Diet selection and performance of horses grazing on different heathland types.

The number of horses in northern Spanish mountains has increased in recent decades, but little is known about their grazing behaviour, performance and potential for foal meat production. This research aimed to study the diet selection, liveweight (LW) changes and parasitic status of dry and lactating mares, and foals' LW gains, grazing on heathlands with different botanical composition. The experimental design consisted of three vegetation types: dominated by heather (Ericaceae) species (H), dominated by gorse (Ulex gallii; G) and co-dominated by gorse and heath-grasses (G-G), with four replicates per treatment (12 paddocks of 1.2 ha). The study lasted three grazing seasons (2010-12). Each year, 24 crossbred mature mares (310±52 kg LW) were used, managing one lactating mare with her foal plus one non-lactating mare per paddock from May to late summer or early autumn. In the case of H paddocks, animals had to be removed before (late August to early September) because of apparent loss of body condition. Animals were periodically weighed. Mares' diet composition was estimated using alkane markers, analysing the discrepancies in alkane concentrations between dietary plant components and faeces. Faecal samples were also analysed for gastrointestinal nematodes ova. Chemical composition of the main plant components (i.e. heather, gorse and grasses) revealed a low nutritive value, averaging 79, 115 and 113 g CP/kg dry matter (DM), respectively, that could restrict livestock performance. Mares initially selected gorse and grasses (0.47 and 0.40, respectively, in 2010), increasing heather consumption over time (from 0.13 in 2010 to 0.29 in 2012) as gorse availability decreased. The performance of both mares and foals was lower in H compared with G and G-G paddocks (-216 v. 347 g/day for mares, P<0.01; 278 v. 576 g/day for foals, P<0.05), whereas LW changes were more favourable in dry mares than in lactating ones (241 v. 78 g/day; P<0.05). Small strongyle (Cyathostominae) egg counts in mares' faeces increased across the grazing season with no differences between treatments. These results indicate that grazing by horses on gorse- and grass-gorse-dominated shrublands could be sustainable at least during part of the year (4 to 6 months). However, heather-dominated heathlands are not able to meet the nutritional needs of horses even for a short time (2 to 4 months). Nevertheless, the low nutritive quality of these vegetation communities, especially in autumn, requires animal access to other pastures with a higher nutritive value, or supplementary feeding, to enhance foals' growth and maintain sustainable grazing systems with productive herds.

Productive responses of breeding Cashmere goats and their kids to different stocking rates on improved upland pastures.

Although goat meat production could be an option for diversification in improved upland pastures in northern Spain, precise information on the optimal grazing management to enhance goat performance and maximize production per unit land area is lacking. The objective of this study was to compare the effects of 3 stocking rates, high stocking rate (HSR; 20 goats/ha), medium stocking rate (MSR; 15 goats/ha), and low stocking rate (LSR; 10 goats/ha), on gastrointestinal (GI) nematode infections and productive responses of Cashmere goats grazing such pastures. Treatments were replicated twice on 6 paddocks sown with and and with a high presence of the native grass . The experiment lasted 3 grazing seasons (from spring to autumn). Pastures were sampled for sward height and botanical and proximate composition. Body weight and BCS changes of goats were monitored and GI nematode infections were assessed by fecal egg counts (FEC). The established treatments resulted in lower mean sward height in the HSR than in the MSR and LSR (9.6, 11.5, and 14.4 cm, respectively; < 0.001). Pasture botanical composition and nutritive quality did not differ between treatments. The mean FEC of does across the 3 grazing seasons were greater ( < 0.05) in the HSR than in the LSR. spp., , and were the most prevalent nematode species identified in coprocultures. Does showed more favorable ( < 0.001) BW and BCS changes in the LSR than in the MSR and HSR (-14, -30, and -52 g/d and -0.1, -0.3, and -0.7 BCS units [scale 1 to 5], respectively). Greater ( < 0.001) kids' BW gains were observed in the LSR and MSR (average 94 g/d) compared with the HSR (70 g/d). Inversely, kid output per unit land area was greater in the HSR than in the MSR and LSR (320, 258, and 192 kg∙ha∙yr, respectively; < 0.001), whereas daily kids' BW gains per hectare were greater ( < 0.001) in the HSR and MSR (average 1.37 kg∙d∙ha) compared with the LSR (0.98 kg∙d∙ha). A medium stocking rate of 15 goats/ha could represent the best compromise between animal health, performance, and productivity per unit land area in this type of upland pastures, but stricter controls of parasite levels during the grazing season would be necessary to avoid production losses, unless alternative nutraceuticals are provided.

Anthelmintic effect of heather in goats experimentally infected with Trichostrongylus colubriformis.

