Review: Closing nutrient cycles for animal production - Current and future agroecological and socio-economic issues.

We face an urgent and complex challenge to produce large amounts of healthful animal and plant foods for an estimated 10 billion people by 2050 while maintaining essential ecosystem services. To compound this challenge, we must do so while not further degrading our environment and conserving essential nutrients such as copper, magnesium, phosphorus, selenium, and zinc that are in short supply for fertilization. Much good research has been done, but to meet this challenge, we need to greatly increase on-farm and watershed-scale research including on-farm evaluations and demonstrations of the putative best combinations of stewardship techniques over multiple years in real-world settings, which are backed by data on nutrient inputs, soil, air, and water chemistry (fluxes) and water discharge. We also need to work with farmers, specialists, and generalists in highly creative interdisciplinary teams that resist forming silos and that use combinations of techniques linked to agroecology and industrial ecology in combination with state-of-the-art engineering. Some of these research and demonstration farms need to be in catchments prone to pollution of aquatic and terrestrial ecosystems with nitrogen, phosphorus, and other nutrients. Some promising approaches include mixed crop-livestock systems, although these alone may not be productive enough without updating to meet the dietary needs of an estimated 10 billion people by 2050. Other approaches could be state-of-the-art multi-trophic production systems, which include several species of plants integrated into production with vertebrates (e.g., ruminants, pigs, poultry), invertebrates (e.g., insects, earthworms) and fish, shrimp, or crayfish to utilize wasted feed and excreta, and recycle nutrients back to the animals (via plants or invertebrates) in the systems. To cut costs and increase desirable outputs, we must recycle nutrients much better within our food production systems and produce both animal and plant foods more efficiently as nutrients cycle through systems.

Preference by sheep for endophyte-infected tall fescue grown adjacent to or at a distance from alfalfa.

Two experiments were conducted to assess preference by sheep for endophyte-infected tall fescue growing in monoculture at least 5 m away from alfalfa (fescue-middle (FM)) over endophyte-infected tall fescue growing adjacent (0.2 to 1 m; fescue-alfalfa (FA)) to alfalfa (FA), and the effect of legume scent on preference for endophyte-infected tall fescue. In Experiment 1, 10 six-month-old lambs were offered for 12 days a choice of freshly harvested FA and FM. On days 13 and 14, lambs were offered the same choice, except cages (to allow access only to scent) containing freshly harvested alfalfa were put in the feeders containing FA, whereas cages containing freshly harvested FM were included with the feeders containing FM. Forage intake was measured 1 h after feeding and at three consecutive 2-h intervals thereafter. FA contained greater (P<0.002) concentrations of the alkaloid ergovaline (360 ± 27 ppm) and CP (8 ± 0.4%) than FM (219 ± 27 ppm and 6 ± 0.4%, respectively). Lambs preferred (P<0.05) FA to FM during the 1st hour of feeding, but the differences became smaller and disappeared in later feeding periods (P<0.005). Lambs offered FA with alfalfa scent or FM with FM scent preferred (P<0.05) FA but only on the 2nd day. In Experiment 2, 10 six-month-old lambs were offered a choice of FM with cages (to allow access only to scent) containing freshly harvested alfalfa or FM for 8 days. During the following 4 days, FM in the cages was replaced with freshly harvested sainfoin. Preference was greater (P<0.05) for FM offered with alfalfa scent than for FM offered with FM scent only on days 4 and 8. When lambs were offered FM with alfalfa or sainfoin in cages, they preferred (P<0.05) tall fescue with sainfoin scent over fescue with alfalfa scent, but intake was variable across hours and days (P<0.001). It is concluded that (1) lambs adjusted their intake of and preference for FA and FM over successive feeding bouts within each day, likely owing to an attempt to balance intakes of nutrients and alkaloids and (2) olfactory cues influenced preference, but to a lesser extent than nutrients and alkaloids in endophyte-infected tall fescue.

Importance of grass-legume choices on cattle grazing behavior, performance, and meat characteristics.

We determined if tall fescue in a mixture with either tannin-containing sainfoin or saponin-containing alfalfa affected cattle foraging behavior, performance, meat quality, and consumer acceptance of meat. Foraging behavior, BW, and pasture biomass before and after grazing were monitored when cattle strip-grazed 3 replications of 2 treatments from May through September 2010 (12 calves/replication) and from June through September 2012 (8 calves/replication). Animals were allowed a choice between tall fescue and sainfoin (SAN) or tall fescue and alfalfa (ALF) growing in strips (fescue, legume, and fescue-legume mixture). Incidence of use (scan samples) of legumes increased from the beginning to the end of the trials (P < 0.05), suggesting that cattle learned to mix legumes with tall fescue. Scan samples and assessments of pasture biomass removal revealed greater use of sainfoin than alfalfa when both legumes were abundant in the spring (P < 0.05); that pattern reversed toward the end of the growing season (P < 0.05) as the abundance of sainfoin declined more than that of alfalfa. Intake of forage per kilogram of gain was greater for SAN than for ALF in 2010 (P = 0.0003) and showed a tendency to be greater for SAN than ALF in 2012 (P = 0.19). There were no differences in ADG between SAN and ALF in either year (1 kg/d; P > 0.05). No incidences of bloat were detected. When cattle (3 calves/treatment in 2010 and 8 calves/treatment in 2012) were slaughtered in September to assess meat quality and consumer acceptance, carcasses were lean (4% to 6% fat content of the longissimus muscle), but compared with ALF, SAN had higher marbling scores, quality grades (Select versus Standard), and back fat thicknesses (P < 0.05) in 2012. Steaks from cattle finished on SAN were redder in color than steaks from cattle finished on ALF (P < 0.05; 2012). Meat samples from SAN and ALF showed some differences in unsaturated fatty acid profiles and volatiles concentrations (P < 0.05), but consumer acceptance did not differ between SAN and ALF (P > 0.05). Thus, cattle incorporated a lower-quality grass into their diets even when legumes were available ad libitum during most grazing trials. Choices between tall fescue and legumes differing in the type of secondary compound (tannins vs. saponins) and bloating potential influenced patterns of forage use by cattle and led to differences in meat quality.

