Article

Sheep self-medicate when challenged with illness-inducing foods

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Abstract

People learn to take aspirin for headaches, antacids for stomach aches and ibuprofen to relieve pain, and often obtain prescriptions from doctors for medications. Is it also possible that herbivores write their own prescriptions? From prehistoric times, people have looked to the presumed self-medicative behaviour of animals for remedies of ailments but it is still not clear whether animals seek medicinal compounds to recuperate from illness. Evidence of self-medication is based almost exclusively on observations rather than experimental analyses. We examined whether lambs select specific protective chemicals to recuperate from ingesting malaise-inducing foods. In pen studies, lambs in a treatment group were conditioned to consume foods and toxins (grain, tannins, oxalic acid) that led to negative internal states and then allowed to eat a substance known to attenuate each state (sodium bentonite, polyethylene glycol and dicalcium phosphate, respectively). Control lambs ate the same foods and medicines, but disassociated temporally, so they did not recuperate from illness. After conditioning, lambs ingested grain or food with tannins or oxalates and then received a choice of the three medicines. Only the treatment animals preferred to eat the specific compound known to rectify the state of malaise induced by the food previously ingested. Control animals never changed their pattern of use of the medicines, regardless of the food consumed before the choice. This first demonstration of multiple malaise–medicine associations supports suppositions in zoopharmacognosy and current hypotheses of homeostatic endeavour, and has implications for the well-being of wild and domestic animals.

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... Other branches of that tree did not end in an area of high centrality (Fig. 3A). The interpretation of these branches is more speculative: they may correspond to excursions as part of the fission-fusion social system , preferential travel routes when foraging for thinly distributed low-quality resources (Schmidt et al., 2016), immediate response to predator attacks (Courbin et al., 2016), prospection for mates (Lovari et al., 2008), or possibly visits to rarely needed resources such as self-medication and trace elements (Villalba, Provenza, & Shaw, 2006). ...
... The special case of resources that are critical to animal fitness but are seldom visited represents a strong limit to this type of approach (Powell & Mitchell, 2012). Examples of such resources include salt licks that the animals may only visit after ingesting toxic forage (Villalba et al., 2006). Visits to salt licks may be too far apart in time to be recorded. ...
... However, as long as adequate information is available to parameterize the simulating model, e.g. knowledge about physiological requirements for trace elements (Villalba et al., 2006), they can be simulated using individual-based models (Wang & Grimm, 2007;Signer, Fieberg, & Avgar, 2017;Fig. 2D). ...
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As technological and statistical innovations open new avenues in movement ecology, I review the fundamental implications of the time frame of home‐range studies, with the aim of associating terminologies consistently with research objectives and methodologies. There is a fundamental distinction between (a) extrapolations of stationary distributions, associated with long time scales and aiming at asymptotic consistency, and (b) period‐specific techniques, aiming at specificity but typically sensitive to the sampling design. I then review the difference between function and utilization in home‐range studies. Most home‐range studies are based on phenomenological descriptions of the time budgets of the study animals, not the function of the visited areas. I highlight emerging trends in automated pattern‐recognition techniques for inference about function rather than utilization.
... Wild herbivores face the same challenges from internal parasites, ticks and disease as domestic ones, whose populations in confinement are maintained through veterinary inputs in the form of synthetic drugs and antibiotics (Acha and Melendez, 1983). However, when domestic herbivores have access to high plant and phytochemical diversity (secondary metabolites) across broad landscapes, they have much less need for veterinary inputs (see Provenza et al., 2003;Villalba et al., 2006;Provenza et al., 2007;Villalba and Provenza, 2007;Provenza and Villalba, 2010;Gessner et al., 2017;Villalba et al., 2017;Provenza, 2018). It follows, therefore, that with wild herbivores having no access to veterinary interventions, their populations, together with genetic adaptation to disease through natural selection, must rely on plant secondary metabolites to remain healthy and productive. ...
... Livestock learn to use different medicines to rectify different maladies. Sheep learn to use sodium bicarbonate to alleviate illness from eating too much grain, polyethylene glycol to counteract food containing high levels of tannins, and dicalcium phosphate to counteract food with high levels of oxalic acid (Villalba et al., 2006). When fed grain or food with tannins or food with oxalic acid, sheep choose the medicine that rectifies the malady. ...
Article
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... However, many drought-tolerant perennial shrubs can contain high levels of minerals (Na, Cl, K, Se and S) and secondary compounds (oxalates, nitrates, tannins and saponins), which can have anti-nutritional properties that limit voluntary feed intake (Norman et al., 2013). Ruminants do learn to consume particular feeds because they raise fitness, for example sheep have been shown to self-medicate when they have consumed illnessinducing feeds (Villalba et al., 2006a; Howery et al., 2010). It is possible that sheep may also learn to overcome the aversive effects of the unfavourable characteristics of the feeds and consume adequate amounts to remediate a vitamin E deficiency. ...
... The association, by animals, of post-ingestive effects with food flavours has been identified as a means through which herbivores learn about the consequences of feeds (Duncan and Young, 2002). Learning about components of the feedbase is an important tool that herbivores use to modify foraging behaviour (either to prefer or to avoid feeds) in order to cope and adapt quickly to the changing internal and external environments for the achievement of nutritional homoeostasis (Villalba et al., 2006a; Howery et al., 2010). It has been experimentally established that foraging behaviour is formed by two learning mechanisms: selflearning from experience and trial and error by relying on post-ingestive feedbacks and learning from peers or from the mother (Provenza and Balph, 1990). ...
Article
Given the capacity of ruminants to modify diet selection based on metabolic needs, we hypothesised that, when given a choice, lambs experiencing a vitamin E deficiency would consume more of a vitamin E-enriched feed than lambs not deficient in vitamin E. Fifty-six Dohne Merino lambs were divided into two groups and fed either a vitamin E-deficient diet over 40 days to induce low plasma vitamin E or a vitamin E-enriched diet to induce high plasma vitamin E. The lambs were then offered a choice of vitamin E-enriched and vitamin E-deficient pellets. For half of the animals, the enriched diet was paired with strawberry flavour and the deficient diet was paired with orange flavour, while the reverse pairings were offered to the others. Lamb preference for the diets was measured daily for the following 15 days. There was a three-way interaction between the high and low vitamin E treatment groups×vitamin E content and type of flavour in the feed×time (days). The lambs preferred pellets flavoured with strawberry but this preference changed to orange flavour in vitamin E-deficient lambs if the orange flavour was paired with high vitamin E. Lambs without a deficiency continued to prefer strawberry-flavoured pellets, regardless of the vitamin E concentrations in the pellets. It is possible that self-learning contributed to the low vitamin E group of lambs changing preference to orange flavour in order to consume more vitamin E, presumably to remediate the deficiency.
... Many studies have postulated or demonstrated that animals eat specific plants to combat or control disease [23,25,60,[64][65][66]. Experimental evidence for self-medication is lacking from free-ranging primates, but the phenomenon that primate species eat plant parts with very low nutritive quality suggests another role for these items, such as fighting against diseases [61]. ...
Article
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Gums produced by trees after injuries are valuable food resources for several primate species. Yet, information on the chemical characteristics of gum is scant and inconsistent. We use gums consumed by lemurs (strepsirrhine primates of Madagascar) as an example to illustrate their possible nutritive and pharmaceutical properties. Exudates from 45 tree species of the dry forests of Madagascar contained 0.38–23.29% protein, 0.46–65.62% sugar, and 0.39–11.86 kJ/g of energy in dry matter. Exemplified by the lemur species Microcebus griseorufus, gum consumption increased with increasing sugar and energy content but was unrelated to protein. But lemurs also fed on gum with very low protein and energy content, suggesting that these exudates were consumed for other reasons. Disk diffusion tests with exudates from five out of 22 tree species consumed by lemurs showed antibacterial activity against Micrococcus spp. and/or Staphylococcus aureus. Exudates with antibacterial activity had lower protein, sugar, and energy contents than samples without antibacterial properties. GC-MS analyses revealed several components with antimicrobial effects that would have the potential for self-medication. This might explain the consumption of gum with very low nutritive value. Possible medicinal effects of tree exudates deserve further attention in view of their pharmaceutical applicability for animals and humans alike.
... Additionally, feeding lambs with chicory or tannin-rich plants like sulla or wilted cassava foliage can enhance their health and decrease their dependency on chemical drugs [98,99]. Additionally, allowing animals to exhibit selfmedication behavior by ensuring they have access to multispecies forages typically part of their diet that contain bioactive ingredients can improve their health [100,101]. ...
... Additionally, feeding lambs with chicory or tannin-rich plants like sulla or wilted cassava foliage can enhance their health and decrease their dependency on chemical drugs [98,99]. Additionally, allowing animals to exhibit selfmedication behavior by ensuring they have access to multispecies forages typically part of their diet that contain bioactive ingredients can improve their health [100,101]. ...
Article
Full-text available
The role of small ruminant production in achieving sustainable and resilient food systems in low- and middle-income countries (LMICs) is yet to be fully explored or incorporated into current agroecological practices and policies. This review examines the principles and practices of agroecology, focusing on circular food systems and the sociopolitical aspects of their implementation for small ruminant production in LMICs. It discusses Gliessman’s five levels of agroecological transition and eight principles for integrating small ruminant production into agroecology: input reduction, animal health, soil health, biodiversity, recycling, synergy, economic diversification, and co-creation of knowledge. The review highlights that, while there are differing interpretations in the scientific literature, there is a growing consensus that agroecological practices applied to small ruminant production have the potential to improve integration and self-sufficiency in farming systems, improve animal health, reduce reliance on external inputs, and promote circularity and biodiversity. This reinforces the view that agroecological approaches to small ruminant production can foster a sustainable and interconnected system that strengthens the relationships between animals, plants, and the environment and enhances circularity. To achieve successful implementation and widespread adoption of these approaches, it is crucial to facilitate greater collaboration and cocreation of knowledge among small ruminant farmers and stakeholders in the small ruminant livestock industry.
... They may also collect and give these resources to their animal themselves (case 2). Experiments conducted on sheep (Villalba et al. 2006), and horse (post-gastric herbivores, like elephants) (Williams 2008) have shown the ability of these herbivores to develop learned self-medication behaviours; thus, a taste-marked forage containing the remedy for a previously induced pathological condition is subsequently preferred to a neutral forage when the same pathological condition is induced in these animals again. If we extend to elephants the ability of other herbivores to associate a taste (including associated aromas) with a perceived therapeutic effect and to remember it, 6 these care practices of the mahouts, when addressed to a 'naïve' elephant, can have the effect in case 1, of guiding its choices in the direction of re-acquiring knowledge from its conspecifics, but above all, in case 2, can constitute a form of transmission of knowledge from 'knowing' elephants to naïve conspecifics via humans. ...
Article
Full-text available
The mahouts and the elephants of Thongmyxay district in Laos form an original hybrid community composed of humans and an animal species represented by both wild and domestic animals (elephants). We investigated in particular the interactions between mahouts' observation of elephants and their own medicinal practices (human and ethnoveterinary), which have been the subject of two previous publications. Based on this material, supplemented by data collected on the status of domestic village elephants and interviews with four local healers, we discuss here with a multispecies approach combining ethnographic data and ethological knowledge, the modalities of construction and possible exchanges of medicinal knowledge between the two species.Elephants have a status characterised by a double hybridity: wild and domestic on the one hand (with ontological circulations from one state to the other), and animal and human on the other. The part of humanity attributed to the elephants is reflected in particular in the self-agency that is recognised in their ability to heal themselves when they are suffering, which leads to the therapeutic use of their dung by Mahouts. The mahouts include in their ethnoveterinary care of the elephants plants that they see the elephants use when they are ill. The medicinal uses they make of some of these plants in their households are more consistent with their observations of these elephant behaviours than it is with the use of the same items by local healers, suggesting a transfer of medicinal knowledge from elephants to mahouts.Since some of the village elephants in Thongmyxay are still periodically released and come into contact with their wild counterparts, the domestication space forms an interface between humans, wild elephants and the forest, and we discuss conversely the possibility of knowledge transfer from mahouts to village elephants through the ethnoveterinary care they receive.This knowledge which is precious for the health and well-being of people and elephants in Laos is threatened by the reduction of the forest cover sheltering the resources used, and by the relocation of village elephants to tourist centres. Thus, emerges the need to think the conservation of the intangible heritage represented by the knowledge of hybrid communities in terms of interspecific heritage, implying that along with the preservation of the ecosystems that host the resources involved in this knowledge, care must be taken to maintain their access to all the populations (human and animal) that use them.
... Ils peuvent aussi collecter et donner eux-mêmes ces ressources à leur animal (cas n° 2). Des expériences menées sur des moutons (Villalba et al. 2006) et des chevaux -herbivores post-gastriques comme les éléphants - (Williams 2008) ont montré la capacité de ces herbivores à développer des comportements d'automédication appris ; ainsi un fourrage marqué par un goût et contenant le remède à un état pathologique induit préalablement est privilégié par la suite à un fourrage neutre lorsqu'on induit à nouveau le même état pathologique chez ces animaux. Si l'on étend aux éléphants ces capacités qu'ont d'autres herbivores d'associer un goût (incluant les arômes associés) à un effet thérapeutique ressenti et d'en garder la mémoire 6 , ces pratiques de soin des cornacs que nous venons de rappeler, lorsqu'elles s'adressent à un éléphant « naïf », peuvent avoir pour effet dans le cas n°1 d'orienter ses choix dans le sens d'une réacquisition de savoirs de ses congénères, mais surtout peuvent constituer dans le cas n° 2 une forme de transmission de savoirs d'éléphants « sachants » vers des congénères naïfs par l'entremise des humains. ...
Article
Full-text available
The mahouts and the elephants of Thongmyxay district in Laos form an original hybrid community composed of humans and an animal species represented by both wild and domestic animals (elephants). We investigated in particular the interactions between mahouts' observation of elephants and their own medicinal practices (human and ethnoveterinary), which have been the subject of two previous publications. Based on this material, supplemented by data collected on the status of domestic village elephants and interviews with four local healers, we discuss here with a multispecies approach combining ethnographic data and ethological knowledge, the modalities of construction and possible exchanges of medicinal knowledge between the two species. Elephants have a status characterised by a double hybridity: wild and domestic on the one hand (with ontological circulations from one state to the other), and animal and human on the other. The part of humanity attributed to the elephants is reflected in particular in the self-agency that is recognised in their ability to heal themselves when they are suffering, which leads to the therapeutic use of their dung by Mahouts. The mahouts include in their ethnoveterinary care of the elephants plants that they see the elephants use when they are ill. The medicinal uses they make of some of these plants in their households are more consistent with their observations of these elephant behaviours than it is with the use of the same items by local healers, suggesting a transfer of medicinal knowledge from elephants to mahouts.Since some of the village elephants in Thongmyxay are still periodically released and come into contact with their wild counterparts, the domestication space forms an interface between humans, wild elephants and the forest, and we discuss conversely the possibility of knowledge transfer from mahouts to village elephants through the ethnoveterinary care they receive.This knowledge which is precious for the health and well-being of people and elephants in Laos is threatened by the reduction of the forest cover sheltering the resources used, and by the relocation of village elephants to tourist centres. Thus, emerges the need to think the conservation of the intangible heritage represented by the knowledge of hybrid communities in terms of interspecific heritage, implying that along with the preservation of the ecosystems that host the resources involved in this knowledge, care must be taken to maintain their access to all the populations (human and animal) that use them.
