Low resting metabolic rate and increased hunger due to ß-MSH and ß-endorphin deletion in a canine model.

Mutations that perturb leptin-melanocortin signaling are known to cause hyperphagia and obesity, but energy expenditure has not been well studied outside rodents. We report on a common canine mutation in pro-opiomelanocortin (POMC), which prevents production of ß-melanocyte-stimulating hormone (ß-MSH) and ß-endorphin but not α-MSH; humans, similar to dogs, produce α-MSH and ß-MSH from the POMC propeptide, but rodents produce only α-MSH. We show that energy expenditure is markedly lower in affected dogs, which also have increased motivational salience in response to a food cue, indicating increased wanting or hunger. There was no difference in satiety at a modified ad libitum meal or in their hedonic response to food, nor disruption of adrenocorticotropic hormone (ACTH) or thyroid axes. In vitro, we show that ß-MSH signals comparably to α-MSH at melanocortin receptors. These data implicate ß-MSH and ß-endorphin as important in determining hunger and moderating energy expenditure and suggest that this role is independent of the presence of α-MSH.

"Feel the Force"-Prevalence of Subjectively Assessed Saddle Fit Problems in Swiss Riding Horses and Their Association With Saddle Pressure Measurements and Back Pain.

Ill-fitting saddles can impair the well-being and performance of horses. Saddle fit is generally assessed subjectively by a trained professional or with an electronic saddle pressure mat, but little is known about the agreement between both methods. The study aims were (1) to assess the prevalence of saddle fit issues in a riding sound Swiss horse population, (2) to investigate how well the subjective assessment correlates with objectively measured pressure magnitude and distribution under the saddle during riding, and (3) how well both correlate with back pain of the horse. Only 10% of the saddles were free of the assessed problems. Pressures exceeded clinically relevant thresholds in 15% of the horses. There was no clear correlation between back pain and pressure magnitude, but back pain was associated with certain subjectively assessed fit problems. Statistically significant associations between fit problems and the expected pressure patterns were found for panel angles, curvature of the saddle, width of the panel channel, and the waist of the saddle. There was no or limited association of pressure patterns with the balance of the saddle, width and angle of the tree head, or the symmetry of the panels. The results revealed that certain fit problems were reflected in the electronically measured pressure distribution and that the subjective assessment can therefore provide relevant information. Pressure magnitude showed only limited association with back pain, which indicates that there are other factors involved in the development of back problems in horses.

Riding Soundness-Comparison of Subjective With Objective Lameness Assessments of Owner-Sound Horses at Trot on a Treadmill.

Lameness is a symptom indicative of pain or injury of the locomotor apparatus. Lame horses generally should not be ridden. However, owners' ability to assess lameness has been questioned. This study's aim was to use subjective lameness assessments and objective gait analysis to generate a descriptive overview of movement and weight-bearing asymmetries of owner-sound riding horses. 235 horses were subjectively assessed in a field study, and the owner's perception of their horse's orthopedic health was recorded through an online survey. 69 horses were re-evaluated by gait analysis at an equine hospital. During trot on an instrumented treadmill, the gait was scored by a veterinarian using lameness grades from 0/5 (sound) to 3/5 (moderate lameness visible at trot). Movement asymmetry of the head (HDmin) and pelvis (PDmin) and weight-bearing asymmetry were quantified simultaneously. The prevalence of subjectively scored lameness grade ≥2/5 in one or more limbs was 55% during study part 1 and 74% during study part 2. Movement asymmetry of the head and/or pelvis exceeding HDmin ≥12 mm and/or PDmin ≥6 mm was found in 57% of the horses. 58% showed weight-bearing asymmetries between contralateral front and/or hind limbs of ≥3% body mass. Gait analysis showed considerable variability of movement and weight-bearing asymmetry values, sometimes independent of the clinical lameness grade, especially in the forehand. Several horses with lameness grade ≤1/5 had asymmetry values greater than mentioned thresholds. The analysis of movement and weight-bearing asymmetry revealed that these objective variables did not necessarily act uniformly and therefore should be interpreted with caution.

Husbandry, Use, and Orthopedic Health of Horses Owned by Competitive and Leisure Riders in Switzerland.