Phytochemical complementarities among endophyte-infected tall fescue, reed canarygrass, birdsfoot trefoil and alfalfa affect cattle foraging.

We determined whether plant diversity and sequence of plant ingestion affected foraging when cattle chose from plants that varied in concentrations of alkaloids, tannins and saponins. We hypothesized cattle that ate high-alkaloid grasses (endophyte-infected tall fescue (TF) or reed canarygrass (RCG)) would prefer forages high in tannins (birdsfoot trefoil, BFT+) or saponins (alfalfa, ALF+), because tannins and saponins can bind to alkaloids, presumably reducing their absorption. We further hypothesized that forages with tannins or saponins consumed before, rather than after, foraging on high-alkaloid grasses would promote greater use of those grasses presumably by binding to alkaloids, thereby reducing their absorption. In Phase 1, cattle (n = 32) grazed on either high (+) or low (-) alkaloid grass (TF or RCG) pastures for 30 min each morning at 0600 h and were then offered a choice of BFT+, BFT-, ALF+ and ALF- for 60 min each day for 12 days. In Phase 2, cattle (n = 32) were first offered a choice of BFT+ or ALF+ for 30 min at 0600 h and then placed on grass (TF+ or -, or RCG+ or -) pastures for 60 min for 12 days. In both phases, we had four spatial replications of four treatments with 2 per calves assigned to each of the 16 replications per treatment combinations. Scan samples of individuals at 2-min intervals were used to determine incidence of foraging on each plant species (%). Cattle grazed more on RCG than on TF in Phases 1 (62% v. 27%; P = 0.0015) and 2 (71% v. 32%; P = 0.0005). In Phase 1, cattle that first foraged on RCG+ or TF- subsequently preferred ALF over BFT, whereas cattle offered RCG- or TF+ foraged on ALF and BFT equally. Foraging by cattle on RCG was cyclic during Phase 1, whereas cattle foraging on TF markedly decreased incidence of use of TF from 41% to only 16% by the end of the 12-day trial (P = 0.0029). Contrary to the cyclic (RCG) or steadily declining (TF) use of grasses in Phase 1, cattle steadily and dramatically increased foraging on both RCG and TF throughout Phase 2, when they first grazed BFT+ or ALF+ followed by high-alkaloid grasses (P = 0.0159). Our findings suggest that in plant species the sequence of ingestion influenced foraging behavior of cattle and that secondary compounds influenced those responses.

Early experience with diverse foods increases intake of nonfamiliar flavors and feeds in sheep.

This study determined whether early experiences by sheep with monotonous or diverse diets influence intake of unfamiliar flavors and feeds later in life. Thirty 2-mo-old lambs were randomly assigned to 3 treatment diets (n = 10): diverse (DIV), diverse with plant toxins (DIV+T), and monotonous (MON). Lambs in DIV received in 9 successive periods of exposure 4-way choice combinations of 2 foods high in energy and 2 foods high in protein from an array of 6 foods: 3 high in energy [beet pulp, oat grain, and a mix of milo:grape pomace (60:40)] and 3 high in digestible protein (DP) (soybean meal, alfalfa, corn gluten meal). Lambs in DIV+T received the same exposure as DIV, but 2 plant toxins, oxalic acid (1.5%) and quebracho tannins (10%), were randomly added to 2 of the feeds in each of the choice combinations. Lambs in MON received a monotonous balanced diet, made with a mixture of all 6 feeds detailed before. All treatments received their feed in 4 separate buckets. During exposure, treatments did not differ in total daily DMI (P = 0.31), but daily intake of ME was less (P < 0.02) and daily intake of DP was greater (P < 0.03) for lambs in DIV and DIV+T than for lambs in MON. Treatments did not differ in ADG or G:F (P > 0.05). After exposure, lambs were offered a familiar feed (wheat bran) containing novel flavors (maple, garlic, or bitter) and 2-way choices of novel feeds (fescue hay vs. corn distillers grains, rice vs. calf manna, and green peas vs. rolled oats). Intake of maple-flavored wheat bran tended (P = 0.08) to be greater for lambs in DIV than for lambs in DIV+T and MON. Intake of bitter-flavored and garlic-flavored wheat bran were greater (P = 0.03 and P = 0.04, respectively) for lambs in DIV and DIV+T than for lambs in MON. During 2-way choice trials, lambs in DIV, but not in DIV+T, showed greater intakes of fescue hay (P = 0.05) and rice (P = 0.04) than lambs in MON. Intake of green peas was greater (P = 0.03) for lambs in DIV and DIV+T than for lambs in MON. At the end of testing, lambs in DIV but not in DIV+T showed greater ADG than lambs in MON (P = 0.05). Thus, early exposure to diverse foods enhanced acceptance of novel flavors relative to early exposure to a monotonous ration. Early experience with diverse feeds plus plant toxins led to a less diverse diet than early experience with diverse feeds. Early exposure to diverse feeds may be beneficial in production systems that require rapid acceptance and high intake of unfamiliar feeds.