... Depending upon the dosage, type, and concentration, some PSC may be toxic and elicit anti-nutritional effects for animal intake (Freeland and Janzen, 1974;Villalba et al., 2017). However, upon learning, ruminants remember and avoid toxins and toxic effects by selecting parts of the plants that do not contain large amounts of these chemicals (Freeland and Janzen, 1974) or selecting specific PSC with pharmaceuticals and prophylactic traits at self-regulated safe levels of intake to selfcounteract toxins (Villalba et al., 2006;Torregrossa and Dearing, 2009). On the other hand, ruminants positively search for and select plants and parts of plants to prevent and medicate illness (Provenza, 1996;Villalba and Provenza, 2007). ...
Article
Full-text available
Grasslands are heterogeneous landscapes composed of a diversity of herbaceous and shrub vegetation that varies not only taxonomically, but biochemically in terms of primary and secondary compounds. Plant Secondary Compounds (PSC) have specific nutritional, medicinal, and prophylactic properties, to which benefits depend upon dosage, type, arrangements, and concentration that changes between and within plants across time and space. The knowledge of the plant content of PSC and their distribution in grazing environments would therefore contribute to the design and creation of healthier foodscapes for ruminants; in other words, healthscapes. Geographic information systems (GIS) have been used extensively for landscape visualization and assessment, through several spatial analysis techniques applied for the creation of virtual maps to add valuable information to a particular environment. Given the knowledge of plants and their composition, GIS emerges as a readily available and low-cost tool to assess and evaluate the distribution of plants with beneficial PSC in large and heterogeneous foodscapes. We present and propose for the very first time, the application and use of GIS to determine the spatial distribution of PSC rich plants with nutraceutical properties to illustrate, visualize, and generate healthscapes for grazing ruminants. We present healthscape maps created using botanical composition analyses and advanced image classification methods to illustrate the distribution of plants regarding their PSC and nutraceutical properties. Such maps add an extra dimension and perspective to plant chemical composition, enabling graziers to visualize in space and time centers of nutrition and prophylactics or medicines, contributing to advanced grazing management decisions toward more productive, sustainable, and healthy grazing systems. The valuable information behind the mapped PSC advances the understanding of the nutritional ecology of grazing environments and foodscapes, introducing a new dimension to the holistic management of pastoral livestock production systems.
... Gastrointestinal parasites may reduce the fitness of hosts by depressing appetite, tissue deposition and skeletal growth (Sykes and Coop 2001), and could more seriously result in poor growth rate, ill-thrift and death (Min et al. 2005). Animals self-medicate by selecting and ingesting substances that may help reduce the infestations of internal parasites (Huffman 2003;Villalba et al. 2006). Bark is rich in plant secondary metabolites (PSMs), especially condensed tannins, which likely play an important role in attenuating the effect of internal parasites. ...
Article
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The bark-stripping behavior of Formosan sambar, Rusa unicolor swinhoii, has become conspicuous in recent years in the Tataka area of Yushan National Park in Taiwan and a cause for concern to visitors and ecologists. We conducted a monthly survey of 537 tagged trees of 21 species and monitored the abundance of sambar using camera traps from October 2018 to January 2021, aiming to interpret possible causes of the bark-stripping behavior in Tataka. We also used a generalized linear model to evaluate factors that may affect the probability of a tree having its bark stripped. Both our observations and the model predictions showed that sambar has a strong preference for bark of Pinus armandii, Photinia niitakayamensis, and Salix fulvopubeseens and for trees with diameter at breast height around 14 cm. Bark stripping mainly occurred between July and October when major forage was most abundant. However, sambar's need for bark surged in May when sambar abundance was moderate and decreased in October when sambar abundance was high. The seasonality of bark stripping was synchronized with the peak periods of antler development, fawn nursing, and spread of gastrointestinal parasites, suggesting that sambar strips bark to ingest minerals for their physiological needs and/or to acquire plant secondary metabolites to repel gastrointestinal parasites. Sambar abundance alone was not sufficient to predict the overall intensity of bark stripping. Rather, the product of sambar abundance and the necessity index (average wound size) were strongly correlated with the overall bark-stripping intensity. Therefore, controlling sambar abundance is essential but it alone may not be the optimal strategy for controlling bark stripping. A combination of population control and relaxing of sambar's parasite loading and/or physiological needs for minerals is an important strategy to control the overall bark stripping. Future research could use the necessity index to investigate the synchronicity of the bark-stripping behavior, deer's physiological state, environmental factors and phenology to better understand the cause of this behavior.
... In addition, ruminants offered functional diversity can self-medicate by ingesting the appropriate dose of a PSC Villalba et al., 2011b;Provenza et al., 2015). For example, Villalba et al. (2006), observed that lambs fed, grain, tannins, and oxalic acid, in toxic quantities could select doseappropriate quantities of three different 'medicinal' substances (sodium bentonite, polyethylene glycol, and dicalcium phosphate) to counteract the aversive effects of toxins. Thereby, functionally diverse diets can allow animals to acquire dose-dependent effects that would be unattainable to animals offered a monoculture sward or be unlikely to occur by chance of resulting species abundances within a mixture, or that would incur a greater detoxification cost. ...
Thesis
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The hypothesis of my research was that providing functional diversity as opposed to dietary monotony will: alter fermentation patterns, increase animal production, and reduce negative environmental impacts, enhance animal welfare, and alter neophobia and partial preference through in utero and early life exposure. Thereby the objective was to determine the effects of altering the functionality of diverse diets (through context, species abundance, species distribution, and temporal availability) on dry matter intake, production, welfare, the environmental impacts, and partial preference compared with animals grazing a monotonous diet. This research was conducted over several experiments. Chapter 3 implemented an in vitro rumen fermentation methodology to determine if diverse versus monotonous diets altered fermentation patterns and allowed for assumptions on production and environmental impacts. The results showed that increasing the portion of chicory, plantain, or a diverse combination (chicory, plantain, and alfalfa) to ryegrass increased 24 hr gas production and branched-chain volatile fatty acid production, while reducing ruminal ammonia concentration. Chapter 4 applied an equal parts dry matter (DM) diverse combination diet to ram lambs and compared their intake, performance, welfare, and urinary nitrogen excretion to those offered a repetitive ryegrass diet. Lambs grazing the diverse diet had a 48% greater dry matter intake (DMI), 92% greater average daily gain (ADG), 25% lower day-to-day coefficient of variation (CV) of intake, and had a 30% lower urinary N concentration. Ram lambs provided a varied diet with set ratios of each species had a 20 and 10% greater DMI and a 29 and 23% reduced DMI CV compared to a diverse diet of the same ratios and a ryegrass diet that were monotonous in presentation in the experiment in chapter 5. The experiment in Chapter 6 explored the diverse and varied diet treatments without the restrictions of set ratios of species and compared them to a monotony of alfalfa. At the same level of intake the varied diet lambs gained 67 and 28% greater than the diverse and alfalfa diet. This greater ADG of the varied lambs occurred with the same intake and diet primary chemistry as the diverse diet, indicating that performance was affected by more than primary chemistry. Lambs consuming the alfalfa treatment conducted 150% more v bouts of stereotypic behavior than the diverse and varied lambs. Chapter 7 provided ewes in the final third of gestation with diverse or monotonous ryegrass diets. I found that diverse ewes birthed heavier lambs and exhibited lower levels of oxidative and metabolic stress 24 hrs after lambing. In chapter 8 the lambs birthed in chapter 7 remained with their dams on their respective treatment until weaning (early life experience) or were removed 24 hrs after lambing to pinpoint the effect of in utero and early life on partial preference and neophobic behaviors in later life. Lambs with early life exposure had partial preference altered more than those with only in utero exposure. Lambs exposed to ryegrass in utero or in utero and in early life spent more time grazing ryegrass than their diverse counterparts. In general, the diverse lambs had reduced latency to graze the diverse species compared to the ryegrass lambs. Further, all lambs chose to comprise a mixed species diet. This chapter demonstrated how in utero and early life experience can reduce neophobic behavior and that when provided the choice animals prefer to comprise a diverse diet even if a familiar forage species is available.
... In experimental settings, sheep have been shown to have the ability to learn to self-medicate. There has been extensive research conducted previously with sheep on their ability to learn to self-medicate for parasitic infection (Villalba et al., 2010;Fishpool et al., 2012;Juhnke et al., 2012) as well as other internal pain not caused by parasites (Provenza et al., 2000;Villalba and Provenza, 2001;Villalba et al., 2006). If sheep can learn to self-medicate, then their choice to ingest medications that are non-addictive would provide a strong indicator that the animal is motivated to alleviate a negative affective state. ...
Article
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Lambs in Australia undergo painful husbandry procedures as part of common husbandry. The magnitude and duration of pain are difficult to assess in lambs. Most currently used methods rely on behavioral expressions and physiological markers that may fail to detect the state of pain an animal experience. This study examined motivation of 12-week-old lambs experiencing chronic pain to self-medicate by consumption of feed containing an analgesic agent as an indicator of pain in lambs. In this study, 36 male Merino lambs were individually penned and acclimated to pelleted feed and two artificial odors: strawberry and banana. Once acclimated to odored feed, lambs were tested for their individual preference for the odors. Lambs were then assigned to one of two groups: Sham—sham handled day 0 and 7 or Ring—Ring castrated day 0 and tail docked day 7. To enable self-medication testing, lambs underwent a conditioning period (day 0–3) followed by the self-medication period (day 7–12). On day 0 lambs were castrated or sham handled, and then offered only medicated feed that contained an odor cue (either strawberry or banana). On day 7, lambs underwent tail-docking or sham handling and were offered both the conditioned medicated feed and non-medicated feed. Amount of each feed consumed was recorded 1 and 12 h after offer each day. Blood samples were taken for cortisol and white blood cell analysis and behavioral observations were recorded for 12 h following treatment. There was no difference in preference for medicated feed between Ring and Sham lambs during the self-medication phase (P = 0.18). Lambs in both groups displayed a significant preference for strawberry cued medicated feed during the self-medicated period when compared to the other testing periods (P = 0.05). Ring lambs displayed more active pain behaviors (mean = 15.1) than Sham (mean = 0.4, P < 0.05). Following castration, Ring lambs had a higher neutrophil/lymphocyte ratio at 6, 24, 48, and 72 h. This study was not able to demonstrate that lambs can self-medicate for a state of pain.
... The results of this study contribute to enhancing the knowledge of zoopharmacognosy and the investigation of natural products with potential for use in both veterinary and botanical pharmacology. Numerous studies have shown the existence of self-medication behavior in vertebrates [39][40][41][42][43] and invertebrates [44][45][46]. The fact that the most active extracts studied here are from coleopteran species found in the Great Bustard diet while the least active ones are not, suggests a self-medication function for the toxic compounds present in these insect species. ...
Article
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Arthropods and specifically beetles can synthesize and/or sequester metabolites from dietary sources. In beetle families such as Tenebrionidae and Meloidae, a few studies have reported species with toxic defensive substances and antiparasitic properties that are consumed by birds. Here we have studied the antiparasitic activity of extracts from beetle species present in the habitat of the Great Bustard (Otis tarda) against four pathogen models (Aspergillus niger, Meloidogyne javanica, Hyalomma lusitanicum, and Trichomonas gallinae). The insect species extracted were Tentyria peiroleri, Scaurus uncinus, Blaps lethifera (Tenebrionidae), and Mylabris quadripunctata (Meloidae). M. quadripunctata exhibited potent activity against M. javanica and T. gallinae, while T. peiroleri exhibited moderate antiprotozoal activity. The chemical composition of the insect extracts was studied by gas chromatography coupled with mass spectrometry (GC-MS) analysis. The most abundant compounds in the four beetle extracts were hydrocarbons and fatty acids such as palmitic acid, myristic acid and methyl linoleate, which are characteristic of insect cuticles. The presence of cantharidin (CTD) in the M. quadripunctata meloid and ethyl oleate (EO) in T. peiroleri accounted for the bioactivity of their extracts.
... Dr Jaquinto, the trusted physician to Queen Ann, wife of James I in 17th century England, is said to have made systematic observations of domestic sheep foraging in the marshes of Essex, which led to his discovery of a successful cure for consumption (Wilson 1962). Recent controlled studies on sheep have demonstrated their ability to learn to associate the curative properties of items paired with the experimentally induced gastrointestinal discomfort, and then select the appropriate 'cure' for that specific discomfort (Villalba et al. 2006). Studies like these have been able to demonstrate what naturalistic studies on other species are unable to do due to ethical constraints, going a long way in revealing the breadth of self-medicative abilities in animals. ...
... There is increasing evidence from other groups of animals that individuals may be capable of adopting specific self medication behaviours in response to parasite infections. Examples of zoopharmacognosy include the ingestion of specific antiparasitic compounds by mammals (Hutchings et al., 2003;Villalba et al., 2006), defensive anointing (Weldon, 2004) and the incorporation of green plants with antiparasitic properties into nests by birds (Gwinner et al., 2000;Petit et al., 2002). However, we are not aware of any similar studies conducted on fish. ...
Chapter
Parasites1 are ubiquitous components of natural and managed ecosystems. Water provides a particularly suitable medium for the support and movement of directly transmitted ectoparasites, and the central role played by fish in aquatic food webs make them ideal intermediate hosts for indirectly transmitted infections. Consequently, parasites use fish as hosts. By definition, parasite infections negatively impact host fish, though very often these detrimental effects are unquantified. However, parasites rely on their hosts for nutrition; so they impose at least energetic demands, and very often they have other physiological impacts. Parasites can be classified as environmental stressors of fish, not only because of the direct impacts they impose after infection, but also because their presence in aquatic environments can reduce the efficiency with which fishes function. The latter effects may arise because of constraints imposed on, for example, the habitat selection or food selection choices that are available to fish attempting to avoid contact with infectious agents.