The use of horses in competitive sports receives increasing criticism from the public, mainly due to the potential for injury. However, it is unclear if orthopedic and other health issues are more common in competition horses than those in leisure horses. The aim of this study was to assess husbandry, use, and orthopedic health in Swiss riding horses and to compare these aspects between horses owned by self-identified competitive riders (CR) and leisure riders (LR) in Switzerland. A total of 237 owners completed an online survey providing information on their athletic ambitions, their horse's husbandry, health, training, and tack. Two experienced veterinarians assessed gait irregularities, muscular development, and back pain in the horses and evaluated saddle fit. Compared with horses owned by competitive riders (CH), a higher proportion of horses kept by leisure riders (LH) were kept unshod, under more natural conditions, and turned out with other horses. LH were exercised less frequently, and LR trained less frequently with instructors. CR reported less time since the last saddle check and the use of more training aids during riding. No differences between the two groups could be found in orthopedic health, muscular development, or back pain, but LH had higher body condition scores and a slightly higher proportion of saddles with at least one fit problem. Our data revealed no increased prevalence of the assessed health problems in competition horses compared with leisure horses in Switzerland. However, suboptimal saddle fit and muscular development, back pain, and gait irregularity are frequent in both groups and deserve more attention.

Influence of Functional Rider and Horse Asymmetries on Saddle Force Distribution During Stance and in Sitting Trot.

Asymmetric forces exerted on the horse's back during riding are assumed to have a negative effect on rider-horse interaction, athletic performance, and health of the horse. Visualized on a saddle pressure mat, they are initially blamed on a nonfitting saddle. The contribution of horse and rider to an asymmetric loading pattern, however, is not well understood. The aim of this study was to investigate the effects of horse and rider asymmetries during stance and in sitting trot on the force distribution on the horse's back using a saddle pressure mat and motion capture analysis simultaneously. Data of 80 horse-rider pairs (HRP) were collected and analyzed using linear (mixed) models to determine the influence of rider and horse variables on asymmetric force distribution. Results showed high variation between HRP. Both rider and horse variables revealed significant relationships to asymmetric saddle force distribution (P < .001). During sitting trot, the collapse of the rider in one hip increased the force on the contralateral side, and the tilt of the rider's upper body to one side led to more force on the same side of the pressure mat. Analyzing different subsets of data revealed that rider posture as well as horse movements and conformation can cause an asymmetric force distribution. Because neither horse nor rider movement can be assessed independently during riding, the interpretation of an asymmetric force distribution on the saddle pressure mat remains challenging, and all contributing factors (horse, rider, saddle) need to be considered.

Ingestive mastication in horses resembles rumination but not ingestive mastication in cattle and camels.

Horses achieve a higher degree of particle size reduction through ingestive mastication than functional ruminants. We characterized mastication using chew-monitoring halters (RumiWatch) in six domestic horses, cattle, and Bactrian camels each. All animals were offered grass hay of the same batch for 15 min. In cattle and camels, measurements were continued after eating until rumination was observed. Except for one horse, 96% of the horses' ingestive mastication data were identified as "rumination" by the proprietary RumiWatch algorithm, whereas ingestion and rumination by cattle and camels were mostly classified correctly. There were no systematic differences between cattle and camels. In cattle and camels, ingestive mastication was less regular than rumination, indicated by significantly higher standard deviations of chewing peak intervals, peak heights, and peak breadths in intraindividual comparisons. The average standard deviations of these measures were lower in horses than in cattle and camel ingestive mastication, indicating a more consistent chewing pattern in horses. Horse values were similar to those of rumination mastication, suggesting equally regular chewing motions. Regular, rhythmic chewing represents a common feature of horses and functional ruminants, but the less uniform ingestive mastication in functional ruminants represents a deviating pattern, the adaptive value of which remains unclear. In particular, it does not appear to promote a higher ingestion rate. A potential cause may be the avoidance of high tooth wear rates by delaying a more regular, systematic mastication until ingesta has been softened and the grit has been washed off in the forestomach.

Influence of ruminal methane on digesta retention and digestive physiology in non-lactating dairy cattle.