Preference for condensed tannins by sheep in response to challenge infection with Haemonchus contortus.

Herbivores prefer feeds that supply required nutrients and avoid those with excess nutrients and plant secondary compounds (PSC). Nevertheless, PSC such as condensed tannins can provide beneficial medicinal effects to herbivores as they act against infective diseases such as parasitism. The objectives of this study were to determine: (1) if parasitized lambs increased preference for a tannin-rich feed after they experienced the beneficial antiparasitic effects of condensed tannins relative to parasitized lambs that did not experience such benefits, and (2) if preference for the tannin-rich feed in the former group decreased when parasite burdens subsided. Twenty two lambs were familiarized with beet pulp and beet pulp+8% quebracho tannins (beet pulp+tannins) and choices were given between the two feeds (initial preference tests). Subsequently, all animals were dosed with 10,000 L(3) stage larvae of Haemonchus contortus. Twenty-two days later, animals were exposed to beet pulp (Control group; n=11) or beet pulp+tannins (Treatment group; n=11) during 24 d. After exposure (during a parasitic infection) animals in both groups were given choices between the two feeds. Lastly, animals in both groups received an antiparasitic drench and were again given a choice between both feeds (after a parasitic infection). Lambs preferred beet pulp to beet pulp+tannins throughout the study (P<0.001) and no difference in preference for the tannin-rich feed was detected between groups during initial preference tests (P>0.05). However, during a parasitic infection, intake of and preference for the tannin-rich feed was higher for lambs that experienced the beneficial effects of condensed tannins while parasitized (Treatment) than for lambs that did not (Control) (P<0.05). When parasitic infections were terminated by chemotherapy, differences between groups disappeared (P>0.05). Preference by the Treatment group for the tannin-rich feed was lower after than during a parasite infection (P<0.05). In contrast, preference by the Control group did not change during these periods (P>0.05). Lambs in the Treatment group displayed lower FEC than lambs in the Control group (Group × Sampling Date; P<0.05). These results show lambs learned about the antiparasitic effects of condensed tannins and increased their preference for the tannin-rich feed, which subsided after chemotherapy, indicating preference was due to parasite infection. These findings suggest management that allows animals to select tannin-rich feeds can enable parasitized animals to self-medicate.

Cattle preferences differ when endophyte-infected tall fescue, birdsfoot trefoil, and alfalfa are grazed in different sequences.

We determined if sequence of ingestion affected use of endophyte-infected tall fescue (TF) when cattle also grazed birdsfoot trefoil (BFT) or alfalfa (ALF). Based on chemical characteristics of TF (alkaloids), BFT (tannins), and ALF (saponins), we hypothesized that cattle first allowed to graze ALF or BFT would subsequently spend more time grazing TF than cattle that first grazed TF followed by ALF or BFT. Sixteen bred heifers (478 ± 39 kg initial BW) were randomly assigned to 4 replicated pasture units. Each replicated unit consisted of 4 treatment sequences (TF → BFT, TF→ALF, BFT → TF, or ALF→TF), with 2 cows per sequence. Pastures were in the vegetative stage of growth at a height of 20 to 30 cm and provided ad libitum forage to cattle. We recorded foraging on TF, BFT, and ALF using scan sampling of individuals at 2-min intervals. The study was conducted in 4 phases run sequentially, for a total of 30 d. In phases 1 and 3, cattle in group 1 grazed TF pastures for 45 min and were then moved to BFT pastures for the next 45 min (TF→BFT); cattle in group 2 grazed in the reverse sequence (BFT → TF). In phases 2 and 4, cattle in group 1 grazed TF pastures for 45 min and then subsequently grazed ALF pastures for the remaining 45 min (TF→ALF); cattle in group 2 grazed in the reverse sequence (ALF→TF). Sequence of plant ingestion affected food selection. In phase 1, scans revealed grazing of TF by heifers was cyclic, and heifers tended to have more scans (P = 0.52) grazing TF when they grazed BFT → TF; scans for heifers grazing TF were consistently greater (P < 0.05) throughout phase 3 of the trial. In phase 2, heifers that grazed in the sequence ALF→TF spent considerably more scans (P = 0.03) foraging on TF from d 4 to 10 than heifers that grazed in the sequence TF→ALF, and they remained greater throughout phase 4 of the trial. Although the sequence ALF→TF appeared to be more effective than BFT → TF, consistent with the hypothesis of a complementary relationship between the steroidal alkaloids in TF and saponins in ALF, tannin concentrations in BFT were minimal (1.8%), which likely reduced the presumed inactivation of alkaloids by tannins. We also speculate that heifers needed to learn about the positive postingestive influence of sequence, a notion consistent with more similar scans spent foraging BFT and TF early in phases 1 (BFT → TF) and 2 (ALF→TF), and with the consistent and marked increase in scans spent foraging on TF for animals foraging in phases 3 (BFT → TF) and 4 (ALF→TF).