... In that way, herbivores may have the ability to select a feed rich in secondary metabolites to alleviate malaise. Some previous experiments have highlighted such abilities in sheep (Villalba et al., 2006), including scenarios when malaise was caused by the presence of GIN (Martínez-Ortíz-de-Montellano et al., 2010;Juhnke et al., 2012;Amit et al., 2013). Conversely, others showed less conclusive results with only limited behavioural modifications or no evidence of self-medication (Lisonbee et al., 2009;Novelo-Chi et al., 2014;Ventura-Cordero et al., 2017 and. ...
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Plant secondary metabolites (PSM) are one of the promising options to control gastrointestinal nematodes in sheep and goats. The objective of this study was to assess the abilities of sheep and goats to self-medicate with tannin-rich sainfoin ( Onobrychis viciifolia ) (SF) when infected with gastrointestinal nematodes, using a cafeteria and an operant conditioning trial. Hypotheses were that parasitized (P) lambs and goat kids would show greater intake and preference for SF than their non-parasitized (NP) counterparts, that kids would eat more SF than lambs (due to their lower resistance against parasites and their greater ability to consume PSM), and that SF intake would increase over time for P animals. We used 20 female kids and 20 ewe lambs aged 3 months. Half of the animals per species ( n = 10) were experimentally infected with 170 L 3 larvae of Haemonchus contortus /kg of BW (P). The other half were free from parasites throughout the study (NP). Five weeks after infection, animals were exposed to a 24-day cafeteria trial (three 8-day periods) offering a free choice between two legume pellets: SF (3.8% condensed tannins) and alfalfa (ALF, Medicago sativa ; no tannin). Subsequently, animals were involved in an operant conditioning trial of two 4-day long sessions, to assess in short-term tests their motivation to walk for a SF reward when offered in choice with freely available ALF. In the cafeteria trial, SF preference was greater in kids than in lambs, particularly in the first two periods. We did not observe a greater preference for SF in P animals, which was even greater in NP animals for periods 1 and 2. Sainfoin intake increased through periods for P animals, which led to similar SF preferences for all groups during period 3. In the operant-conditioning trial, motivation to get the SF reward was similar between P and NP animals. These results support the hypotheses that goats are more willing to consume tanniferous feeds than sheep, and that P animals increased SF intake through time. However, the emergence of a curative self-medicative behaviour was not supported, as P individuals did not show greater SF intake, preference, nor a greater motivation to get SF than NP animals, regardless of animal species. These findings are discussed with previous results and some explanations are presented.
... From ethnographic evidence one can attest the constant and careful monitoring that people practice over their herds (Perezgrovas, 2004;Wyndham, 2009;Perezgrovas et al., 2014;Molnar, 2017). Moreover, pastoralists are well aware of their animals' abilities for self-medication (Huffman, 2003;Villalba et al., 2005). From this, we hypothesize that at least some of the species that people have learned to use (i.e., domesticate) were at first tried because they saw livestock species consuming them. ...
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Understanding both domestication processes and agricultural practices is an interdisciplinary endeavor. Ethnographic research is potentially helpful for reconstructing past events. Such knowledge is also crucial for documenting the links between biological and cultural diversity, as well as for future purposes such as innovation in food production and sustainability. Here, we review six ethnographic case studies in different pastoral socioecological systems of the American continent. The livestock species involved include the native South American camelids and Arctic reindeer, as well as some Old World species (mainly goats, sheep, and cattle). Starting with the Columbian exchange (15th-16th centuries) and continuing up to the present, Old World herbivores launched novel uses of the local flora which resulted in entirely new livelihoods and cultures, i.e., pastoralism with its variants. Three of these case studies approach specifically how herding ecologies (human–animal–plant relationships) stirred specific management practices (human–plant relationships) that in some instances have moved toward conscious human selection of plant phenotypes. The other examples correspond to three potential instances of similar ongoing processes that we propose on the basis of ethnobotanical and ethnozoological data that were produced separately by other authors. Based on the studies we have reviewed, along with additional information from other parts of the world, we are able to conclude that: (a) New World pastoralist societies are/have been continuously adding species to the humanity’s portfolio of useful plants; (b) animals have been aiding in this processes in different ways; and, (c) how human–animal–plant relationships unfold in the present could have been similar in the past, thus analogies may be proposed for explaining prehistoric multispecies interactions and their outcomes. With our review, we intend to bring more attention to contemporary pastoralists as plant managers, animals as agents in human-plant interactions, and domestication as a behavioral complex and multispecies process that is as important in the present or future as it was in the past. Our understanding of food production practices is not only fundamental for improving our current frameworks of governance, conservation, and restoration of useful species populations, but also of biocultural diversity altogether.
... The study of feed preferences has seen several approaches, including the following: (1) Campbell (1976) pioneered the study of flavour conditioning in farm animals by testing aroma transfer from the sow feed to the piglet potentially through maternal fluids; (2) Black and coauthors studied, in a systematic approach, feed choices of sheep base on the particle size, density of forage and levels of moisture among other characteristics Black 1984a, 1984b;Kenney et al. 1984), ultimately resulting in the development of mathematical models predicting feed intake in farm animals ( Black 1995Black , 2009Black , 2014); (3) Rubenheimer and Simpson introduced the concept of nutritional geometric framework Simpson and Raubenheimer 1993), integrating nutritional requirements, environment and ingredient constraints and physiological status (mainly health, immune responses, reproduction) and predicting food intake in a diversity of animal and human subpopulations (Simpson and Raubenheimer 2012); (4) the modelling of feed choices developed by Forbes and co-workers ( Forbes and Provenza 2000;Forbes 2007) was based on the theory that animals select feed to minimise the discomfort generated by imbalanced dietary regimes (the concept particularly stressed potential deleterious effects of excess nutrients); (5) a complementary contribution by Provenza and coauthors showed that animals develop dietary preferences by associating post-ingestive events, particularly related to health attributes (self-medication) and its relationship with immune response ( Villalba et al. 2006Villalba et al. , 2013Provenza 2007, 2009;Lisonbee et al. 2009; Villalba and Landau 2012); (6) additional knowledge in this area has been delivered by Kyriazakis and contributors, refering to diet selection based on specific nutrient requirements during gastrointestinal parasitism in sheep ( Kyriazakis et al. 1994Kyriazakis et al. , 1996. Furthermore, from a methodological view, the adoption of electrophysiological methods (studying the electrical stimulation of cranial chorda timpany and glossopharengial nerves), gene sequencing and gene-expression techniques and the use of brain imaging (such as magnetic resonance or positron emission tomography) have significantly improved our understanding of the physiological principals underpinning responses to feed sensory cues in farm animals ( Danilova et al. 1999;Glaser et al. 2000;Sauleau et al. 2009;da Silva et al. 2014). ...
Article
The fact that most farm animals have no dietary choice under commercial practices translates the dietary decisions to the carers. Thus, a lack of understanding of the principles of dietary choices is likely to result in a high toll for the feed industry. In healthy animals, diet selection and, ultimately, feed intake is the result of factoring together the preference for the feed available with the motivation to eat. Both are dynamic states and integrate transient stimulus derived from the nutritional status, environmental and social determinants of the animal with hard-wired genetic mechanisms. Peripheral senses are the primary inputs that determine feed preferences. Some of the sensory aspects of feed, such as taste, are innate and genetically driven, keeping the hedonic value of feed strictly associated with a nutritional frame. Sweet, umami and fat tastes are all highly appetitive. They stimulate reward responses from the brain and reinforce dietary choices related to essential nutrients. In contrast, aroma (smell) recognition is a plastic trait and preferences are driven mostly by learned experience. Maternal transfer through perinatal conditioning and the individual's own innate behaviour to try or to avoid novel feed (often termed as neophobia) are known mechanisms where the learning process strongly affects preferences. In addtition, the motivation to eat responds to episodic events fluctuating in harmony with the eating patterns. These signals are driven mainly by gastrointestinal hormones (such as cholecystokinin [CCK] and glucagon-like peptide 1 [GLP-1]) and load. In addition, long-term events generate mechanisms for a sustainable nutritional homeostasis managed by tonic signals from tissue stores (i.e. leptin and insulin). Insulin and leptin are known to affect appetite by modulating peripheral sensory inputs. The study of chemosensory mechanisms related to the nutritional status of the animal offers novel tools to understand the dynamic states of feed choices so as to meet nutritional and hedonic needs. Finally, a significant body of literature exists regarding appetite driven by energy and amino acids in farm animals. However, it is surprising that there is scarcity of knowledge regarding what and how specific dietary nutrients may affect satiety. Thus, a better understanding on how bitter compounds and excess dietary nutrients (i.e. amino acids) play a role in no-choice animal feeding is an urgent topic to be addressed so that right choices can be made on the animal's behalf.Journal compilation
... Associated with this, the concept of providing a range of plant species with which animals can self-medicate when given the opportunity is gaining credence (Clayton and Wolfe, 1993;Huffman, 2003;Huffman and Seifu, 1989). The literature surrounding the effectiveness of traditional plant remedies and the ability of farmed animals to treat themselves is growing, for example in Africa (Githiori et al., 2004), South America (Lans and Brown, 1998), India (Sharma and Singh, 1989), Europe (Pieroni et al., 2004) and the United States (Villalba et al., 2006). ...
... In non-human animals, self-medication remains a controversial subject, because evidence is mostly anecdotal. A few experimentally verified cases of self-medication support the theoretical expectation that animals can and do make specific foraging decisions that function specifically to remediate illness (Huffman and Caton, 2001;Villalba et al., 2006;Singer et al., 2009). In zoological medicine, this concept has first been implemented by primate keeping institutions. ...
... There is increasing evidence from other groups of animals that individuals may be capable of adopting specific self medication behaviours in response to parasite infections. Examples of zoopharmacognosy include the ingestion of specific antiparasitic compounds by mammals (Hutchings et al., 2003;Villalba et al., 2006), defensive anointing (Weldon, 2004) and the incorporation of green plants with antiparasitic properties into nests by birds (Gwinner et al., 2000;Petit et al., 2002). However, we are not aware of any similar studies conducted on fish. ...
... Il existe quelques observations et preuves expérimentales dans des contextes particuliers qui laissent à penser que les herbivores seraient capables de s'automédiquer mais elles sont à l'heure actuelle trop peu nombreuses pour permettre une reconnaissance générale de cette capacité. Expérimentalement, Villalba et al. (2006) ont montré que des agneaux apprennent à choisir la substance, parmi trois, qui rectifie le malaise induit par un aliment récemment ingéré. Ils sont aussi capables d'augmenter volontairement leur consommation de polyéthylène glycol, une substance qui se fixe aux tannins des plantes et en réduit l'effet toxique, lorsque la teneur en tannins de leur régime augmente . ...
... This detoxification can explain the consumption, in times of hunger, with soil, of items that might not be edible or generally consumed. An experimental study on sheep confirms that animals may learn to associate toxic food with its antidote to decrease malaise (Villalba et al 2006). Three reasons appear to reject hunger or poor diet quality as a major reason for soil eating by Kanyawara chimpanzees: (1) the amount of clay material ingested is very small (usually less than a handful, which is likely not to be enough to give a sensation of fullness to the stomach), (2) geophagy occurs mainly in the afternoon when chimpanzees have already eaten large amounts of food and (3) soil eating does not seem to be restricted to periods of food scarcity. ...
... Sheep learn to ingest polyethylene glycol (PEG) to attenuate aversive effects of tannins; they discriminate the benefits of PEG from non-medicinal substances, they ingest PEG after a meal high in tannins (Villalba & Provenza, 2001), and they titrate the dose of PEG in accord with the amount of tannin in the diet (Provenza, Burritt, Perevolotsky, & Silanikove, 2000). In the most complex studies to date, sheep learned to selectively ingest three medicines e sodium bentonite, polyethylene glycol, dicalcium phosphate e that attenuate illnesses from eating too much grain, tannins, or oxalic acid, respectively (Villalba, Provenza, & Shaw, 2006). ...
Article
We contend that palates link herbivores and humans with landscapes and consider how these relationships have changed historically. An attuned palate, which enables herbivores to meet needs for nutrients and self-medicate to rectify maladies, evolves from three interrelated processes: flavor-feedback associations, availability of phytochemically rich foods, and learning in utero and early in life to eat nourishing combinations of foods. That occurs when wild or domestic herbivores forage on phytochemically rich landscapes, is less common when domestic herbivores forage on monoculture pastures, is close to zero for herbivores in feedlots, and is increasingly rare for people who forage in modern food outlets. Unlike our ancestors, the palates of many individuals are no longer linked in healthy ways with landscapes. Industrial farming and selection for yield, appearance, and transportability diminished the flavor, phytochemical richness, and nutritive value of fruits and vegetables for humans. Phytochemically impoverished pastures and feedlot diets can adversely affect the health of livestock and the flavor and nutritive value of meat and milk products for humans. While flavors of produce, meat, and dairy have become blander, processed foods have become more desirable as people have learned to link synthetic flavors with feedback from energy-rich compounds that obscure nutritional sameness and diminish health. Thus, the roles plants and animals once played in nutrition have been usurped by processed foods that are altered, fortified, and enriched in ways that can adversely affect appetitive states and food preferences. The need to amend foods, and to take nutrient supplements, could be reduced by creating phytochemically rich plants and herbivores and by creating cultures that know how to combine foods into meals that nourish and satiate. Copyright © 2015. Published by Elsevier Ltd.
... Sheep also learn to discriminate among minerals such as phosphorus, calcium and sodium and they learn to select the flavoured 'food' previously paired with recovery from deficiencies of each of these minerals . In addition to learning preferences for nutritious foods, herbivores learn to prefer medicinally beneficial foods, as post-ingestive feedback from medicines provides positive post-ingestive experiences to sick animals (Villalba et al. 2006a;Villalba and Landau 2012). These studies have shown that herbivores can discriminate among many different internal states and that they learn to ingest foods that ameliorate nutrient deficits and that rectify illness. ...
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Conventional models of foraging, such as optimal foraging theory, generally take the univariate approach to explain the decisions of consumers on the basis of the intrinsic properties of foods, including nutrient concentration and abundance. However, the food environment is inherently diverse and, as a consequence, foraging decisions are influenced by the interactions among multiple food components and the forager. Foraging behaviour is affected by the consumer's past experiences with the biochemical context in which a food is ingested, including the kinds and amounts of nutrients and plant secondary compounds in a plant and its neighbours. In addition, past experiences with food have the potential to influence food preference and intake through a mechanism, namely, food hedonics, which is not entirely dependent on the classical homeostatic model of appetite control. Research on the impacts of experience with food context and its behavioural expression in natural settings should pioneer innovative management strategies aimed at modifying food intake and preference of herbivores to enhance their nutrition, health and welfare, as well as the health and integrity of the landscapes they inhabit.