The effects of heather (composed primarily of Calluna vulgaris with a smaller content of Erica umbellata and Erica cinerea) consumption on the establishment of incoming infective larvae (experiment 1, preventive treatment) and an adult worm population (experiment 2, curative treatment) were investigated in Cashmere goats experimentally infected with Trichostrongylus colubriformis. In experiment 1, 12 castrated male goats were divided into two groups: heather-supplemented vs. non-supplemented animals. After 2 weeks of adaptation to the diet, all goats were experimentally infected per os with 6,000 T. colubriformis third-stage larvae. Three weeks post-infection, the goats were slaughtered, and worm counts as well as female worm fecundity and development were determined. Heather consumption was associated with a close to significant (P=0.092) reduction (mean 14 %) in larvae establishment. No effect on fecundity was observed, but the length of female worms in supplemented goats was greater (P<0.001). In experiment 2, 15 non-lactating does were experimentally infected with 6,000 T. colubriformis third-stage larvae. At 6 weeks post-infection, three groups were established: control, heather-supplemented and heather-supplemented with polyethylene glycol. Individual faecal nematode egg output was measured twice weekly to assess gastrointestinal nematode egg excretion. The goats were slaughtered 5 weeks after heather administration (11 weeks post-infection), and worm counts as well as female worm fecundity and development were subsequently determined. Heather administration was associated with a significant (P<0.001) decrease (between 47 and 66 % compared with control group) in egg excretion from 45 to 76 days post-infection. Although worm counts and female fecundity were lower in supplemented goats, no significant differences were observed. Overall, the results showed a reduction in T. colubriformis larvae establishment and a decrease in nematode egg excretion when heather was administered in experimentally infected goats. The heather plus polyethylene glycol treatment reduced nematode egg excretion levels at the same proportion as heather, thereby suggesting that the threshold of tannins required for an anthelmintic effect is most likely quite low.

In vitro effect of heather extracts on Trichostrongylus colubriformis eggs, larvae and adults.

This study was carried out to evaluate the in vitro effects of different heather species on Trichostrongylus colubriformis eggs, larvae and adult worms, and obtain scientific evidence to attribute these effects to the action of their phenolic compounds and/or tannins. Total phenolic extracts of three heather species (Calluna vulgaris, Erica cinerea, and Erica umbellata) and an equal mixture of these three extracts were tested in vitro in the three development stages of T. colubriformis using an egg hatching assay (EHA), larval exsheathment inhibition assay (LEIA), and adult motility inhibition assay (AMIA). The egg hatching rate was measured after incubation with heather extracts for 48 h at 25°C. Infective third-stage larvae (L3) were incubated for 3h at 25°C with heather extracts. The evolution of artificial exsheathment over time was measured with repeated observations at 20-min intervals for 60 min. Adult worms were obtained from one donor goat and incubated with the extracts at 37°C for 5 days in 48-multiwell plates. Worm motility was measured at 0, 19, 24, 43, 48, 67, 72, 96 and 115 h after the beginning of the experiment. The extracts were tested at concentrations of 75, 150, 300, 600 and 1,200 µg/ml. All extracts significantly (P<0.001) inhibited egg hatching and the effect was dose dependent. All extracts inhibited or delayed the exsheathment of T. colubriformis L3, and the effect was dose dependent for C. vulgaris. Incubation with heather extracts induced a reduction in adult worm motility compared to control, although significant (P<0.05) differences were only found at the highest concentrations. Additional studies showed that purified tannins of the same heather species disturbed T. colubriformis larval exsheathment. All these results confirm the anthelmintic properties of heather against T. colubriformis, and suggest that not only tannins but also some other phenolic compounds might be involved.

In vitro effect of heather (Ericaceae) extracts on different development stages of Teladorsagia circumcincta and Haemonchus contortus.

The aim of the present study was to evaluate the in vitro effects of heather (Ericaceae) phenolic extracts on the abomasal nematodes Teladorsagia circumcincta and Haemonchus contortus. Extracts of three heather species (Calluna vulgaris, Erica cinerea, Erica umbellata and a balanced mixture of all three) were tested in vitro on different development stages of T. circumcincta (eggs, infective larvae and adult worms) and H. contortus (eggs and infective larvae) using an egg hatching assay (EHA), a larval exsheathment inhibition assay (LEIA) and an adult motility inhibition assay (AMIA). The egg hatching rate was measured after incubation with heather extracts for 48 h at 25 °C. Ensheathed infective larvae were incubated for 3h at 20 °C with heather extracts. Artificial exsheathment was induced in vitro by adding hypochloride solution to the larval suspension. The progress of exsheathment over time was measured by repeated observations at 10-min (T. circumcincta) and 20-min (H. contortus) intervals for 60 min. Adult T. circumcincta worms were obtained from two donor goats and incubated with the extracts at 37 °C for 3 days in 48-well multiwell plates. Worm motility was measured at 0, 19, 24, 43, 48, 67 and 72 h. The extracts were tested at concentrations of 75, 150, 300, 600 and 1200 µg/mL. Incubation with E. cinerea, E. umbellata and mixed heather extracts had a significant (P<0.01) dose-dependent effect on T. circumcincta egg hatching. H. contortus egg hatching was significantly (P<0.01) inhibited only by the E. cinerea extract. All extracts had a significant (P<0.01) dose-dependent effect on the exsheathment of T. circumcincta and H. contortus infective larvae. The incubation with all heather extracts induced a reduction in adult T. circumcincta motility compared to the control, although significant (P<0.05) differences were only found at the highest concentration (1200 µg/mL). The effect of the mixed extract was significant at all concentrations and significant effects were also observed for C. vulgaris and E. umbellata at 600 µg/mL. These results show anthelmintic properties of heather phenolic extracts against T. circumcincta and H. contort, thus confirming observations from previous in vivo studies.