Enteric methane (CH4) production is a side-effect of herbivore digestion, but it is unknown whether CH4 itself influences digestive physiology. We investigated the effect of adding CH4 to, or reducing it in, the reticulorumen (RR) in a 4×4 Latin square experiment with rumen-fistulated, non-lactating cows, with four treatments: (i) control, (ii) insufflation of CH4 (iCH4), (iii) N via rumen fistula, (iv) reduction of CH4 via administration of bromochloromethane (BCM). DM intake (DMI), apparent total tract digestibility, digesta mean retention times (MRT), rumen motility and chewing activity, spot breath CH4 emission (CH4exhal, litre/kg DMI) as well as CH4 dissolved in rumen fluid (CH4RRf, µg/ml) were measured. Data were analysed using mixed models, including treatment (or, alternatively, CH4exhal or CH4RRf) and DMI relative to body mass0·85 (rDMI) as covariates. rDMI was the lowest on the BCM treatment. CH4exhal was highest for iCH4 and lowest for BCM treatments, whereas only BCM affected (reduced) CH4RRf. After adjusting for rDMI, CH4RRf had a negative association with MRT in the gastrointestinal tract but not in the RR, and negative associations with fibre digestibility and measures of rumination activity. Adjusting for rDMI, CH4exhal had additionally a negative association with particle MRT in the RR and a positive association with rumen motility. Thus, higher rumen levels of CH4 (CH4exhal or CH4RRf) were associated with shorter MRT and increased motility. These findings are tentatively interpreted as a feedback mechanism in the ruminant digestive tract that aims at mitigating CH4 losses by shortening MRT at higher CH4.

Digesta retention patterns of solute and different-sized particles in camelids compared with ruminants and other foregut fermenters.

The mean retention times (MRT) of solute or particles in the gastrointestinal tract and the forestomach (FS) are crucial determinants of digestive physiology in herbivores. Besides ruminants, camelids are the only herbivores that have evolved rumination as an obligatory physiological process consisting of repeated mastication of large food particles, which requires a particle sorting mechanism in the FS. Differences between camelids and ruminants have hardly been investigated so far. In this study we measured MRTs of solute and differently sized particles (2, 10, and 20 mm) and the ratio of large-to-small particle MRT, i.e. the selectivity factors (SF(10/2mm), SF(20/2mm), SF(20/10mm)), in three camelid species: alpacas (Vicugna pacos), llamas (Llama glama), and Bactrian camels (Camelus bactrianus). The camelid data were compared with literature data from ruminants and non-ruminant foregut fermenters (NRFF). Camelids and ruminants both had higher SF(10/2mm)FS than NRFF, suggesting convergence in the function of the FS sorting mechanism in contrast to NRFF, in which such a sorting mechanism is absent. The SF(20/10mm)FS did not differ between ruminants and camelids, indicating that there is a particle size threshold of about 1 cm in both suborders above which particle retention is not increased. Camelids did not differ from ruminants in MRT(2mm)FS, MRTsoluteFS, and the ratio MRT(2mm)FS/MRTsoluteFS, but they were more similar to 'cattle-' than to 'moose-type' ruminants. Camelids had higher SF(10/2mm)FS and higher SF(20/2mm)FS than ruminants, indicating a potentially slower particle sorting in camelids than in ruminants, with larger particles being retained longer in relation to small particles.

Digesta kinetics in gazelles in comparison to other ruminants: Evidence for taxon-specific rumen fluid throughput to adjust digesta washing to the natural diet.

Digesta flow plays an important role in ruminant digestive physiology. We measured the mean retention time (MRT) of a solute and a particle marker in the gastrointestinal tract (GIT) and the reticulorumen (RR) of five gazelles and one dikdik species. Species-specific differences were independent from body mass (BM) or food intake. Comparative evaluations (including up to 31 other ruminant species) indicate that MRT GIT relate positively to BM, and are less related to feeding type (the percentage of grass in the natural diet, %grass) than MRT RR. The MRTparticleRR is related to BM and (as a trend) %grass, matching a higher RR capacity with increasing BM in grazers compared to browsers. MRTsoluteRR is neither linked to BM nor to %grass but shows a consistent phylogenetic signal. Selectivity factors (SF; MRTparticle/MRTsolute, proxies for the degree of digesta washing) are positively related to %grass, with a threshold effect, where species with >20% grass have higher SF. These findings suggest that in different ruminant taxa, morphophysiological adaptations controlling MRTsoluteRR evolved to achieve a similar SF RR in relation to a %grass threshold. A high SF could facilitate an increased microbial yield from the forestomach. Reasons for variation in SF above the %grass threshold might represent important drivers of ruminant diversification and await closer investigation.

