this improvement has been believed to be a result of the higher presence of reportedly-beneficial genera such as the bifidobacteria. As such, functional food ingredients such as prebiotics and probiotics could effect a beneficial modification in the composition and activities of gut microflora of infants by increasing positive flora components. The prebiotic approach aims to increase resident bacteria that are considered to be beneficial for human health, e.g. bifidobacteria and lactobacilli, while probiotics advocates the use of the live micro-organisms themselves in the diet. Both approaches have found their way into infant formula feeds and aim to more closely simulate the gut microbiota composition seen during breast-feeding.

Effects of phosphates and citrate on sediment formation in UHT goats' milk

Sediment formation was investigated during UHT treatment of goats' milk, subjected to indirect treatment at 140 degrees C for 2 s, with upstream homogenisation. Stabilisers evaluated were sodium hexametaphosphate (SHMP), trisodium citrate (TSC), disodium hydrogen orthophosphate (DSHP), and sodium dihydrogen orthophosphate (SDHP). With no added stabiliser, goats' milk produced a heavy sediment on UHT treatment. Addition of SDHP reduced pH, had little effect on ionic calcium and did not substantially reduce sediment. However, addition of SHMP, DSHP and TSC each reduced ionic calcium, increased ethanol stability and reduced sediment. Following stabiliser additions, there was a good correlation between ethanol stability and ionic calcium (R-2=0.85) but not between ethanol stability and pH (R-2=0.08). Overall, reducing ionic calcium reduced the amount of sediment formed for all these three stabilisers, although there was no single trend line between sediment formation and ionic calcium concentration. Sediment formation was not well correlated with pH for TSC or for SHMP, but it was for DSHP, making it the only stabiliser where sediment formation correlated well both with ionic calcium and pH, which might account for its effectiveness at higher ionic calcium levels. Sediment was much reduced when the temperature was reduced from 140 degrees C to 125 degrees C and 114 degrees C. There were no further changes in sediment on storage for two weeks. Analysis of the sediment showed that it was predominantly fat and protein, with a mass ratio ranging between 1.43:1 and 1.67:1. Its mineral content was usually less than 5% of dry weight. The maximum amounts of P and Ca were found to be 2.32% and 1.63%, respectively.