1. Primary growth vegetative Lotus pedunculatus containing 46 and 106 g/kg dry matter (DM) of total condsnsed tannin and 3 and 14 g/kg DM of free condensed tannin, was cut and fed fresh at hourly intervals (750 g DM/d) to sheep fitted with permanent cannulas into the rumen and duodenum. Low-and high-tannin lotus contained respectively 41.3 and 31.6 g total nitrogen/kg DM and 132 and 152 g lignin/kg DM. The two forms of lotus were similar in carbohydrate composition.2. Nutrient intake was recorded and faecal output measured by direct collection. Digesta flow to the duodenum was estimated by measuring dilution at the duodenum of inert ruthenium phenanthroline (Ru-P) and chromium-EDTA markers continuously infused into the rumen.3. Effects attributable to condensed tannins were assessed by comparing the digestion of the two diets, and by comparing the digestion of each with predicted values for non-tannin-containing fresh forages fed at similar intakes. 4.Apparent digestibility of all nutrients measured was less for high-than for low-tannin lotus (P < 0.01). The levels of cellulose digested ruminally and post-ruminally in both forms of lotus were similar to predicted values. However, less hemicellulose and readily fermentable carbohydrate (RFC; soluble carbohydrate +pectin) was digested in the rumen in sheep given both forms of lotus than would be predicted for non-tannin-containing fresh forage diets, but this was compensated for by greater post-ruminal absorption of both nutrients. 5.Total N gains across the rumen (duodenal N flow -total N intake) were 1.8 and 10.5 g/d for low-and high-tannin lotus v. predicted losses of 3.7 and 2.1 g/d for non-tannin-containing fresh forages given at the same total N intakes. Post-ruminal digestion of non-amonia-N (NAN; proportion NAN flowing at duodenum) was 0.71 and 0.67 for low-and high-tannin lotus respectively v. 0.76 for comparable non-tannin-containing fresh forages. Energy absorbed as amino acids from the small intestine was calculated to be 0.29 of metabolizable energy for both forms of lotus, compared with 0.17 and 0.21 for perennial ryegrass and white clover.6 . It was concluded that the presence of condensed tannins in lotus markedly increased post-ruminal NAN absorption compared with non-tannin-containing fresh forage diets, but depressed ruminal digestion of RFC and hemicellulose.

SUMMARYAreas of sulla (Hedysarum coronarium) and pasture (Lolium perenne/Trifolium repens/Holcus lanatus) were grazed by young sheep (29·5–34·8 kg initial liveweight) in four experiments, and effects upon body growth, wool growth and rumen metabolism were measured.Sulla contained 40–50 g condensed tannins (CT)/kg DM, whilst the pasture contained small amounts of CT (2–6 g/kg DM). After chewing during eating, a lower proportion of total CT was readily extractable and greater proportions were protein-bound and fibre-bound. Nutritional effects attributable to CT were assessed by oral administration of polyethylene glycol (PEG), which binds and inactivates CT, to half the animals grazing each forage. Rates of body growth were consistently higher for lambs grazing sulla than those grazing pasture, with the CT concentration in sulla being neither stimulatory or inhibitory to body growth or voluntary feed intake (VFI). The action of CT reduced carcass fatness in sheep fed both diets, in the one experiment where this was measured. CT present in both sulla and pasture decreased rumen ammonia concentration and decreased molar proportions of iso-butyrate and iso and n-valerate. During spring and early summer, when wool growth rates were highest, CT present in both pasture and sulla increased wool growth rate; when wool growth rates were low during winter, CT had no effect upon the wool growth of sheep grazing either forage. Numbers of protozoa and molar proportions of n-butyrate in rumen fluid were increased by CT in sheep grazing sulla but not pasture.It was concluded that the higher rates of body growth and VFI in lambs grazing sulla was most likely to be due to its very high ratio of readily fermentable: structural carbohydrate.

Interrelationships between the concentrations of total condensed tannin (TCT), free condensed tannin (FCT) and lignin were studied to gain knowledge of how to manipulate nutritive value of fresh herbages containing condensed tannins fed to ruminants. FCT was defined as condensed tannin not bound by macerates of fresh plants, with both FCT and TCT being determined with vanillin HCI. Effects of spraying lotus with polyethylene glycol (mol. wt 3350; PEG) upon the relationship between FCT and TCT was also studied. Increasing soil nutrient and climatic stress caused large and similar increases in the concentrations of TCT and of lignin. Over the range &90 g kg-' DM, 10% of TCT in Lotus sp. was detected as FCT, with increments in TCT above 90gkg-' D M being released almost entirely as FCT. PEG formed much stronger chemical bonds with condensed tannins than did plant proteins, and did not release FCT; consequently PEG application reduced the concentration of condensed tannin that was detectable with vanillin HCI. After disintegration of plant material, it is proposed that most condensed tannin is bound and co-precipitated as an insoluble complex with protein, that FCT is in equilibrium with this complex, and that bound and free tannin are indices of nutritionally beneficial and detrimental effects produced by condensed tannins in fresh forages eaten by ruminants. It was concluded that growing Lotus pedunculatus under conditions of stress leads to depressions in nutritive value through simultaneously increasing concentrations of lignin and FCT, both of which depress rumen carbohydrate digestion and voluntary intake, and that treatment with PEG offers a convenient method of separating effects due to condensed tannins from other factors influencing nutritive value.