... A growing body of literature in the behavioral, ecological and pharmacological sciences suggests that animals use certain plants for the control of parasite infection and related illnesses (Cousins et al., 2002). There are evidences of animal self-medication (Engel, 2002;Huffman, 2006Huffman, , 2003Huffman, , 1997Plotkin, 2000;Janzen, 1978;Burritt and Provenza, 2000;Provenza, 2001, 2002;Phy and Provenza, 1998;Villalba et al., 2010Villalba et al., , 2006Krief, 2012). However, we have little knowledge about the abilities of animals to self-medicate, and many of the observations are anecdotal and equivocal (Clayton and Wolfe, 1993;Lozano, 1998;Houston et al., 2001). ...
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Present study documents distinctive service-beneficiary complex relationship which exists between Berberis and mountain wildlife. The synergized co-evolved epigenetic relationship observed in high mountain areas is reported for the first time. Ethno-veterinarian and zoopharmacological practices from the area have also been discussed. It is a common observation of local communities that when wildlife (especially avian and ungulates) gets physically hurt, they rush towards Berberis and eat berries and/or chew leaves or bark. Various stakeholders including hunters (57.77±5.47%), shepherds (70.74±4.20%) and local population (29.52±4.19%) have either personally observed or believe in Berberis-wildpharma interactive phenomenon. Similarly traditional communities use Berberis spp. to treat various veterinarian ailments including fracture of limbs (42.23±1.73%), back bone injury (10.42±1.04%), external injury (7.76±1.06), internal wounds (22.10±1.27%) and delivery incisions (7.23±1.48%). During present investigations, no observation was recorded regarding utilization of Berberis by herpetofauna and carnivores animals.
... A growing body of literature in the behavioral, ecological and pharmacological sciences suggests that animals use certain plants for the control of parasite infection and related illnesses (Cousins et al., 2002). There are evidences of animal self-medication (Engel, 2002; Huffman, 2006 Huffman, , 2003 Huffman, , 1997 Plotkin, 2000; Janzen, 1978; Burritt and Provenza, 2000; Provenza, 2001, 2002; Phy and Provenza, 1998; Villalba et al., 2010 Villalba et al., , 2006 Krief, 2012). However, we have little knowledge about the abilities of animals to self-medicate, and many of the observations are anecdotal and equivocal (Clayton and Wolfe, 1993; Lozano, 1998; Houston et al., 2001). ...
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Berberis, a member of family Berberidaceae serves as ‘wildlife-clinic’ in nature. Berberis pseudumbellata subsp. gilgitica is endemic to the area and has become critically endangered. For the first time research documented unique epigenetic behaviour of different mountain wildlife and zoopharmacological practices of traditional communities from CKNP. When wildlife gets physical hurts, it rushes towards Berberis plants and eat berries, chew leaves or bark, which is found significant (p<0.001). Such behaviour is untaught, not learned, and automatic but seems pre-programmed instinctively triggered reflexed. This is observed in avifauna (House Sparrow, Chakor, Ram Chakor) and angulates (Flare horned Markhor, Himalayan Ibex) only. 79.03%, 36.51% hunters (SD±30.02), 83.24%, 58.23% shepherds (SD±17.68) and 41.92%, 17.11% local population (SD±17.54) believe and have personal observations respectively for Berberis-wildpharma interaction. Traditionally it is used zoopharmacologically for bone healing of limbs (93.67%), other bone fracture and injury (90.91%), other internal injuries (85.53%), external injuries (73.33%), delivery of sheep (38.12%), goat (17.36%) and cow (19.87%). No such observation was recorded with respect to herpetofauna and carnivores.
... A growing body of literature in the behavioral, ecological and pharmacological sciences suggests that animals use certain plants for the control of parasite infection and related illnesses (Cousins et al., 2002). There are evidences of animal self-medication (Engel, 2002; Huffman, 2006 Huffman, , 2003 Huffman, , 1997 Plotkin, 2000; Janzen, 1978; Burritt and Provenza, 2000; Provenza, 2001, 2002; Phy and Provenza, 1998; Villalba et al., 2010 Villalba et al., , 2006 Krief, 2012). However, we have little knowledge about the abilities of animals to self-medicate, and many of the observations are anecdotal and equivocal (Clayton and Wolfe, 1993; Lozano, 1998; Houston et al., 2001). ...
Article
Full-text available
Berberis, a member of family Berberidaceae serves as 'wildlife-clinic' in nature. Berberis pseudumbellata subsp. gilgitica is endemic to the area and has become critically endangered. For the first time research documented unique epigenetic behaviour of different mountain wildlife and zoopharmacological practices of traditional communities from CKNP. When wildlife gets physical hurts, it rushes towards Berberis plants and eat berries, chew leaves or bark, which is found significant (p<0.001). Such behaviour is untaught, not learned, and automatic but seems pre-programmed instinctively triggered reflexed. This is observed in avifauna (House Sparrow, Chakor, Ram Chakor) and angulates (Flare horned Markhor, Himalayan Ibex) only. 79.03%, 36.51% hunters (SD±30.02), 83.24%, 58.23% shepherds (SD±17.68) and 41.92%, 17.11% local population (SD±17.54) believe and have personal observations respectively for Berberis-wildpharma interaction. Traditionally it is used zoopharmacologically for bone healing of limbs (93.67%), other bone fracture and injury (90.91%), other internal injuries (85.53%), external injuries (73.33%), delivery of sheep (38.12%), goat (17.36%) and cow (19.87%). No such observation was recorded with respect to herpetofauna and carnivores. epigenetic behavior of mountain wildlife towards Berberis species.
... A beneficial effect such as the anti-inflammatory activity was demonstrated for C. pachystachya (Juan-Hikawczuk et al. 1998). In addition to current worldwide research programs almost exclusively dedicated to the study of antioxidant substances in the human diet (Stevenson & Hurst, 2007;Virgili & Marino, 2008;Van den Ende et al. 2011;Wootton-Beard & Ryan, 2011), efforts towards a better understanding of the role of non-nutritional metabolites in wild animal diet may offer promising complementary issues for the medicinal applications of plants (Lozano, 1998;Huffman, 2003;Villalba et al. 2006). ...
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The major part of plants foraged by endangered pampas and marsh deer at the Brazilian Pantanal Wetland belongs to botanical genera containing medicinal species used mainly for healing infectious and inflammatory diseases. In this study, extracts and fractions of 19 plant species from deer diet were in vitro screened against bovine and suid herpesviruses, avian reovirus and infectious bursal disease virus (IBDV). At non-cytotoxicity concentrations, the extract and/or fractions from 14 plant species presented antiviral activity against at least one of these viruses except IBDV. Cecropia pachystachya, Melochia villosa and Polygonum acuminatum presented the most relevant results against bovine and suid herpesviruses while Andira cuyabensis was the most active plant against avian reovirus. C. pachystachya extract and fractions showed virucide effect, and kept their inhibitory activity towards both herpesviruses independent of the addition time in cell culture. Considering the benefit of the antioxidant activity of food and medicinal plants for the health balance, and its important role in viral infections, the extracts were also screened in a DPPH assay. Our findings show that several plants foraged by pampas and marsh deer possess antiviral activity against some pathogenic viruses for mammals. This study proposed an innovative strategy by adopting plants consumed in the diet of wild non-primate mammals for the search of potentially therapeutic substances.
... A growing body of literature in the behavioral, ecological and pharmacological sciences suggests that animals use certain plants for the control of parasite infection and related illnesses (Cousins et al., 2002). There are evidences of animal self-medication (Engel, 2002; Huffman, 2006 Huffman, , 2003 Huffman, , 1997 Plotkin, 2000; Janzen, 1978; Burritt and Provenza, 2000; Provenza, 2001, 2002; Phy and Provenza, 1998; Villalba et al., 2010 Villalba et al., , 2006 Krief, 2012). However, we have little knowledge about the abilities of animals to self-medicate, and many of the observations are anecdotal and equivocal (Clayton and Wolfe, 1993; Lozano, 1998; Houston et al., 2001). ...
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Full-text available
Berberis, a member of family Berberidaceae serves as 'wildlife-clinic' in nature. Berberis pseudumbellata subsp. gilgitica is endemic to the area and has become critically endangered. For the first time research documented unique epigenetic behaviour of different mountain wildlife and zoopharmacological practices of traditional communities from CKNP. When wildlife gets physical hurts, it rushes towards Berberis plants and eat berries, chew leaves or bark, which is found significant (p<0.001). Such behaviour is untaught, not learned, and automatic but seems pre-programmed instinctively triggered reflexed. This is observed in avifauna (House Sparrow, Chakor, Ram Chakor) and angulates (Flare horned Markhor, Himalayan Ibex believe and have personal observations respectively for Berberis-wildpharma interaction. Traditionally it is used zoopharmacologically for bone healing of limbs (93.67%), other bone fracture and injury (90.91%), other internal injuries (85.53%), external injuries (73.33%), delivery of sheep (38.12%), goat (17.36%) and cow (19.87%). No such observation was recorded with respect to herpetofauna and carnivores. epigenetic behavior of mountain wildlife towards Berberis species. Life Sci J Introduction Over thousands of years and generations after generations through 'trial and error' human race has gained inclusive experience and knowledge of zoopharmacology (Rodriguesa et al., 2003) and wildpharma. Standing on the giant shoulders and critical observations, traditional communities are able to recognize plants and their uses for a variety of human and animals' ailments. In case of wildlife, transmission of learning to next generations is not a common phenomenon and therefore, every generation and individual rely on a complex intelligent but supra-instinct-lead behaviour to deal with abnormal situations like accidents, wounds, illness etc. (Chakraborty et al., 2013; Gradé et al., 2009). Dealing with such unusual situations, animals graze specific plants (Gradé et al., 2009). According to Brunfels (1532), the leaves of Laurus nobilis are used by pigeons and chickens to cure constipation. Noordhuis (2005) reported that Erasmus observed a toad, who, after he was bitten by a spider, took instantly a leave of plantago. Wildlife is specific in choosing fruit of a plant for their food and welfare (Dersal et al., 1938). Besides, food-web interplay, animals have been a source of learning for humanity on how to cure several bodily disorders with plants. Shepherds notice that sheep tend to eat dandelion flowers, plantago or burdock leaves when they were weak (Krief, 2012; Asseldonk, 2006). In the foothills of the Himalaya near Mt. Everest the use of the roots of 'chota-chand' as a potent antidote for a snake bite is said to have been learned by observing mongooses feeding on the plant before fighting with cobra (Balick and Cox, 1996). Berberis lyceum with active components viz; flavonoids, alkaloids including Berberine, tannins, saponins and triterpenoids has found effective for skin affections and wound healing in animals (Chakraborty et al., 2013; Mittal et al., 2013; Asif et al., 2007). A growing body of literature in the behavioral, ecological and pharmacological sciences suggests that animals use certain plants for the control of parasite infection and related illnesses (Cousins et al., 2002). There are evidences of animal self-medication Krief, 2012). However, we have little knowledge about the abilities of animals to self-medicate, and many of the observations are anecdotal and equivocal (Clayton and Wolfe, 1993; Lozano, 1998; Houston et al., 2001). Contrary to the observations of Lozano (1998) recorded here from hunters, shepherds and local mountain communities establish cause and effect to a greater extend that wildlife rely upon Berberis and other medicinal plants for certain hurts, injuries and other illnesses. These unlearned adaptability and accidental behaviour patterns in animals are neither learned nor
... A growing body of literature in the behavioral, ecological and pharmacological sciences suggests that animals use certain plants for the control of parasite infection and related illnesses (Cousins et al., 2002). There are evidences of animal self-medication (Engel, 2002; Huffman, 2006 Huffman, , 2003 Huffman, , 1997 Plotkin, 2000; Janzen, 1978; Burritt and Provenza, 2000; Provenza, 2001, 2002; Phy and Provenza, 1998; Villalba et al., 2010 Villalba et al., , 2006 Krief, 2012). However, we have little knowledge about the abilities of animals to self-medicate, and many of the observations are anecdotal and equivocal (Clayton and Wolfe, 1993; Lozano, 1998; Houston et al., 2001). ...
Article
Full-text available
Berberis, a member of family Berberidaceae serves as ‘wildlife-clinic’ in nature. Berberis pseudumbellata subsp. gilgitica is endemic to the area and has become critically endangered. For the first time research documented unique epigenetic behaviour of different mountain wildlife and zoopharmacological practices of traditional communities from CKNP. When wildlife gets physical hurts, it rushes towards Berberis plants and eat berries, chew leaves or bark, which is found significant (p<0.001). Such behaviour is untaught, not learned, and automatic but seems pre-programmed instinctively triggered reflexed. This is observed in avifauna (House Sparrow, Chakor, Ram Chakor) and angulates (Flare horned Markhor, Himalayan Ibex) only. 79.03%, 36.51% hunters (SD±30.02), 83.24%, 58.23% shepherds (SD±17.68) and 41.92%, 17.11% local population (SD±17.54) believe and have personal observations respectively for Berberis-wildpharma interaction. Traditionally it is used zoopharmacologically for bone healing of limbs (93.67%), other bone fracture and injury (90.91%), other internal injuries (85.53%), external injuries (73.33%), delivery of sheep (38.12%), goat (17.36%) and cow (19.87%). No such observation was recorded with respect to herpetofauna and carnivores.
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Domestic sheep (Ovis aries) are among the earliest animals domesticated for human use. They are consumed worldwide as mutton, hogget, and lamb, kept as wool and milk producers, and used extensively in scientific research. The popular stereotype is that sheep are docile, passive, unintelligent, and timid, but a review of the research on their behavior, affect, cognition, and personality reveals that they are complex, individualistic, and social.
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Pasture-based production systems represent a significant sustainable supplier of animal source foods worldwide. For such systems, mounting evidence highlights the importance of plant diversity on the proper functioning of soils, plants and animals. A diversity of forages and biochemicals –primary and secondary compounds- at appropriate doses and sequences of ingestion, may lead to benefits to the animal and their environment that are greater than grazing monocultures and the isolated effects of single chemicals. Here we review the importance of plant and phytochemical diversity on animal nutrition, welfare, health, and environmental impact while exploring some novel ideas about pasture design and management based on the biochemical complexity of traditional and non-traditional forage sources. Such effort will require an integration and synthesis on the morphology, ecophysiology, and biochemistry of traditional and non-traditional forage species, as well as on the foraging behavior of livestock grazing diverse pasturelands. Thus, the challenge ahead entails selecting the “right” species combination, spatial aggregation, distribution and management of the forage resource such that productivity and stability of plant communities and ecological services provided by grazing are enhanced. We conclude that there is strong experimental support for replacing simple traditional agricultural pastures of reduced phytochemical diversity with multiple arrays of complementary forage species that enable ruminants to select a diet in benefit of their nutrition, health and welfare, whilst reducing the negative environmental impacts caused by livestock production systems.