Selection of tannins by sheep in response to gastrointestinal nematode infection.

Herbivores learn to select compounds that attenuate the aversive effects of plant secondary metabolites (PSM), but can they increase intake of PSM they typically avoid when these PSM provide medicinal effects? We hypothesized that herbivores learn to increase intake of PSM-containing feeds when experiencing a gastrointestinal parasitic infection. Ten lambs with natural gastrointestinal parasitic burdens (PB) and 10 nonparasitized lambs (NP) were offered a choice of alfalfa (Medicago sativa) and alfalfa mixed with 10% quebracho tannin (Schinopsis quebracho-colorado; alfalfa:tannins) before and after they were conditioned with the postingestive effects of tannins. Preference for alfalfa:tannins did not differ between groups before experiencing the postingestive effects of tannins (P = 0.85) or when parasite loads were terminated due to the administration of ivermectin (P = 0.18). In contrast, when tested with a parasite burden, lambs in PB consumed more alfalfa:tannins (P = 0.08), showed greater preference for alfalfa:tannins (P = 0.07), and consumed less alfalfa than lambs in NP (P = 0.06). Ingestion of tannins by lambs in PB was followed by reduced fecal egg counts (FEC; P = 0.006), and there was a direct proportional relationship between preference for alfalfa:tannins and FEC (P = 0.07). In summary, parasitized lambs increased their intake of alfalfa:tannins when they experienced a parasite burden, which suggests they self-medicated with tannins against parasites. Self-selection of PSM has implications for the quest for alternatives to chemoprophylaxis in the treatment and well-being of parasitized wild and domestic animals grazing in pasturelands and in confinement.

Links between ruminants' food preference and their welfare.

Nutrition is an important aspect of welfare, and in most recommendations for the welfare of animals adequate nutrition is a primary requirement. However, in intensive livestock production systems the decision for adequate nutrition is made based on traditional paradigms of feeding monotonous rations or plant monocultures, frequently with excesses or imbalances of nutrients relative to the individuals' physiology, which can compromise welfare. Individual ruminants can better meet their needs for nutrients and regulate their intake of secondary compounds when offered a variety of foods than when constrained to a single food, even if the food is nutritionally balanced. The concept of food variety is central because monotonous flavors and feeds and excess nutrients all cause animals to satiate, which in turn causes animals to eat a variety of foods. When offered a variety of foods, satiety for single foods stimulates the selection of a diverse diet and thus food intake, but when restricted to a monotonous diet satiety is aversive and limits food intake. Moreover, if a monotonous diet is aversive to animals then this could be stressful, even if monotony implies consuming a balanced diet. A diverse diet may also increase resistance to disease in ruminants, by allowing consumption of small amounts of compounds with antimicrobial/antiparasitic effects and immunity-enhancing properties. Herbivores also experience the benefits of ingesting compounds with medicinal (i.e. antiparasitic) benefits and they learn to prefer foods containing such compounds as their preferences are associatively conditioned by the food's homeostatic utility to the body. Such learned patterns of behavior begin in utero and feeding experiences early in life cause changes - neurological, morphological and physiological - in animals, which influence on their subsequent behavior and welfare. Such experiences with the environment enable animals to adapt to local diets and stressors and reduce the levels of fear. Finally, feeding behavior in farm animals could be an aid in the early detection and mitigation of pain or sickness, and as such become an important tool in the identification of welfare and health of animals before the appearance of clinical signs. Management strategies in ruminant production systems could benefit by allowing animals to manifest their feeding preferences, thereby acknowledging the animals' role as active players in feeding systems, instead of regarding them as passive entities that just respond to prescriptions and formulations.

Neem-tree (Azadirachta indica Juss.) extract as a feed additive against the American dog tick (Dermacentor variabilis) in sheep (Ovis aries).