1. Voluntary intake was determined with vegetative Lotus pedunculatus cut and fed fresh to growing sheep of 42-46 kg live weight. Effects attributable to condensed tannins were assessed by growing the plant under high and low levels of soil fertility, inducing low and high concentrations of tannin (Expt l), or by binding the tannins through spraying the herbage with polyethylene glycol (molecular weight 3350, PEG; Expt 2). Primary-growth lotus was used in Expt 1 and secondary-growth lotus in Expt 2. Concentrations of total and free condensed tannin were determined in fresh lotus, free tanning being defined as that not bound by mascerates of the plant.2. In Expt 1 the herbages fed contained respectively 46 and 106 g total condensed tannin/kg dry matter (DM) and 3 and 14 g free condensed tannin/kg DM. Mean metabolizable energy (ME) intakes were 0.89 and 0.77 MJ/kg live 3. The lotus used in Expt 2 contained 63 and 5 g total reactive condensed tannin and free condensed tannin/kg DM respectively. After spraying with PEG at 2.4 g/g total condensed tannin, these values were reduced to 7 and 0.5 g/kg DM respectively. PEG addition increased apparent digestibility (proportion of each nutrient ingested) of cellulose, hemicellulose and nitrogen by 0.05,0.08 and 0.26, and increased ME intake from 0.48 to 0.69 MJ/kg live ~eightO'?~ per d.4. It was concluded that high concentrations of condensed tannins depressed ME intake, due to depressions in both the voluntary intake and digestion of organic matter. The stimulation of hemicellulose and cellulose digestion by PEG addition suggests that the condensed tannin was depressing rumen digestion.5. It is considered that free tannins are most likely to have been responsible for the depressive effects in the present study. This could be due to their reaction with microbial enzymes in the rumen, the tannin content of the plant exceeding the capacity of the plant protein to bind it, or to reaction with enzymes secreted into the small intestine following the absorption of amino acids. In either instance free tannins could also react with proteins of the gut wall.Reponses in protein deposition in body tissue and in wool to abomasal infusion of casein + methionine in growing lambs fed on fresh ryegrass (Lolium perenne)-based herbage ad lib. have shown that absorption of one or more essential amino acids from the small intestine is below animal requirements (Barry, 1981). A subsequent review (Barry, 1982) concluded that lactating ewes and dairy cows consuming similar fresh pasture ad lib. of high metabolizable energy (ME; 11.3 MJ ME/kg dry matter (DM)) and total nitrogen (33 g/kg DM) content were also likely to be deficient in the absorption of essential amino acids, relative to the requirements for optimum production at the level of voluntary ME intake achieved. One of the two main predisposing causes of this deficit was identified as the exceptionally high degradation rate of the proteins in fresh forages in the rumen (0.7).

The condensed tannin concentration in Lotus pedunculatus (cv. Grasslands 'Maku') was 8-1 1 % D M when grown in acid soils without fertiliser application and 2-3 % D M when grown in high fertility soils. Application of P & S fertiliser to the acid soils increased DM yield and reduced condensed tannin content to 6 5 % DM, with over 88 % of the variation in condensed tannin content being explained by variation in DM yield; it also increased plant total N concentration and halved the molar ratio of condensed tannin : protein (MR). Condensed tannins quantitatively precipitated soluble protein in lotus and also effectively precipitated protein in mixtures of lotus with white clover. The minimum concentration of condensed tannin necessary to precipitate protein was 2 4 % DM, corresponding to MR values of 6-13. However, protein from such mixtures could be deaminated by fermentations with rumen fluid in uituo, but potentially soluble protein in pure lotus (MR 29) was protected from deamination by condensed tannins. A portion of lotus tannin could not be bound by plant constituents in finely ground fresh herbage and was designated 'free' tannin. This fraction was linearly related to total plant tannin content, was predicted to be zero at 1.8% DM total tannin, and increased at 0.15 units per unit increase in total tannin content above this figure. Maximum 'free' tannin comprised 17% of total condensed tannin. Effects of total and 'free' tannin content upon the intake and digestion of lotus by sheep are discussed.