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Self-medication behaviour is the use of natural materials or chemical substances to manipulate behaviour or alter the body’s response to parasites or pathogens. Self-medication can be preventive, performed before an individual becomes infected or diseased, and/or therapeutic, performed after an individual becomes infected or diseased. We summarized all available reports of self-medication in mammals and reconstructed its evolution. We found that reports of self-medication were restricted to eutherian mammals and evolved at least four times independently. Self-medication was most commonly reported in primates. Detailed analyses of primates suggest that self-medication is a life-history trait associated with body size, absolute brain size and longevity, but we found no support for the hypothesis that self-medication evolved to reduce the costs of social living. Large, longer-lived species might thus benefit uniquely from self-medication.
Article
Background Poisoning with Acer pseudoplatanus L. in horses contradicts the hypothesis of coexistence between plants and vertebrate herbivores being mediated through antipastoral traits as toxins. However, incidental observations showed that horses evaded Acer seedlings with primary leaves. The objective of the present cross-discipline study was (i) to analyse whether developmental stages of A. pseudoplatanus L. differed as to phenolics hypothesised as antipastoral traits, and (ii) to observe systematically the selection behaviour of pastured horses towards A. pseudoplatanus seedlings. Methods Phenolic profiles of five developmental stages from fruits to seedlings of progressing age up to adult leaves of A. pseudoplatanus and Acer campestre L. were characterised. Video recordings of grazing behaviour of 29 pastured horses towards seedlings of A. pseudoplatanus resulted into 117 sequences as additional field data. Results The horses ingested 19.1 per cent of juvenile seedlings with cotyledons (1.65 mg total phenolics/g fresh weight (FW), 82 compounds, 0.02 mg total gallic acid/g FW) yet only 5.46 per cent of older seedlings with primary leaves (8.48 mg total phenolics/g FW, 120 compounds, 3.13 mg total gallic acid/g FW). Conclusion Horses distinguished between seedlings in distinct stages that could be chemically distinguished, too. Acer seedlings with primary leaves provide a strong, but not complete antipastoral effect that correlates with dramatic changes in phenolic compounds.
Article
Pimelea poisoning of cattle, historically known as St. George Disease or Marree Disease, is a prevailing issue in arid grazing regions of inland Australia. Ingestion of the toxic native Pimelea species that contain the secondary metabolite simplexin, a diterpene orthoester with potent protein kinase C activity, induces diarrhoea, characteristic oedema and potentially fatal right-sided heart failure in cattle. Outbreaks of toxic Pimelea in the grazing field depend on seasonal prevalence. However, all stages of the plant carry the toxin, from seeds, juvenile plants to dead plant material. Livestock generally avoid consuming green Pimelea plants and only consume toxic Pimelea when pasture is minimal or where Pimelea growing through grass tussocks results in inadvertent ingestion. Our knowledge base of Pimelea poisoning has greatly improved with past research, yet the health hazards for livestock grazing in Pimelea affected pastures remains a significant issue whilst the ongoing search to develop effective strategies to mitigate poisoning continues. The goal of this review is to collate historical and recent research giving an overview of the current understandings of Pimelea poisoning, the toxin, its toxic effects and progress made towards remedies to alleviate the effects of Pimelea intoxication.
Article
Density‐dependent behavior underpins white‐tailed deer ( Odocoileus virginianus ) theory and management application in North America, but strength or frequency of the phenomenon has varied across the geographic range of the species. The modifying effect of stochastic environments and poor‐quality habitats on density‐dependent behavior has been recognized for ungulate populations around the world, including white‐tailed deer populations in South Texas, USA. Despite the importance of understanding mechanisms influencing density dependence, researchers have concentrated on demographic and morphological implications of deer density. Researchers have not focused on linking vegetation dynamics, nutrition, and deer dynamics. We conducted a series of designed experiments during 2004–2012 to determine how strongly white‐tailed deer density, vegetation composition, and deer nutrition (natural and supplemented) are linked in a semi‐arid environment where the coefficient of variation of annual precipitation exceeds 30%. We replicated our study on 2 sites with thornshrub vegetation in Dimmit County, Texas. During late 2003, we constructed 6 81‐ha enclosures surrounded by 2.4‐m‐tall woven wire fence on each study site. The experimental design included 2 nutrition treatments and 3 deer densities in a factorial array, with study sites as blocks. Abundance targets for low, medium, and high deer densities in enclosures were 10 deer (equivalent to 13 deer/km ² ), 25 deer (31 deer/km ² ), and 40 deer (50 deer/km ² ), respectively. Each study site had 2 enclosures with each deer density. We provided deer in 1 enclosure at each density with a high‐quality pelleted supplement ad libitum , which we termed enhanced nutrition; deer in the other enclosure at each density had access to natural nutrition from the vegetation. We conducted camera surveys of deer in each enclosure twice per year and added or removed deer as needed to approximate the target densities. We maintained >50% of deer ear‐tagged for individual recognition. We maintained adult sex ratios of 1:1–1:1.5 (males:females) and a mix of young and older deer in enclosures. We used reconstruction, validated by comparison to known number of adult males, to make annual estimates of density for each enclosure in analysis of treatment effects. We explored the effect of deer density on diet composition, diet quality, and intake rate of tractable female deer released into low‐ and high‐density enclosures with natural nutrition on both study sites (4 total enclosures) between June 2009 and May 2011, 5 years after we established density treatments in enclosures. We used the bite count technique and followed 2–3 tractable deer/enclosure during foraging bouts across 4 seasons. Proportion of shrubs, forbs, mast, cacti, and subshrubs in deer diets did not differ ( P > 0.57) between deer density treatments. Percent grass in deer diets was higher ( P = 0.05) at high deer density but composed only 1.3 ± 0.3% (SE) of the diet. Digestible protein and metabolizable energy of diets were similar ( P > 0.45) between deer density treatments. Likewise, bite rate, bite size, and dry matter intake did not vary ( P > 0.45) with deer density. Unlike deer density, drought had dramatic ( P ≤ 0.10) effects on foraging of tractable deer. During drought conditions, the proportion of shrubs and flowers increased in deer diets, whereas forbs declined. Digestible protein was 31%, 53%, and 54% greater ( P = 0.06) during non‐drought than drought during autumn, winter, and spring, respectively. We studied the effects of enhanced nutrition on the composition and quality of tractable female deer diets between April 2007 and February 2009, 3 years after we established density treatments in enclosures. We also estimated the proportion of supplemental feed in deer diets. We used the 2 low‐density enclosures on each study site, 1 with enhanced nutrition and 1 with natural nutrition (4 total enclosures). We again used the bite count technique and 2–3 tractable deer living in each enclosure. We estimated proportion of pelleted feed in diets of tractable deer and non‐tractable deer using ratios of stable isotopes of carbon. Averaged across seasons and nutrition treatments, shrubs composed a majority of the vegetation portion of deer diets (44%), followed by mast (26%) and forbs (15%). Enhanced nutrition influenced the proportion of mast, cacti, and flowers in the diet, but the nature and magnitude of the effect varied by season and year. The trend was for deer in natural‐nutrition enclosures to eat more mast. We did not detect a statistical difference ( P = 0.15) in the proportion of shrubs in diets between natural and enhanced nutrition, but deer with enhanced nutrition consumed 7–24% more shrubs in 5 of 8 seasons. Deer in enhanced‐nutrition enclosures had greater ( P = 0.03) digestible protein in their overall diet than deer in natural‐nutrition enclosures. The effect of enhanced nutrition on metabolizable energy in overall diets varied by season and was greater ( P < 0.04) for enhanced‐nutrition deer during summer and autumn 2007 and winter 2008. In the enhanced‐nutrition treatment, supplemental feed averaged 47–80% of the diet of tractable deer. Of non‐tractable deer in all density treatments with enhanced nutrition, 97% ( n = 128 deer) ate supplemental feed. For non‐tractable deer averaged across density treatments, study sites, and years, percent supplemental feed in deer diets exceeded 70% for all sex and age groups. We determined if increasing deer density and enhanced nutrition resulted in a decline in preferred forbs and shrubs and an increase in plants less preferred by deer. We sampled all 12 enclosures via 20, 50‐m permanent transects in each enclosure. Percent canopy cover of preferred forbs was similar ( P = 0.13) among deer densities averaged across nutrition treatments and sampling years (low density: = 8%, SE range 6–10; medium density: 5%, 4–6; high density: 4%, 3–5; SE ranges are presented because SEs associated with backtransformed means are asymetrical). Averaged across deer densities, preferred forb canopy cover was similar between nutrition treatments in 2004; but by 2012 averaged 20 ± 17–23% in enhanced‐nutrition enclosures compared to 10 ± 8–13% in natural‐nutrition enclosures ( P = 0.107). Percent canopy cover of other forbs, preferred shrubs, other shrubs, and grasses, as well as Shannon's index, evenness, and species richness were similar ( P > 0.10) among deer densities, averaged across nutrition treatments and sampling years. We analyzed fawn:adult female ratios, growth rates of fawns and yearlings, and survival from 6 to 14 months of age and for adults >14 months of age. We assessed adult body mass and population growth rates (lambda apparent, λ APP ) to determine density and nutrition effects on deer populations in the research enclosures during 2004–2012. Fawn:adult female ratios declined ( P = 0.04) from low‐medium density to high density in natural‐nutrition enclosures but were not affected ( P = 0.48) by density in enhanced nutrition enclosures although, compared to natural nutrition, enhanced nutrition increased fawn:adult female ratios by 0.15 ± 0.12 fawns:adult female at low‐medium density and 0.44 ± 0.17 fawns:adult female at high density. Growth rate of fawns was not affected by deer density under natural or enhanced nutrition ( P > 0.17) but increased 0.03 ± 0.01 kg/day in enhanced‐nutrition enclosures compared to natural nutrition ( P < 0.01). Growth rate of yearlings was unaffected ( P > 0.71) by deer density, but growth rate increased for males in some years at some density levels in enhanced‐nutrition enclosures. Adult body mass declined in response to increasing deer density in natural‐nutrition enclosures for both adult males ( P < 0.01) and females ( P = 0.10). Enhanced nutrition increased male body mass, but female mass did not increase compared to natural nutrition. Survival of adult males was unaffected by deer density in natural‐ ( P = 0.59) or enhanced‐ ( P = 0.94) nutrition enclosures. Survival of adult females was greatest in medium‐density enclosures with natural nutrition but similar at low and high density ( P = 0.04). Enhanced nutrition increased survival of females ( P < 0.01) and marginally for males ( P = 0.11). Survival of fawns 6–14 months old was unaffected ( P > 0.35) by density in either natural‐ or enhanced‐nutrition treatments but was greater ( P = 0.04) under enhanced nutrition. Population growth rate declined ( P = 0.06) with increasing density in natural‐nutrition enclosures but not ( P = 0.55) in enhanced nutrition. Enhanced nutrition increased λ APP by 0.32. Under natural nutrition, we found only minor effects of deer density treatments on deer diet composition, nutritional intake, and plant communities. However, we found density‐dependent effects on fawn:adult female ratios, adult body mass, and population growth rate. In a follow‐up study, deer home ranges in our research enclosures declined with increasing deer density. We hypothesized that habitat quality varied among home ranges and contributed to density‐dependent responses. Variable precipitation had a greater influence on deer diets, vegetation composition, and population parameters than did deer density. Also, resistance to herbivory and low forage quality of the thornshrub vegetation of our study sites likely constrained density‐dependent behavior by deer. We posit that it is unlikely that, at our high‐density (50 deer/km ² ) and perhaps even medium‐density (31 deer/km ² ) levels, negative density dependence would occur without several wet years in close association. In the past century, this phenomenon has only happened once (1970s). Thus, density dependence would likely be difficult to detect in most years under natural nutrition in this region. Foraging by deer with enhanced nutrition did not result in a reduction in preferred plants in the vegetation community and had a protective effect on preferred forbs because ≤53% of deer diets consisted of vegetation. However, enhanced nutrition improved fitness of individual deer and deer populations, clearly demonstrating that nutrition is limiting for deer populations under natural conditions in western South Texas. © 2019 The Authors. Wildlife Monographs published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.
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OBJECTIVE: In rodents, progesterone (P4) pretreatment increases anxiety and response to stressors. Social isolation is a stressor that generates physiological and behavioural stress responses in sheep. The aim of the study was to compare the stress response of anoestrous ewes previously treated or not with P4 to the social isolation test. DESIGN: Ten ewes received P4 treatment during 13 d (group P4-W) and another 10 remained untreated as controls (group Con). The ewes were individually isolated in a novel place during 10 min, 24 h after the end of P4 pretreatment and their behaviours were recorded. Cortisol and P4 concentrations as well as body surface temperature were recorded before and after the test. RESULTS: Ewes of the P4-W group presented higher Cortisol levels 0, 10, 20 and 30 min after the social isolation and had greater area under the curve of Cortisol compared to Con ewes (41,785%±4,156% vs. 25,682%±4,565% during 75 min). Progesterone and body surface temperature increased after social isolation, with no differences between P4-W and Con ewes. There were no differences in behavioural responses to social isolation. CONCLUSIONS: P4 pretreatment appears to augment the stress response to social isolation in anoestrous ewes.
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Biodiversité et santé comble le fossé entre l’écologie de la santé et les concepts issus des initiatives internationales comme EcoHealth, One Health ou Planetary Health. La biodiversité et la santé offrent une occasion unique de montrer comment les sciences écologiques, les sciences de l’environnement, les sciences médicales et les sciences sociales peuvent contribuer à améliorer la santé et le bien-être humains en préservant la biodiversité et ses services rendus aux sociétés. Cet ouvrage offre un aperçu général et intégré des disciplines scientifiques contribuant à lier la santé à la biodiversité : de l’écologie évolutive des maladies infectieuses et non infectieuses à l’éthique, au droit, et aux politiques publiques.
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Palates link animals with landscapes. An attuned palate, which enables animals to meet needs for nutrients and self-medicate, evolves from flavour-feedback associations, availability of biochemically rich foods, and learning in utero and early in life to eat nourishing combinations of foods. Unlike our ancestors who ate biochemically rich diets, the palates of many individuals are no longer linked in healthy ways with landscapes. Selection for yield, appearance and transportability diminish phytochemical richness of vegetables and fruits, which adversely affects the flavour and nutritive value of produce for humans. Likewise, phytochemically impoverished pastures and feedlot diets can unfavourably affect the health of livestock and the flavour and nutritive value of meat and dairy for humans. Not coincidentally, as the flavours of meat, dairy and produce became blander, processed foods became more desirable as people in industry learned to link feedback from energy-rich compounds with artificial flavours that obscure nutritional sameness and diminish health. Thus, the roles plants and animals once played in nutrition and health have been usurped by processed foods fortified and enriched in ways that adversely affect preferences by stimulating appetite for processed over wholesome foods. The health of soil, plants, herbivores and humans could be improved by creating cultures that know how to produce and combine biochemically rich foods into meals that nourish and satiate.