Acaricides can be conveyed to ticks via the blood of their hosts. As fruit and kernel extracts from the Meliaceae family, and, in particular the tetranortriterpenoid azadirachtin (AZA) inhibits tick egg production and embryogenesis in the Ixodidae ticks, we investigated the effects of Neem Azal, an extract containing 43% AZA, given as a feed additive to lambs artificially infested with engorging adult Dermacentor vairiabilis ticks. After tick attachment, the lambs were allotted to three dietary treatments: AZA0 (control, n=10), AZA0.3 (n=5), and AZA0.6 (n=5), with feed containing 0%, 0.3%, and 0.6% AZA on DM basis, respectively. In half of the AZA0 lambs, ticks were sprayed on day 4 after attachment with an ethanol:water:soap emulsion containing 0.6% AZA (AZA0S). In spite of its very pungent odor, the neem extract was well accepted by all but one lamb. No differences were found between treatment groups in liver enzymes in blood, and there was no indication of toxicity. The plasma AZA concentrations after 7 and 14 days of feeding AZA were (4.81 and 4.35 microg/mL) for the AZA0.6 and (3.32 and 1.88 microg/mL) for the AZA0.3 treatments, respectively (P<0.0001). Treatments were not lethal to ticks, but tick weights at detachment were 0.64, 0.56, 0.48, and 0.37 g for ticks from the AZA0, AZA0.3, AZA0S, and AZA0.6 treatments (P<0.04), respectively, suggesting that blood AZA impaired blood-feeding. The highest mortality rate after detachment was for AZA0.6 (P<0.09). As AZA affects embryo development and ticks at the molting stages, we expect that following treatments of hosts for longer periods, one-host ticks will be more affected than the three-host tick D. variabilis.

Rumen distension and contraction influence feed preference by sheep.

Distension of the rumen limits feed intake by livestock. Ruminal dysfunctions due to bloat, which causes distension by accumulation of excessive gas within the rumen, also reduce feeding. We hypothesized that excessive levels of rumen distension cause feed aversions and that preference increases for feeds eaten in association with recovery from bloat. To test these hypotheses, we determined whether 12 commercial crossbred lambs (average initial BW of 43 +/- 2 kg) could associate ingestion of specific feeds with the consequences of increased intraruminal pressure and its subsidence. Six of the lambs were fitted with rumen cannulas and offered ground alfalfa for 30 min after a rubber balloon was inserted into the rumen of each animal and distended with air to volumes of 1.8, 2.5, or 4.5 L. Subsequently, balloons were deflated and alfalfa was offered again for a second period of 30 min. Feed intake was not affected when the balloon was not distended (P = 0.45 to 0.93), but distension reduced feed intake (P < 0.001) in direct proportion to the magnitude of distension at all 3 volumes (R(2) = 0.70). Relief from distension promoted a compensatory increase in feed intake (P = 0.006). During conditioning to determine if lambs acquired a preference for a feed associated with recovery from distension, fistulated lambs were offered novel feeds: wheat bran (group 1; n = 3) and beet pulp (group 2; n = 3), and the balloon was distended for 30 min. Feeds were then switched and the balloons were deflated (recovery). Control lambs (n = 6) received the same feeding protocol without the balloons. Lambs formed strong aversions to feeds associated with distension and preferred feeds associated with recovery (P = 0.001 to P = 0.10). No preferences or avoidances were observed in control lambs conditioned without rumen distension (P = 0.17 to P = 0.87). Thus, rumen distension and recovery from distension induced feed aversions and preferences, respectively, which may be critical in learning avoidance of bloat-inducing plants and preferences for plants and supplements that relieve the incidence of bloat.

Supplemental polyethylene glycol affects intake of and preference for sericea lespedeza by cattle.

We tested the hypothesis that supplemental polyethylene glycol (PEG), a polymer that neutralizes the effects of tannins, would increase intake and preference of cattle for fresh-cut sericea lespedeza (SL; Lespedeza cuneata), a legume with high concentrations of condensed tannins. Sixteen crossbred steers (initial BW of 249 +/- 6.6 kg) were randomly assigned to 2 treatments: 1) grain supplement with PEG (PEG-S) and 2) grain supplement without PEG (control). To assess intake, we conducted a trial with 6 sample periods, each 6 d in duration, with steers fed SL and prairie hay (PH) in separate meals. Steers were fasted overnight and fed their respective supplements (with and without PEG) at 0800 h. Animals were then offered fresh-cut SL from 1050 to 1550 h, PH was fed from 1600 to 2000 h, and they were without food from 2000 to 0800 h the next day. To assess preference, we conducted 1-d tests in which steers had simultaneous access to SL and PH on the day following periods 2 to 6. As with the intake experiment, steers were fasted overnight and fed their respective supplements at 0800 h, but from 1050 to 1250 h all steers had access to SL and PH simultaneously. We weighed steers before and at the completion of the study and calculated ADG. Steers treated with PEG consumed more SL per unit of BW than control steers in periods 2 to 6 (period x treatment interaction, P < 0.001). In contrast, controls consumed more PH than steers given PEG (period x treatment x day interaction, P = 0.009). The PEG-S steers consumed more total DM (SL + PH) than controls in periods 3, 5, and 6 but not in periods 1, 2, and 4 (period x treatment interaction, P = 0.004). Sericea lespedeza intake as a percentage of total DMI was greater for the PEG-S steers every day except d 1 and 2 of period 1 (period x treatment x day interaction, P = 0.03). Averaged across the 5 preference tests, PEG-S steers selected a greater proportion of SL than did control steers (39 vs. 9%), and the magnitude of the difference was greater in the later tests (test x treatment interaction, P = 0.004). The PEG-S steers had greater ADG than controls (0.44 vs. 0.24 kg/d; P = 0.005). Our results indicate PEG increases intake of and preference for SL and suggest that PEG supplementation of cattle may increase intake of SL and improve ADG in pastures that contain SL.