Effect of the consumption of heather on incoming larvae and established population of Teladorsagia circumcincta in experimentally infected Cashmere goats.

This study was performed in Cashmere goats that were experimentally infected with Teladorsagia circumcincta to investigate the effects of heather consumption on the establishment of incoming infective larvae (experiment 1) and on an adult nematode population (experiment 2). In experiment 1, 24 non-lactating goats were divided into 2 groups: heather-supplemented vs. non-supplemented. After 2 weeks of adaptation to the diet, all of the goats were experimentally infected with 6000 infective larvae of T. circumcincta. Twelve animals (6 controls and 6 supplemented with heather) were slaughtered at 6 days and at 3 weeks post-infection (pi). After slaughter, the worms were counted and the female worm fecundity and development were determined. Heather consumption was associated with a significant reduction in larval establishment at 6 days (P=0.033) and at 3 weeks (P=0.041) pi. No differences in worm counts between the slaughter times were found. In the goats slaughtered at 3 weeks pi, the number of eggs in utero and length of the female worms were significantly (P<0.001) lower than those of control group. In experiment 2, 24 non-lactating goats were experimentally infected with 10,000 T. circumcincta infective larvae daily for 5 consecutive days (total infection of 50,000 larvae). After 3 weeks, 2 groups were established: control and heather-supplemented. The faecal egg output of each animal was measured at 2-days interval during the experimental period. The goats were slaughtered after 3 weeks of heather administration for parasitological studies. Heather administration was associated with a significant decrease in egg excretion between 25 and 29 days pi. The worm counts were similar in both groups, but the female length and fecundity were significantly (P<0.001) lower in supplemented goats. These results show that heather consumption reduces the establishment of T. circumcincta larvae in goats and the development and fecundity of female adult parasites.

Is the anthelmintic effect of heather supplementation to grazing goats always accompanied by anti-nutritional effects?

To test the hypothesis that the beneficial anthelmintic effect of consuming moderate amounts of tannins may not always be accompanied by anti-nutritional effects in goats, two experiments were conducted. In the first, 48 Cashmere goats were randomly assigned to two treatments: supplementation with tannin-containing heather (6.4% total tannins) and non-supplementation. All goats grazed continuously from May to September under farm conditions in a mountainous area of northern Spain. The mean percentage of heather incorporated into the diet of the supplemented animals was 29.1%. Supplementation reduced the mean number of nematode eggs in faeces (P < 0.001) and the goat mortality rate (P < 0.05). The rumen ammonia concentration was markedly reduced in the goats receiving the heather supplement (160 v. 209 mg/l; P < 0.01), while volatile fatty acid (VFA) concentrations were significantly greater (63.0 v. 53.6 mmol total VFA/l; P < 0.05). The heather-supplemented goats also showed a lower loss of live weight (P < 0.01) and body condition score (P < 0.001). In the second experiment, batch cultures of rumen microorganisms with rumen fluid from nine goats whose diet included 29% heather - or not, were used to incubate three substrates (pasture, pasture + heather and pasture + heather + polyethylene glycol) to investigate in vitro ruminal fermentation. Differences (P < 0.01) among substrates were observed in terms of dry matter disappearance (DMD), in vitro true substrate digestibility (ivTSD), gas production and ammonia concentration, the greatest values always associated with the pasture substrate. Cultures involving rumen inoculum derived from goats receiving the heather-containing diet showed slightly lower DMD (46.9 v. 48.5 g/100 g; P < 0.05), ivTSD (64.6 v. 65.9 g/100 g; P < 0.10) and gas production (105 v. 118 ml/g; P < 0.001) values, but much greater total VFA concentrations (48.5 v. 39.3 mmol/l; P < 0.05), and suggest that the efficiency of ruminal fermentation in these animals was probably improved. Together, the results support the absence of a clear nutritional cost counteracting the beneficial anthelmintic effect of supplementing the diet of grazing goats with tannin-containing heather.