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Ruminants select nutritious feeds and avoid deleterious ones by relying on a learning process. We tested the hypothesis that they may similarly be able to select feeds for their medicinal properties, i.e. to self-medicate. Our experimental model was lambs infected with gastrointestinal nematodes (GIN), and the temperate legume sainfoin (Onobrychis viciifolia) containing condensed tannins (CT). Parasitism by GIN is a major constraint of grazing livestock and interest is growing for bioactive forages with anthelminthic (AH) properties, as an alternative to chemotherapy. We predicted that parasitized lambs will express greater intake and preference for tannin-rich sainfoin than non-parasitized lambs, and that lambs will adjust sainfoin intake as a function of their parasitic status. We also tested the importance of a conditioning period in helping parasitized lambs to experience the beneficial effects of tannin-rich sainfoin. We used 40 ewe lambs and two types of sainfoin pellets with low (2.16%; T-) or moderate (4.06%; T+) CT contents. Lambs were split into two groups (n = 20), one being kept non-parasitized (NP) while the other was experimentally infected with 3000 larvae of the blood-sucking nematode Haemonchus contortus (parasitized, P). Preference tests (4-d long) between T+ and T- were performed before infection (Test 1), after parasite development (Test 2), after conditioning (21 day long period with only T+ offered, Test 3), and after change in parasitic status (Test 4). Lambs' health status was monitored via faecal egg counts and haematocrit. We did not observe a self-medication behaviour as P and NP lambs expressed similar T+ preferences in all tests (Group, P = 0.98; Group x Test, P = 0.92), and P lambs did not increase their T+ preference from Tests 1 to Test 2 (0.27 and 0.23 ± 0.036, respectively). Initially, lambs expressed an aversion towards T+, but they switched to a preference after conditioning (Test 3: 0.62 ± 0.036), which was maintained after chemical deworming (Test 4: 0.66 ± 0.059) (Test, P < 0.0001). Collectively, these results indicate that (1) the moderate CT content in T+ was sufficient to induce an initial sensorial aversion; (2) T+ induced a positive reward in all lambs, whatever their parasitic status, which outweighed its potential AH effects; and (3) the conditioning period was important as it gave animals the opportunity to overcome their initial reluctance to consume a CT-containing forage, facilitating the learning process and allowing memorization of the sensorial and post-ingestive characteristics of the feed. Even if animals may not use temperate fodder legumes with moderate CT contents autonomously in a curative way, the absence of a trade-off between positive (medicinal, nutritional) and negative (toxicity of CT) effects may favour their use prophylactically, once animals have learned about their post-ingestive consequences.
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Plant secondary metabolites can adversely affect cellular and metabolic processes in herbivores, but also at low doses and in appropriate mixtures, they can have beneficial effects on animal nutrition, health and other therapeutic impacts. In this paper, we demonstrate the potential effects of plants with medicinal properties on animal foraging behavior as a function of the consequences they experience after ingestion of Mediterranean shrubs that are rich in phytotoxins. This mechanism - behavior by consequences - suggests animals are able to meet nutritional requirements and self-select certain plants with medicinal properties if they are offered the opportunity to do so when foraging in diverse Mediterranean shrub communities. Understanding the feeding behavior of ruminants when offered a variety of plant species is necessary to be able to improve their health and well-being by reducing levels of stress and fear; it may also lead to the early detection of illness. Thus, management strategies in biochemically/biologically diverse ecosystems should benefit from allowing ruminants to manifest their feeding preferences.
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Objective: In rodents, progesterone (P4) pretreatment increases anxiety and response to stressors. Social isolation is a stressor that generates physiological and behavioural stress responses in sheep. The aim of the study was to compare the stress response of anoestrous ewes previously treated or not with P4 to the social isolation test. Design: Ten ewes received P4 treatment during 13 d (group P4-W) and another 10 remained untreated as controls (group Con). The ewes were individually isolated in a novel place during 10 min, 24 h after the end of P4 pretreatment and their behaviours were recorded. Cortisol and P4 concentrations as well as body surface temperature were recorded before and after the test. Results: Ewes of the P4-W group presented higher cortisol levels 0, 10, 20 and 30 min after the social isolation and had greater area under the curve of cortisol compared to Con ewes (41,785%±4,156% vs. 25,682%±4,565% during 75 min). Progesterone and body surface temperature increased after social isolation, with no differences between P4-W and Con ewes. There were no differences in behavioural responses to social isolation. Conclusions: P4 pretreatment appears to augment the stress response to social isolation in anoestrous ewes.
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Purpose – The purpose of this paper is to suggest a practical means of incorporating ecological capital into the framework of business entities. Investors and shareholders need to be informed of the viability and sustainability of their investments. Ecological (natural) capital risks are becoming more significant. Exposure to material risk from primary industry is a significant factor for primary processing, pharmaceutical, textile and the financial industry. A means of assessing the changes to ecological capital assets and their effect on inflows and outflows of economic benefit is important information for stakeholder communication. Design/methodology/approach – This paper synthesises a body of literature from accounting, ecological economics, ecosystem services, modelling, agriculture and ecology to propose a way to fill current gaps in the capability to account for ecological capital. It develops the idea of the ecological balance sheet (EBS) to enable application of familiar methods of managing built and financial capital to management of ecological assets (ecosystems that provide goods and services). Findings – The EBS is possible, practical and useful. A form of double-entry bookkeeping can be developed to allow accrual accounting principles to be applied to these assets. By using an EBS, an entity can improve its capability to increase inflows and avoid future outflows of economic benefit. Social implications – Although major efforts are under-way around the world to improve business impact on natural resources, these efforts have been unable to satisfactorily help individual businesses elucidate the practical economic and competitive advantages conferred by investment in ecological capital. This work provides a way for businesses to learn about what the impact of changes to ecological assets has on inflows and outflows of economic benefit to their enterprise and how to invest in ecological capital to reduce their enterprise’s material risk and create competitive advantage. Originality/value – No one has synthesised knowledge and practice across these disciplines into a practical approach. This approach is the first demonstration of how ecological assets can be managed in the same way as built capital by using proven practices of accounting.
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The aim of this study was to determine the rate, variability and repeatability of intake by grazing sheep of a medicated feed block (MFB) containing fenbendazole and investigate if infection with gastrointestinal nematodes altered consumption patterns of the MFB in the same grazing mob. In Experiment 1, 30 Merino wethers were given access to an MFB for two separate 1-week periods, with blood sampling at Days 2, 4 and 6 of each period to determine MFB intake. In Experiment 2, the wethers were selected based on previous MFB intake and allocated to receive an oral dose of 10 000 Trichostrongylus colubriformis and 3000 Haemonchus contortus (anthelmintic susceptible) or a long acting anthelmintic. After 5 weeks, sheep were given access to an MFB (1.5 mg fenbendazole/g) and eight blood samples were taken over 2 weeks to determine intake. In Experiment 1, individual MFB intake in Week 1 and Week 2 was positively correlated (P = 0.002, R-2 = 0.287). Mean individual MFB intake in Experiment 1 and Experiment 2 was positively correlated (P = 0.008, R-2 = 0.047). In Experiment 2, more infected wethers (95%) consumed the MFB than did uninfected wethers (79%) (P < 0.001) and infected wethers ate significantly more MFB over the first 4 days (P = 0.041) of access. All infected sheep consumed sufficient MFB to receive a therapeutic dose and worm egg counts in infected sheep declined from 2165 epg to 120 epg in the first week of access to MFB. The decline in differences in MFB intake between infected and uninfected sheep corresponded to the decline in worm egg count, suggesting the existence of self-medication with parasitism accounting for intake differences.
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Vitamin E deficiency is common in sheep during summer and autumn in Mediterranean environments because of the lack of green feed. Deficiency of Vitamin E can lead to the development of nutritional myopathy, a condition causing heart and skeletal muscle damage which, in severe cases, can lead to death of the animal. Saltbush (Atriplex spp.) contains high concentrations of Vitamin E, so providing sheep with access to saltbush during summer may improve their Vitamin E status and prevent Vitamin E deficiency. We wished to determine whether backgrounding lambs on saltbush over summer and autumn (i.e. graze saltbush-based pastures for several weeks before finishing them to condition suitable for slaughter) would prevent Vitamin E deficiency and nutritional myopathy and compared the effectiveness of this strategy in preventing Vitamin E deficiency to a commercially available synthetic Vitamin E supplement. Ten-month-old cross-bred lambs (n ≤ 48) were backgrounded on dry, senesced (control) or saltbush-based pastures for 8 weeks during summer. After backgrounding they were fed a grain-based finishing ration containing low levels of Vitamin E for a further 5 weeks. We found that while grazing saltbush the plasma Vitamin E concentrations of lambs increased from 1.1 to 2.6 mg/L within 3 weeks, concentrations that were significantly higher than the concentrations in the lambs that did not have access to saltbush during backgrounding (P < 0.001). The improved Vitamin E concentrations corresponded with a reduction in the incidence of nutritional myopathy, with none of the lambs grazing saltbush showing any biochemical signs of myopathy, whereas 17% of lambs backgrounded on control pastures had elevated plasma concentrations of creatine kinase that were indicative of subclinical nutritional myopathy. During the subsequent finishing phase, lambs that had not had access to saltbush during backgrounding were all Vitamin E deficient and, of these, 8.5% were diagnosed with subclinical nutritional myopathy. By contrast, none of the lambs backgrounded on saltbush was Vitamin E deficient nor did they have any biochemical evidence of Vitamin E-responsive myopathy. The present study demonstrated that saltbush is a valuable source of Vitamin E for livestock that can reduce the incidence of subclinical nutritional myopathy in lambs during summer and prevent plasma Vitamin E concentrations becoming deficient for up to 5 weeks after saltbush is removed from the diet.
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The article which follows this introduction was originally published as a Special (Cover) Article in the American Journal of Psychiatry in the November, 1985 issue, the same month in which the First International Drug Symposium, sponsored by The Bahamas Ministry of Health and The Embassy of the United States of America, was convened to discuss the rock-cocaine epidemic in the Bahamas and other Caribbean Islands. Based on my article, I was invited to participate in the Symposium and to speak about some of my views on the psychological predispositions for drug dependence in general, and in particular, on the psychological predisposition for cocaine dependence. At first, I did not grasp the seriousness and scope of the cocaine problem, but I accepted the invitation, believing I might make a contribution to the Symposium. I was not long in attendance at the Symposium before I realized that the Bahamian citizens, professionals, and health care leaders were facing a major crisis as a consequence of the cocaine epidemic.
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Ansrnacr.-Mule deer (Odocoileus hemionus), domestic sheep (Oois aries), and American black bears (Ursus americanus) were fed quebracho (Schinopsis sp.) tannin to determine the contribution of salivary proteins to nitrogen- and fiber-digestive efficiencies and tannin metabolism. These values were compared to previously published values for laboratory rats (Ratfus rattus) and prairie voles (Microtus ochrogaster). Mule deer, black bears, and laboratory rats consumrrrg this condensed tannin produced tannin-binding salivary proteins that reduced fecal-nitrogen losses per unit of ingested tannin and reduced tannin metabolism relative to domestic sheep and prairie voles. Digestibility of the plant fiber was reduced significantly by tannins in domestic sheep, but not in mule deer. Although virtually all ingested tannin (98.3 + 5.0%) was recovered in feces of mule deer and black bears, ca. 25% was not recovered in feces of domestic sheep and presumably was metabolized. The defensive role of tannins as digestion inhibitors or toxins is dependent upon the molecular characteristics of the tannin interacting with the physiological capability of the animal. Results from one plant-mammal interaction cannot be used to interpret others without an understanding of the characteristics of the tannins and the physiology, ecology, and evolution of the animal.
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We gathered data on the amount, composition, and rate of ingestion of foods and soils by the provisioned Japanese macaques (Macaca fuscata fuscata) at Arashiyama, Japan. Behavioral observations spanned one year on 8 adult females, using focal animal sampling. We analyzed a subsample of their foods for nutritional and toxic secondary compound content. We also analyzed soils eaten by the macaques for several physical-chemical properties and tested their adsorption affinity to tannins and alkaloids. Geophagy occurred at a high rate of 2.97 g/indiv./day with an elevated frequency in the afternoon. About two-thirds of their foods (by fresh weight) were provisioned items, which are extremely rich in proteins and soluble carbohydrates. The soils that they ingested were generally poor in mineral elements, the bio-availability of which was low. The soils had a high adsorption capacity for plant alkaloids but were poorly absorptive for tannins. They were rich in clay minerals of proven buffering capacity. Geophagy at Arashiyama may improve the health of macaques via buffering gastric upset. We discuss the results from the viewpoint of several hypotheses on geophagy.
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Nonhuman primates represent a major component of the frugivore biomass in several rain-forest communities. Although there is considerable evidence that prosimians, monkeys, and apes serve as dispersal agents for many tropical trees, little attention has been paid to the more basic questions of why certain species of primates swallow and void seeds, and what, if any, are the advantages to an animal of having a large, hard, bolus pass through its digestive tract. We examine patterns of fruit-eating and seed-swallowing in two species of free-ranging tamarins: Saguinus mystax and Saguinus geoffroyi. Fruits commonly eaten by tamarins contain large seeds surrounded by a fibrous and adhesive pulp or arilate seed coat. They generally swallow seeds and pulp together. Intact seeds are voided over a 1- to 3-h period. Measurements of 132 seeds naturally voided by Panamanian tamarins average 11.2 mm in length and 0.3 g. The greatest number of large seeds contained in the digestive tract of a single animal at one time was 13. In the case of moustached tamarins, we collected 220 seeds. Average seed length is 11.9 mm and average seed weight is 0.3 g. At the time of capture, one animal had 26 seeds in its digestive tract. In both tamarin species, there is evidence of sex-based differences in feeding behavior. Adult female moustached and Panamanian tamarins swallowed and voided seeds of larger size than adult males did. Seed size is positively correlated with pulp weight (p
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When physiological adaptation is insufficient, hosts have developed behavioral responses to avoid or limit contact with parasites. One such behavior, leaf-swallowing, occurs widely among the African great apes. This behavior involves the slow and deliberate swallowing without chewing of whole bristly leaves. Folded one at a time between tongue and palate, the leaves pass through the gastro-intestinal (GI) tract visibly unchanged. Independent studies in two populations of chimpanzees (Pan troglodytes schweinfurthii) showed significant correlations between the swallowing of whole leaves and the expulsion of the nodule worm Oesophagostomum stephanostomum and a species of tapeworm (Bertiella studeri). We integrate behavioral, parasitological and physiological observations pertaining to leaf-swallowing to elucidate the behavioral mechanism responsible for the expulsion and control of nodule worm infections by the ape host. Physical irritation produced by bristly leaves swallowed on an empty stomach, increases motility and secretion resulting in diarrhea which rapidly moves leaves through the GI tract. In the proximal hindgut, the site of third-stage larvae (L3) cyst formation and adult worm attachment, motility, secretion and the scouring effect of rough leaves is enhanced by haustral contractions and peristalsis-antiperistalsis. Frequently, at the peak of reinfection, a proportion of nonencysted L3 is also predictably vulnerable. These factors should result in the disruption of the life cycle of Oesophagostomum spp. Repeated flushing during peak periods of reinfection is probably responsible for long-run reduction of worm burdens at certain times of the year. Accordingly, leaf-swallowing can be viewed as a deliberate adaptive behavioral strategy with physiological consequences for the host. The expulsion of worms based on the activation of basic physiological responses in the host is a novel hitherto undescribed form of parasitic control.