The effect of polyethylene glycol on intake of Mediterranean shrubs by sheep and goats.

Poor nutritional quality and increased content of secondary compounds can reduce consumption of Mediterranean shrubs by herbivores. In 2 sequential trials, we examined the effect of polyethylene glycol (PEG) and number of shrub species offered on daily intake of Mediterranean shrubs by 12 sheep and 12 goats. The PEG (25 g) was fed to experimental animals with barley. In trial 1 (6 shrubs), goats ate more (P = 0.0008) daily total shrub biomass than did sheep (60.7 vs. 45.9 +/- 2.6 g/kg of BW). There was a trend (P = 0.08) toward a positive PEG effect on total shrub intake, with PEG-supplemented animals consuming more total shrubs than controls (56.7 vs. 50.0 +/- 2.6 g/kg of BW). Trial 2 (using 3 shrubs) was a continuation of trial 1, except that animals were given less barley and treatment animals were given more PEG (50 g). Both sheep and goats showed a numerical decrease in total shrub intake from trial 1 to trial 2. Sheep receiving PEG ate more (P = 0.002) total shrubs than did controls, but no PEG effect was found for goats. Thus, PEG had a greater influence on sheep than goats when only 3 shrubs were offered, a result that may be related to the fact that fewer shrubs with complementary secondary compounds were offered and that goats appear to have a greater ability to consume and detoxify secondary compounds from Mediterranean shrubs. Overall, as the number and diversity of shrubs offered increased, supplemental PEG had less effect on increasing intake for both goats and sheep.

Learned appetites for calcium, phosphorus, and sodium in sheep.

If supplemental minerals are needed to promote optimal animal performance, what is the best way of providing them: free choice or in the diet? We hypothesized that herbivores discriminate among feeds containing Na, P, and Ca and modify their choices as a function of need. One group of lambs was fed a basal diet low in P and high in Ca (low P-high Ca), whereas another group was fed a basal diet high in P and low in Ca (high P-low Ca). After 73 d of exposure to the unbalanced diets, the lambs were conditioned by offering flavored grape pomace containing NaCl, CaCO(3), or NaH(2)PO(4). Preference for pomace + minerals was determined when all lambs were fed a basal diet of alfalfa pellets and barley grain (initial preference) and during 4 phases. Phases 1 and 2 occurred after 40 and 67 d of feeding the unbalanced basal diets, phase 3 occurred after conditioning with NaCl, CaCO(3), or NaH(2)PO(4), and phase 4 occurred 22 d after the groups were moved to 2 new (separate) locations so the animals in the different groups could not eat dirt, urine, or feces from the other pen. Preference for pomace did not differ between the groups during the initial preference tests (P = 0.62); both groups preferred NaCl > CaCO(3) = NaH(2)PO(4) (P < 0.001). As the study progressed, and lambs fed low P-high Ca had lower P and greater Ca concentrations in serum than lambs fed high P-low Ca (P < 0.001), the preference between groups diverged. In phase 2, lambs in high P-low Ca continued to prefer NaCl (P < 0.001), but lambs in low P-high Ca preferred NaH(2)PO(4) (P < 0.05). After conditioning, both groups preferred NaCl = NaH(2)PO(4) > CaCO(3) (P < 0.01 to 0.11). After the groups were moved to different locations, lambs fed low P-high Ca showed the lowest concentration (3.7 mg/dL) of inorganic P in serum for all phases (P < 0.001), and they preferred NaH(2)PO(4) > NaCl = CaCO(3) (P < 0.001). In contrast, lambs in high P-low Ca avoided NaH(2)PO(4) (P < 0.05). Lambs offered high P-low Ca showed a greater preference for CaCO(3) (P = 0.12) and NaCl (P < 0.05) and a lower preference for NaH(2)PO(4) compared with lambs fed low P-high Ca (P < 0.001). In summary, lambs discriminated among different flavored feeds containing NaCl, CaCO(3), and NaH(2)PO(4) and displayed preferences as a function of the mineral imbalance in their basal diets. Thus, it may be possible to feed Ca and P supplements free choice, such that individual animals within a group can manifest preferences based on their specific needs.

What does it mean to be locally adapted and who cares anyway?

The availability of fossil fuels will likely decline dramatically during the first half of the twenty-first century, and the massive deficits probably will not be alleviated by alternative sources of energy. This seeming catastrophe will create opportunities for communities to benefit from foods produced locally in ways that nurture relationships among soil, water, plants, herbivores, and people to sustain their collective well being. Agriculture will be much more at the heart of communities than it is currently, but by necessity, it will no longer be so dependent on fossil fuels to power machinery or to produce fertilizers, herbicides, and insecticides to grow and protect plants in monocultures, antibiotics, and anthelmintics to maintain the health of herbivores, or nutritional supplements and pharmaceuticals to sustain humans. Rather, from soils and plants to herbivores and people, we will have to learn once again what it means to be locally adapted to the landscapes we inhabit. In the process of relearning these skills, plants will become more important as nutrition centers and pharmacies, their vast arrays of primary (nutrients) and secondary (pharmaceuticals) compounds useful in nutrition and health. There also will be a need, as in times past before our heavy reliance on fossil fuels, to produce livestock in easy-care systems that match seasonally-available forages with production needs and that match animals anatomically, physiologically, and behaviorally to local landscapes. This will mean reducing inputs of fossil fuels to increase profitability by 1) matching animal needs to forage resources; 2) selecting for animals that are adapted anatomically, physiologically, and behaviorally to local environments; 3) culling animals unable to reproduce with minimal help from humans; and 4) creating grazing systems that enhance the well-being of soils, plants, herbivores, and people.