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Earth from a termite mound in the Mahale Mountains National Park, Tanzania, eaten by chimpanzees, was analyzed to determine the possible stimulus, or stimuli, for geophagy. The termite mound sample contains relatively high aluminum (10.0%), iron (3.0%), and sodium (0.5%). This correlates well with the mineralogy of the clay (<2 µm) fraction, which is high in metahalloysite, a 1:1 (Si:Al=1:1) clay mineral similar in chemical composition to the clay mineral kaolinite, and smectite (montmorillonite), which is a 2:1 expandable clay mineral. The combination of metahalloysite and smectite produces a substance much like the pharmaceutical Kaopectate™ widely used by humans as an anti-diarrheal agent. These analyses and preliminary observations linking geophagy with instances of severe diarrhea, and other signs of gastrointestinal upset in the Mahale chimpanzees, suggest that one function for the ingestion of this substance by chimpanzees may be to help provide temporary relief from gastrointestinal ailments. Further detailed investigations into the relationship between health and geophagy should provide important insights into the diverse roles of this behavior as a form of self-medication.
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Along with competition and predation, parasitism is one of the main sources of biotic stress facing all organisms. Recently, it has become recognized that animal diets may also be shaped by the need for protection from parasites. Foraging behavior evolves primarily to meet the need of a nutritionally adequate diet. However, just as foraging behavior can be affected by predators and competitors, some features of diet selection seem to have evolved to stave off or reduce parasitism. These adaptations can include the avoidance of foods that are also potential sources of parasitic infection, the use of prophylactic substances, and the consumption of therapeutic substances. This chapter incorporates self-medication into the broader phenomena—namely, the effects of plant chemicals across several trophic levels—and categorizes self-medicating behavior into two basic forms: prophylactic and therapeutic. It reviews evidence in the published literature for the occurrence of self-medication in nonhuman vertebrates. The chapter also discusses behavioral mechanisms that may play a role in self-medication and highlights potential implications for other areas of research.
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We studied the ability of lambs to select safe foods in the presence of harmful foods. In a series of feeding experiments, 3- to 6-mo-old lambs were offered a choice between 1) a familiar, safe and a novel, harmful food, 2) a novel, safe and a novel, harmful food and 3) a familiar, harmful and a novel, safe food. All harmful foods were palatable feeds that had been treated with lithium chloride (LiCl), a non-lethal gastrointestinal poison. When lambs ingested a meal composed of a novel food containing LiCl and a familiar, safe food, lambs subsequently avoided the novel food. Lambs avoided the novel food even when the familiar food contained LiCl. The response of lambs varied when lambs were given a choice between a novel, safe food and a novel food containing LiCl. Lambs either avoided the novel food containing LiCl and ingested the safe food or they limited their intake of both novel foods. Their response was dependent on the novelty of the food containing LiCl. When a novel, palatable food contained 2% LiCl, lambs always ate some of the food, even after experiencing illness from ingesting it. Lambs experienced with foods containing LiCl displayed greater food neophobia than lambs naive to LiCl-treated foods. Thus, novelty was the major criterion that lambs used to associate foods with gastrointestinal illness.
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We hypothesized that feed preference depends on the interplay between flavour and postingestive effects, and we tested two predictions based on this hypothesis: (1) lambs acquire preferences for poorly nutritious feeds paired with starch; and (2) preferences persist when starch is no longer administered. Twenty lambs were randomly allocated to two groups and conditioned as follows: on odd-numbered days, lambs in group 1 received onion-flavoured wheat straw and lambs in group 2 received oregano-flavoured wheat straw. On even-numbered days, the flavours were switched and starch (2.5-9.4% of the digestible energy received/d) was infused into the rumen of all animals during straw consumption. Four periods of 8 d of conditioning were performed and on the 9th day of each period all animals were offered a choice between onion- and oregano-flavoured straw. After conditioning, starch administration was suspended and lambs were offered onion- and oregano-flavoured straw at weekly intervals for 8 weeks (extinction). Lambs strongly preferred the flavoured straw paired with starch, and this preference persisted during extinction. Thus, these results suggest that the postingestive effects of energy play an important role in the development of feed preferences of ruminants.
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We suggested that food preference depends on the interplay between flavour and post-ingestive effects, and we predicted that protein-restricted lambs would acquire preferences for foods paired with supplemental sources of N, including urea (Expts 1 and 2), casein (Expt 3), and gluten (Expt 4). In each experiment, twenty lambs, in two groups of ten, were conditioned as follows: on odd-numbered days, lambs in group 1 received wheat straw (Expts 1, 3, and 4) or ground barley (Expt 2) flavoured with a distinctive flavour, and lambs in group 2 received the same food but with a different flavour. On even-numbered days, flavours were switched and lambs received capsules containing different amounts of urea (ranging from 0.12 to 0.92 g N/d), casein (ranging from 0.23 to 0.69 g N/d), or gluten (ranging from 0.23 to 0.69 g N/d). After conditioning period of 8 d, lambs were given a two-choice test to determine preference for flavours paired with N. In Expts 1 and 2, lambs preferred the flavours conditioned with urea at lower doses (0.12 g N/d in Expt 1, 0.23 and 0.46 g N/d in Expt 2), but they avoided the flavour associated with urea at the highest dose (0.23 g N/d in Expt 1 and 0.92 g N/d in Expt 2). In Expts 3 and 4, lambs avoided the flavours associated with the lowest doses of casein or gluten (0.23 g N/d), but they preferred the flavours paired with casein or gluten at higher doses (0.46 and 0.69 g N/d). After conditioning, N administrations were suspended and lambs in Expts 3 and 4 were offered a choice of the two flavours at weekly intervals for 2 weeks (extinction); preferences persisted during extinction. Collectively, these results suggest that the post-ingestive effects of N in different forms and concentrations influenced the development of food preferences by lambs.
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Tannins occur in many plant species, and they often suppress intake by reducing nutrient availability or by causing malaise. Polyethylene glycol (PEG) binds to tannins and may thereby increase the availability of macronutrients and decrease malaise. Supplemental PEG increases intake of tannin-containing plants by sheep, goats, and cattle. Given the strong response to supplemental PEG, we speculated that animals might self-regulate their intake of PEG when offered foods high in tannins. The objective of the first experiment was to determine if the amount of supplemental PEG (0, 25, 50, 75, or 100 g; molecular weight, 3,350) affected intake by lambs of a food (milo-tannin mix) containing 20% quebracho tannin. There was a linear relationship (Y = 272 + 1.2X; R2 = .86; P = .023) between the amount of supplemental PEG ingested and the subsequent intake of milo-tannin food by lambs. The objective of the second experiment was to determine whether lambs self-regulated intake of PEG when fed a ration that contained 0, 5, 10, 15, or 20% quebracho tannin and whether they adjusted their intake of PEG when tannin was removed from the diet. There was a positive relationship between the amount of PEG ingested and intake of food and tannin (P = .0001). Lambs fed high-tannin diets ate more PEG than controls (P = .03). Lambs fed the 20% tannin diet ate the most PEG, and controls ate the least PEG. Tannin limited intake of the diets, but PEG attenuated the response to a great degree (P = .065). Immediately after tannin was removed from the ration, lambs that formerly had been fed the 20% tannin ration ate more PEG than lambs fed the other rations (P = .0075). Ten of the lambs (5 from the 20% tannin group, 1 from the 15% tannin, and 2 each from the 10 and 5% groups) continued to eat PEG for 7 d after tannin was removed from their ration. When they were tested again 6 wk after the trial and offered tannin-free diets, their intake of PEG had decreased.
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Although we understand how fearful memories are stored in the brain, how they are extinguished remains a mystery. The answers may lie with the cannabinoid compounds our bodies produce.
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To quantify the use of self-treatment and to determine the actions taken to manage malaria illness. A cross-sectional study was undertaken in six peasant associations in Butajira district, southern Ethiopia, between January and September 1999. Simple random sampling was used to select a sample of 630 households with malaria cases within the last six months. Overall, 616 (>97%) of the study households acted to manage malaria, including the use of antimalarial drugs at home (112, 17.8%), visiting health services after taking medication at home (294, 46.7%), and taking malaria patients to health care facilities without home treatment (210, 33.3%). Although 406 (64.5%) of the households initiated treatment at home, the use of modern drugs was higher (579, 92%) than that of traditional medicine (51, 8%). Modern drugs used included chloroquine (457, 73.5%) and sulfadoxine-pyrimethamine (377, 60.6%). Malaria control programmes were the main sources of antimalarials. In most cases of malaria, treatment was started (322, 52.3%) or health services visited (175, 34.7%) within two days of the onset of symptoms. Cases of malaria in the lowland areas started treatment and visited health services longer after the onset of malaria than those in the midland areas (adjusted odds ratio, 0.44; 95% confidence interval (CI), 0.30-0.64; and adjusted odds ratio, 0.37; 95% CI, 0.25-0.56, respectively). Similarly, those further than one hour's walk from the nearest health care facility initiated treatment later than those with less than one hour's walk (adjusted odds ratio, 0.62; 95% CI 0.43-0.87). This might be because of inaccessibility to antimalarial drugs and distant health care facilities in the lowland areas; however, statistically insignificant associations were found for sex, age, and religion. Self-treatment at home is the major action taken to manage malaria. Efforts should be made to improve the availability of effective antimalarials to communities in rural areas with malaria, particularly through the use of community health workers, mother coordinators, drug sellers, and shop owners.
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Early in the co-evolution of plant-animal relationships, some arthropod species began to utilize the chemical defences of plants to protect themselves from their own predators and parasites. It is likely, therefore, that the origins of herbal medicine have their roots deep within the animal kingdom. From prehistoric times man has looked to wild and domestic animals for sources of herbal remedies. Both folklore and living examples provide accounts of how medicinal plants were obtained by observing the behaviour of animals. Animals too learn about the details of self-medication by watching each other. To date, perhaps the most striking scientific studies of animal self-medication have been made on the African great apes. The great ape diet is often rich in plants containing secondary compounds of non-nutritional, sometimes toxic, value that suggest medicinal benefit from their ingestion. Chimpanzees (Pan troglodytes), bonobos (Pan paniscus) and gorillas (Gorilla gorilla) are known to swallow whole and defecate intact leaves. The habit has been shown to be a physical means of purging intestinal parasites. Chimpanzees and man co-existing in sub-Saharan Africa are also known to ingest the bitter pith of Vernonia amygdalina for the control of intestinal nematode infections. Phytochemical studies have demonstrated a wide array of biologically-active properties in this medicinal plant species. In light of the growing resistance of parasites and pathogens to synthetic drugs, the study of animal self-medication and ethno-medicine offers a novel line of investigation to provide ecologically-sound methods for the treatment of parasites using plant-based medicines in populations and their livestock living in the tropics.
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Host-parasite interactions are often seen as an arms race, with parasites attempting to overcome host resistance to infection. Herbivory is a common route of transmission of parasites that represents the most pervasive challenge to mammalian growth and reproduction. The present paper reviews the foraging skills of mammalian herbivores in relation to their ability to exploit plant properties to combat parasites. The starting point is that foraging behaviour may ameliorate the impact of parasitism in three ways; hosts could: (1) avoid foraging in areas contaminated with parasites; (2) select diets which increase their resistance to parasites; (3) select for foods containing anti-parasitic properties (self-medication). Details are given of the pre-requisite skills needed by herbivores if they are to combat parasitism via behaviour, i.e. herbivores are able to: (a) determine their parasitic state and alter their behaviour in relation to that state (behaviours 1, 2 and 3); (b) determine the environmental distribution of parasites (behaviour 1); (c) distinguish plant species or plant parts that increase their resistance to parasites (behaviour 2) or have anti-parasitic properties (behaviour 3). Mammalian herbivores cannot detect the presence of the parasites themselves and must rely on cues such as faeces. Despite the use of these cues contacting parasites may be inevitable and so mechanisms to combat parasitism are necessary. Mammalian herbivores have the foraging skills needed to exploit the heterogeneous distributions of nutrients and parasites in complex foraging environments in order to avoid, and increase their resistance to, parasites. Current evidence for the use of plant secondary metabolites (PSM) by herbivores for self-medication purposes remains equivocal. PSM have both positive (anti-parasitic) and negative (toxic) effects on herbivores. Here details are given of an experimental approach using tri-trophic (plant-herbivore-parasite) interactions that could be used to demonstrate self-medication in animals. There is strong evidence suggesting that herbivore hosts have developed the foraging skills needed to take advantage of plant properties to combat parasites and thus use behaviour as a weapon in the host-parasite arms race.
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Chimpanzees in the wild swallow the rough hispid leaves of certain plant species as a means of physically expelling intestinal parasites. A plant with such a leaf texture was introduced in 36 trial sessions to a captive group of 11 healthy adult chimpanzees to investigate the possible origin and acquisition of leaf swallowing behavior. One male (housed separately from the group during testing) and one female, both captive born, spontaneously exhibited the behavior on their first trial without prior opportunity to observe others with this plant. Six other chimpanzees on their first trial displayed a phobic response to these leaves and rejected them entirely, while another two chewed and swallowed the leaves in a normal way. Four individuals eventually exhibited the behavior, after having approached and closely observed the leaf swallowing of the first female to exhibit the behavior in the group. Four of the six individuals that initially avoided the leaves never overcame their phobia toward this plant and were not in proximity to a chimpanzee performing leaf swallowing during test sessions. Individuals born to wild chimpanzee mothers were no more likely to perform the behavior than captive-reared group mates. These results suggest that the acquisition of this behavior is based in part on a propensity to fold and swallow rough, hispid leaves, but that the acquisition and spread of leaf swallowing within a group is likely to be socially influenced. This study provides support for the hypothesis that leaf swallowing originated in the wild from opportunistic feeding behavior and was later passed down in the form of a self-medicative behavioral tradition.