Self-medication and homeostatic behaviour in herbivores: learning about the benefits of nature's pharmacy.

Traditional production systems have viewed animals as homogeneous 'machines' whose nutritional and medicinal needs must be provided in a prescribed manner. This view arose from the lack of belief in the wisdom of the body to meet its physiological needs. Is it possible for herbivores to select diets that meet their needs for nutrients and to write their own prescriptions? Our research suggests it is. Herbivores adapt to the variability of the external environment and to their changing internal needs not only by generating homeostatic physiological responses, but also by operating in the external environment. Under this view, food selection is interpreted as the quest for substances in the external environment that provide homeostatic utility to the internal environment. Most natural landscapes are diverse mixes of plant species that are literally nutrition centres and pharmacies with vast arrays of primary (nutrient) and secondary (pharmaceutical) compounds vital in the nutrition and health of plants and herbivores. Plant-derived alkaloids, terpenes, sesquiterpene lactones and phenolics can benefit herbivores by, for instance, combating internal parasites, controlling populations of fungi and bacteria, and enhancing nutrition. Regrettably, the simplification of agricultural systems to accommodate inexpensive, rapid livestock production, coupled with a view of secondary compounds as toxins, has resulted in selecting for a biochemical balance in forages favouring primary (mainly energy) and nearly eliminating secondary compounds. There is a global need to create a more sustainable agriculture, with less dependence on external finite resources, such as fossil fuels and their environmentally detrimental derivatives. Self-medication has the potential to facilitate the design of sustainable grazing systems to improve the quality of land as well as the health and welfare of animals. Understanding foraging as the dynamic quest to achieve homeostasis will lead to implementing management programs where herbivores have access not only to diverse and nutritious foods but also to arrays of medicinal plants.

Terpenes and carbohydrate source influence rumen fermentation, digestibility, intake, and preference in sheep.

We hypothesized that toxins and nutrients in foods interact to influence foraging behavior by herbivores. Based on this hypothesis we predicted that 1) terpenes in big sagebrush (Artemisia tridentata) influence intake and preference in sheep for diets varying in sources of nonstructural (barley grain) and structural (sugar beet pulp) carbohydrates, and 2) these effects are due to the differential effects of terpenes on fermentation products and apparent digestibility of each class of carbohydrates. Lambs were fed 2 isoenergetic and isonitrogenous diets with varying proportions of the same ingredients (beet pulp- and barley grain-based diet) or offered a choice between the 2 diets; all feeds were fed without and with terpenes, in consecutive periods. We also compared intake and preference of the beet pulp-and barley-based diets before and after the lambs ate a meal of sagebrush. Finally, we assessed the effect of terpenes on ruminal variables and in vivo digestibility. Lambs ate less when fed beet pulp or when they were offered a choice of diets with terpenes (P < 0.001), and intake of the beet pulp-based diet was the most affected (P < 0.05). Lambs preferred the beet pulp-to the grain-based diet with terpenes, but their preference reversed when terpenes were removed from the diets (P < 0.05). When lambs were offered both diets, intake and preference did not differ (P > 0.20) before eating sagebrush, but they preferred the beet pulp-based diet after eating sagebrush (P < 0.05). Intake of sagebrush did not differ among groups consuming the test diets (P = 0.21). Addition of terpenes to both diets increased the digestibility of DM, NDF, and ADF and decreased concentrations of total VFA and acetate (P < 0.05). Terpenes also depressed butyrate concentration in the barley-based diet (P < 0.05). Propionate concentrations were not affected by terpenes in either feed (P = 0.63). In summary, the predominant type of feed ingredient (beet pulp, grain) ingested with terpenes influenced fermentation products, intake, and preference in lambs. The source of energy from supplements, or other plants in the diet, is likely to influence intake and preference for sagebrush in sheep foraging on rangelands. Moreover, ingesting terpenes from sagebrush may also influence intake and preference for other plant species or supplements.

Phosphorus appetite in sheep: dissociating taste from postingestive effects.