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Ruminants select nutritious diets from a diverse array of plant species that vary in kinds and concentrations of nutrients and toxins, and meet their nutritional requirements that vary with age, physiological state, and environmental conditions. Thus, ruminants possess a degree of nutritional wisdom in the sense that they generally select foods that meet nutritional needs and avoid foods that cause toxicosis. There is little reason to believe that nutritional wisdom occurs because animals can directly taste or smell either nutrients or toxins in foods. Instead, there is increasing evidence that neurally mediated interactions between the senses (i.e., taste and smell) and the viscera enable ruminants to sense the consequences of food ingestion, and these interactions operate in subtle but profound ways to affect food selection and intake, as well as the hedonic value of food. The sensation of being satisfied to the full (i.e., satiety) occurs when animals ingest adequate kinds and amounts of nutritious foods, and animals acquire preferences (mild to strong) for foods that cause satiety. Unpleasant feelings of physical discomfort (i.e., malaise) are caused by excesses of nutrients and toxins and by nutrient deficits, and animals acquire aversions (mild to strong) to foods that cause malaise. What constitutes excesses and deficits depends on each animal's morphology, physiology, and nutritional requirements. This does not mean that ruminants must maximize (optimize) intake of any particular nutrient or mix of nutrients within each meal or even on a daily basis, given that they can withstand departures from the normal average intake of nutrients (i.e., energy-rich substances, nitrogen, various minerals, and vitamins). Rather, homeostatic regulation needs only some increasing tendency, as a result of a gradually worsening deficit of some nutrient or of an excess of toxins or nutrients, to generate behavior to correct the disorder. Extreme states should cause herbivores to increase diet breadth and to acquire preferences for foods that rectify maladies. From an evolutionary standpoint, mechanisms that enable animals to experience feedback, sensations such as satiety and malaise, should be highly correlated with nutritional well being, toxicosis, and nutritional deficiencies, which are directly related with survival and reproduction.
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The effect of feeding quebracho tannin, a mixture of condensed tannins, on dietary nutrient utilisation and nitrogen (N) retention and its effects on the gastrointestinal (GI) tract was investigated in sheep and rats. Sheep (n = 24) were fed on a pelleted diet of dried grass alone (controls) or containing quebracho tannin at 50 g kg−1 diet dry matter (DM) (tannin-fed animals) at a level sufficient to achieve a daily liveweight gain (DLWG) of 100 g day−1. Complete collections of faeces and urine were made for two seven-day periods after two and six weeks of feeding these diets (n = 6 per group). Apparent digestibilities of dry matter, N and neutral detergent fibre (NDF) were significantly (P < 0.001) reduced in tannin-fed animals at both measurement periods. No evidence was obtained to suggest that rumen micro-organisms can adapt to the presence of dietary tannins with prolonged feeding. Tannin-fed animals excreted significantly (P < 0.01) more N in faeces and less in urine than controls suggesting an alteration in N metabolism. Histological examination of samples of the GI tract obtained from pairs of sheep slaughtered after two, five and seven weeks of feeding the diets indicated ulceration and an increase in mucosal histiocytes, particularly in the jejunum and ileum of most tannin-fed animals. In a subsequent experiment, rats were fed ad libitum a ground chow containing either cellulose or quebracho tannin at 40 g kg−1 DM. Tannin-fed rats had significantly (P < 0.05) reduced feed intakes, DLWG, N retention and body fat deposition compared to controls. Protein synthesis rates in the duodenal mucosa were not increased in tannin-fed rats suggesting that enterocyte proliferation was not stimulated in this region of the GI tract.These studies indicate that feeding quebracho tannin to ruminants has both ruminal and post-ruminal effects that, together, result in reduced nutrient utilisation and impaired animal performance.© 1999 Society of Chemical Industry
Article
In an earlier study it was observed that the rumen microbes from cattle which had never consumed tannin-containing diets do not have enzymes for degrading condensed tannins. In this study, rumen microbes were exposed to small amounts of quebracho tannins for 8 days using a rumen simulation technique (RUSITEC). The levels maintained in the RUSITEC were 0.1, 0.2 and 0.4 mg of the spray-dried quebracho powder (SDQT) per ml of the medium. After 8 days of tannin exposure, the liquor containing ‘exposed/adapted’ microbes from RUSITEC was incubated for 40 h without and with purified tannins of quebracho and Dichostachys cinerea leaves in in vitro gas method. There was no degradation of condensed tannins. The enzymes for degradation of condensed tannins were not induced in rumen microbes by exposure to different concentrations of tannins for 8 days in the RUSITEC. In the RUSITEC, SDQT significantly reduced the number of total protozoa, entodiniomorphs and holotrichs; effect was higher on holotrichs. There was no significant change in the levels of short chain fatty acid but the molar proportion of propionate was significantly higher and of butyrate significantly lower at 0.4 mg SDQT ml−1. Significantly lower levels of ammonia in the medium was also observed on injection of tannins. Microbial mass production, calculated using 15N incorporation, was similar at 0.1 and 0.2 mg SDQT ml−1 but significantly lower (13%) at 0.4 mg SDQT ml−1. The dry matter digestibility of the feed (80% hay and 20% barley) was not significantly affected by SDQT.
Article
In vitro studies were made to determine the nature of complexes formed between the condensed tannins (i.e. pro-anthocyanidins or flavolans) of sainfoin (Onobrychis viciifolia) and either the major soluble dietary protein, Fraction 1 protein, of green leaves, or bovine salivary mucoprotein. Fraction 1 protein, uniformly labelled with 14C, formed only insoluble complexes with these tannins. Bovine salivary mucoprotein produced insoluble complexes only at temperatures below 25°C. Fraction 1 protein complexes were stable between pH 3.5 and pH 7.0. At pHs below 3.0 greater than 90% of the protein was solubilised in the presence of PEG but at pH 8.0 only 30% of the protein was released from the complex. In vivo experiments with sheep fed sainfoin, confirmed the stability of the tannin-Fraction 1 complexes in the rumen (pH 6.5) and break-up of the complexes in anterior duodenal samples (pH 2.5) as shown by the extractability of condensed tannin from the complex. The significance of these results with respect to (a) bloat in cattle, (b) nitrogen metabolism of ruminants, is discussed. Polyethylene-glycol, mol.wt. 4000 (PEG) exchanged with protein in the complex releasing protein into solution. The effectiveness of PEG was dependent on the amount of tannin in the complex and the age of the complex at the time of addition of PEG. The results explain the observation that PEG is unsuitable as a marker for rumen volume determination when animals are fed herbage that contains condensed tannins. A method for determining any deleterious effects of tannins on nitrogen metabolism by ruminants, is suggested.
Article
This study tests hypotheses on the biochemical functions of geophagy in parrots: mechanical enhancement of digestion, acid buffering capacity, mineral supplementation, adsorption of dietary toxins, and gastrointestinal cytoprotection. Parrots showed clear preferences for specific soil horizons. Comparisons of preferred and nonpreferred soils from several sites suggest that soils have little ability to enhance grinding and no measurable ability to buffer gastric pH. Soils offered insignificant mineral supplementation since most minerals occurred at similar levels in samples regardless of preference, and the minerals were generally more plentiful in the birds' diets. Sodium was available in moderate levels at some sites (>1000 ppm), but was well below sodium detection thresholds of parrots. X-ray diffraction, cation exchange capacity, and in vitro adsorptive trials showed that the preferred soils are capable of exchanging substantial quantities of cations and are capable of adsorbing low-molecular-weight secondary compounds. In captive Amazona parrots, orally administered clay reduced the bioavailability of the alkaloid quinidine by roughly 60%, demonstrating that in vivo adsorption of potentially toxic compounds may be a biologically important function of geophagy. Labeled clay remained in the lower gastrointestinal tract of captive parrots for >12 hr, which along with high adsorptive capacities, further suggests a potential role in protecting the gastrointestinal lining from various biological and chemical insults. Detoxification and cytoprotection are the most likely functions of geophagy for parrots and herbivores with similar ecologies. Given the variety of chemically defended seeds consumed by these herbivores, geophagy likely protects consumers from dietary toxins, allowing increased diet breadth and/or enhancing digestibility.
Article
We studied the effects of mother's preference and lamb age on selection of Amelanchier alnifolia and Cercocarpus montanus by lambs. Thirty-six ewes and their lambs were divided into treatment and control groups. Mothers in the treatment group were conditioned to eat A. alnifolia, but avoid C. montanus, by pairing ingestion of C. montanus with lithium chloride (LiCl), a compound that causes gastrointestinal distress. Mothers in the control group were allowed to eat both foods with impunity. After ewes were conditioned, lambs were exposed daily for 5 days with their mothers to either A. alnifolia or C. montanus for 5 min followed by a 5 min exposure to the other food. Half of the lambs in the treatment and control groups were 6 weeks of age during exposure, while the other half were 12 weeks of age. Treatment lambs of both ages ate less C. montanus than control lambs during exposure. Lambs were weaned after exposure and tested 10 days later for preference. Preference indices (PI) for C. montanus were calculated as the percentage of total bites contributed by C. montanus. Treatment lambs exposed at 6 weeks of age ate less C. montanus than A. alnifolia compared with control lambs during trials of 5 min day−1 for 7 days (PI=0.07 vs. 0.44; P<0.05). Treatment lambs exposed at 12 weeks of age also ate less C. montanus than A. alnifolia compared with control lambs (PI=0.26 vs. 0.45; P<0.05). Lambs that were 12 weeks of age during exposure, however, ate more C. montanus than lambs exposed at 6 weeks of age (PI=0.26 vs. 0.07; P<0.05). Nine weeks after exposure, treatment lambs exposed at 6 weeks of age still showed lower preference for C. montanus than for A. alnifolia compared with control lambs (PI=0.10 vs. 0.48; P<0.05). Treatment and control lambs that were 12 weeks of age during exposure, however, showed equal preference for C. montanus and A. alnifolia (PI=0.42 vs. 0.42; P>0.05). We conclude that lambs 6 weeks of age were influenced more by their mothers dietary habits than were lambs 12 weeks of age.
Article
Not all pharmacists are human; other species also use medicinal substances to combat pathogens and other parasites. Self-medicating behaviour is a topic of rapidly growing interest to behaviourists, parasitologists, ethnobotanists, chemical ecologists, conservationists and physicians. Although most of the pertinent literature is anecdotal, several studies have now attempted to test the adaptive function of particular self-medicating behaviours. We discuss the results of these studies in relation to simple hypotheses that can provide a framework for future tests of self-medication.
Article
African Elephants Loxodonta africana regularly eat soil. At some sites, such as Mount Elgon on the Kenya-Uganda border, extensive caves have been excavated in the mountainside by their quarrying activity (Redmond, 1984; Bowell, 1992). The material eaten often has little organic content, and this might better be described as a weathered, soft, friable rock, but for convenience we refer to it here as soil. Such soils have been shown to offer sodium, calcium and other mineral nutrients which may be lacking from their normal plant diet (Weir, 1969, 1972, 1973 Moe, 1992; Bowell, 1992; Eksteen & Bornman, 1990). Elephants living in the cloud forest on the eastern escarpment of Ngorongoro in northern Tanzania regularly visit a number of low cliffs to prise away lumps of soil which they eat. This material does not taste 'salty' to the human tongue, and we present here an analysis of this material which suggests that one of its functions may be to assist the animals in the digestion of forest browse through its ability to detoxify the high concentration of plant secondary compounds found in tropical forest trees.
Article
The use of polyethylene glycol in deter- mining rumen fluid volume of cows fed liberal amounts of roughage and grain (control) or high-grain, low-fiber diets was studied. The polyethylene glycol method accurately measured rumen fluid volume in cows fed either diet. The rumen fluid turnover time was 5.0 and 11.2 hr for animals fed control and high-grain, low-fiber diets. For cows fed the control diet, 20.6% of the tureen fluid volume turned over every hour while the mean for cows fed the high-grain, low-fiber diet was 9.6%.
Article
Ruminants eat an array of plant species that vary in nutrients and toxins. This selection makes intuitive sense, but no theories adequately explain this diversity. Some maintain it reduces the likelihood of overingesting toxins, whereas others contend it meets nutritional needs. Nevertheless, herbivores seek variety even when toxins are not a concern and nutritional needs are met. I offer another explanation for this behavior, one which encompasses the avoidance of toxins and the acquisition of nutrients. A key concept in this theory is aversion, the decrease in preference for food just eaten as a result of sensory input (a food's taste, odor, texture, i.e., its flavor) and postingestive effects (effects of nutrients and toxins on chemo-, osmo-, and mechano-receptors) unique to each food. Aversions are pronounced when foods contain toxins or high levels of rapidly digestible nutrients; they also occur when foods are deficient in specific nutrients. Aversions occur even when animals eat nutritionally adequate foods because satiety (satisfied to the full) and surfeit (filled to nauseating excess) represent points along a continuum, and there is a fine line between satiety and aversion. Thus, eating any food is likely to cause a mild aversion, and eating a food too frequently or in excess is likely to cause a strong aversion. Aversions are involuntary and are not the result of conscious decisions by an animal. Aversions yield benefits (e.g., obtain a balanced diet, reduce ingestion of toxic foods, optimize foraging and rumination times, sample foods, maintain a diverse microflora in the rumen) that are often mistaken as the cause of varied diets. In this article, I discuss the subtle ways in which aversions diminish preference and cause animals to eat a variety of foods.
Article
We hypothesized that volatile fatty acids are feedback signals that condition food preferences in ruminants, and we tested two predictions based on this hypothesis: 1) low doses of propionate condition preferences for low-quality foods (Exp. 1 and 2) preferences are not caused by osmotic load (Exp. 2). In Exp. 1, lambs were offered chopped wheat straw flavored with either oregano or onion on odd days, whereas on even days flavors were switched and lambs received capsules containing sodium propionate. During four 8-d conditioning periods, the amounts of propionate delivered ranged from .7 to 1.4% of the daily DE intake (Period 1) or were fixed at .7% (Period 2) and 1% of the daily DE intake (Periods 3 and 4). After each 8-d conditioning period, lambs were offered oregano- and onion-flavored straw. Conditioning was then suspended and lambs were offered onion- and oregano-flavored straw at weekly intervals for 1 mo (extinction). Lambs preferred the flavor paired with propionate during conditioning (P < .001) and extinction (P < .07). During Exp. 2, a different group of lambs was conditioned as in Exp. 1, but sodium chloride was delivered at osmotic loads equivalent to those when propionate supplied .7% and 1% of the daily DE intake. Lambs strongly avoided the flavor paired with sodium chloride (P < .001). Thus, lambs acquired preferences for straw conditioned with doses of propionate typically considered ineffective in the regulation of food intake, and osmolalities generated by propionate did not cause, but probably attenuated, food preferences.