We hypothesized that lambs discriminate the postingestive effects of P and associate those effects with feed flavor to modify feed choices. Three predictions were tested based on this hypothesis: 1) lambs will modify preference for arbitrary flavors eaten during intraruminal infusions of NaH2PO4, 2) changes in preferences are more specific than changes in osmotic load induced by salts of Na; and 3) preference for P is inversely related to the concentration of inorganic P in blood. Thirty lambs were depleted of P by the offer of a P-deficient diet, allocated to 3 groups (10 lambs/group), and conditioned during 3 periods as follows: During conditioning period 1, lambs in each of 3 groups ate a poorly nutritious feed (grape pomace), flavored differently for each group, while water was infused into the rumen. During conditioning periods 2 and 3, lambs again ate grape pomace, with 2 new flavors now paired with infusion of an aqueous solution (126 mmol) of NaCl (conditioning period 2) or NaH2PO4 (conditioning period 3), rather than with water. After conditioning, all lambs were offered a choice of the 3 flavors during preference tests immediately after conditioning (period 1) and every 3 wk thereafter (periods 2, 3, and 4). During period 1, when serum inorganic P levels were greatest, lambs preferred flavors paired with water > NaCl > NaH2PO4 (P < 0.05). During periods 2 and 3, as inorganic P concentrations decreased in serum, lambs preferred flavors paired with NaH2PO4 > NaCl (period 2, P = 0.10; period 3, P = 0.05). Lambs preferred flavors paired with water > NaH2PO4 in period 2 (P < 0.001), but those differences disappeared in periods 3 and 4 (P > 0.05). During period 4, lambs preferred flavors paired with NaCl > NaH2PO4 (P < 0.10). The estimate of the slope for the linear relationship between intake of flavors paired with NaH2PO4 and serum inorganic P was negative (P < 0.0001), whereas estimates of the slopes for the relationships between intake of flavors paired with NaCl or water and serum inorganic P were not different from 0. Thus, preference for P was inversely related to the concentration of serum inorganic P. Our results suggest lambs discriminated among the postingestive effects of NaH2PO4, NaCl, and water and associated those effects with specific flavors. Lambs avoided flavors paired with NaH2PO4 during periods of P replenishment, and they increased preference for those flavors during periods of P need.

Influence of macronutrients and activated charcoal on intake of sagebrush by sheep and goats.

We determined if supplemental macronutrients or activated charcoal influenced intake of sagebrush, a shrub that contains terpenes. Sheep (lambs 3 mo age, 33 kg) and goats (kids 5 mo age, 28 kg) were fed supplements high in energy, high in protein, or offered a choice between the two supplements before and after receiving sagebrush for 4 h/d. The effect of activated charcoal, a compound that reduces bioavailability of toxins, was assessed by offering charcoal mixed with a carrier or the carrier alone while animals consumed sagebrush for 4 h/d. The effects of supplemental macronutirents were tested before and after charcoal supplementation. Activated charcoal did not increase intake of sagebrush by sheep or goats (P > 0.05), but supplemental macronutrients influenced intake of sagebrush by both species (P < 0.05). Intake of sage-brush was markedly higher for animals fed the high-protein supplement than for animals fed the high-energy supplement (P < 0.05). Lambs (before and after charcoal supplementation) and kids (after charcoal supplementation) offered a choice between the two supplements used sagebrush at levels comparable to animals supplemented with protein. They consumed more CP than animals fed the high-energy supplement and more ME than lambs fed the high-protein supplement (P < 0.05). Thus, providing a choice between supplements enhanced intake of sagebrush and enabled animals to achieve a more balanced intake of macronutrients. Preference for supplement and sagebrush also depended on an animal's immediate past history with a supplement. Sheep and goats previously fed the high-energy or the high-protein supplement preferred, respectively, the high-protein or the high-energy supplement when given a choice (P < 0.05). The changes in preference for supplement caused a decreased intake of sagebrush by groups that preferred the energy concentrate, and an increased intake of sagebrush by groups that preferred the protein concentrate. Thus, supplements of different energy and protein densities consistently altered intake of sagebrush by sheep and goats.

Polyethylene glycol influences selection of foraging location by sheep consuming quebracho tannin.

Tannins are a heterogeneous group of phenolic polymers that can induce detrimental effects when consumed by herbivores. Polyethylene glycol (PEG) binds to tannins and thus attenuates their negative effects. Our objective was to determine whether sheep actively seek PEG when fed tannins and thus modify their foraging location as a function of the spatial distribution of PEG. Lambs were first trained to recognize the beneficial effects of PEG by offering a meal high in quebracho tannin (QT), which presumably caused malaise, and then PEG (MW, 3,350), which presumably led to recovery from malaise. Animals were then tested in an experimental area where they could forage at two different locations that contained in Trial 1 1) PEG and QT in adjacent food boxes (PEG+QT) or 2) QT and in Trial 2 1) PEG or 2) QT. Preference for foraging locations was tested under three conditions: 1) no preload meal, 2) a meal high in quebracho tannin 1 h before testing, and 3) a basal diet high in quebracho tannin. Lambs spent more time (P = 0.012) and ate more tannin-containing food (P = 0.022) at locations where PEG was present (PEG+QT) than where it was absent (QT; Trial 1). Lambs responded to increased tannins in their bodies (Conditions 2 and 3) by reducing intake (P = 0.0001; Trial 2) at sites containing only QT and by increasing intake (P = 0.0001, Trial 1; P = 0.001, Trial 2) and foraging time (P = 0.072, Trial 1; P = 0.0001, Trial 2) at locations where PEG was available. Thus, PEG influenced foraging location by sheep consuming quebracho tannin. Accordingly, it may be possible to formulate range blocks that enable herbivores to ingest PEG in tannin-rich habitats. Strategic distribution of PEG in those habitats may attract animals to underutilized feeding sites.