Methane emission by adult ostriches (Struthio camelus).

Ostriches (Struthio camelus) are herbivorous birds with a digestive physiology that shares several similarities with that of herbivorous mammals. Previous reports, however, claimed a very low methane emission from ostriches, which would be clearly different from mammals. If this could be confirmed, ostrich meat would represent a very attractive alternative to ruminant-and generally mammalian-meat by representing a particularly low-emission agricultural form of production. We individually measured, by chamber respirometry, the amount of oxygen consumed as well as carbon dioxide and methane emitted from six adult ostriches (body mass 108.3±8.3 kg) during a 24-hour period when fed a pelleted lucerne diet. While oxygen consumption was in the range of values previously reported for ostriches, supporting the validity of our experimental setup, methane production was, at 17.5±3.2 L d(-1), much higher than previously reported for this species, and was of the magnitude expected for similar-sized, nonruminant mammalian herbivores. These results suggest that methane emission is similar between ostriches and nonruminant mammalian herbivores and that the environmental burden of these animals is comparable. The findings furthermore indicate that it appears justified to use currently available scaling equations for methane production of nonruminant mammals in paleo-reconstructions of methane production of herbivorous dinosaurs.

Methane emission by camelids.

Methane emissions from ruminant livestock have been intensively studied in order to reduce contribution to the greenhouse effect. Ruminants were found to produce more enteric methane than other mammalian herbivores. As camelids share some features of their digestive anatomy and physiology with ruminants, it has been proposed that they produce similar amounts of methane per unit of body mass. This is of special relevance for countrywide greenhouse gas budgets of countries that harbor large populations of camelids like Australia. However, hardly any quantitative methane emission measurements have been performed in camelids. In order to fill this gap, we carried out respiration chamber measurements with three camelid species (Vicugna pacos, Lama glama, Camelus bactrianus; n = 16 in total), all kept on a diet consisting of food produced from alfalfa only. The camelids produced less methane expressed on the basis of body mass (0.32±0.11 L kg⁻¹ d⁻¹) when compared to literature data on domestic ruminants fed on roughage diets (0.58±0.16 L kg⁻¹ d⁻¹). However, there was no significant difference between the two suborders when methane emission was expressed on the basis of digestible neutral detergent fiber intake (92.7±33.9 L kg⁻¹ in camelids vs. 86.2±12.1 L kg⁻¹ in ruminants). This implies that the pathways of methanogenesis forming part of the microbial digestion of fiber in the foregut are similar between the groups, and that the lower methane emission of camelids can be explained by their generally lower relative food intake. Our results suggest that the methane emission of Australia's feral camels corresponds only to 1 to 2% of the methane amount produced by the countries' domestic ruminants and that calculations of greenhouse gas budgets of countries with large camelid populations based on equations developed for ruminants are generally overestimating the actual levels.

Human-introduced long-term traditions in wild redfronted lemurs?

Behavioural traditions have only been described for a small subset of species, and the factors responsible for the maintenance of traditions over time are unclear. Redfronted lemurs are known to learn socially but traditions have not been described in the wild. We conducted a social diffusion experiment over three experimental years with artificial feeding boxes that could be opened in two different ways (pushing or pulling a door). Six out of 14 individuals that participated in at least 2 years exhibited a stable preference: five lemurs maintained a pull and one lemur a push preference, suggesting that habit formation and reinforcement learning may have lead to preferences over time. The remaining individuals exhibited fluctuating preferences and switched between showing a preference or no preference, but never switched between preferences. This instability might have been due to the low level of difficulty and/or the low object specificity of the task. The majority of lemurs additionally scrounged. Scrounging was not influenced by age, sex or success in manipulating the boxes. Thus, redfronted lemurs appear to use the two techniques flexibly but also scrounged opportunistically to get access to the rewards, indicating that traditions might be stabilized by multiple factors.

Energy requirements and metabolism of the Phillip's dikdik (Madoqua saltiana phillipsi).

Basal metabolic rates in mammals are mainly determined by body mass, but also by ecological factors. Some mammalian species inhabiting hot, dry environments were found to have lower metabolic rates compared to temperate species. We studied energy metabolism in Phillip's dikdik (Madoqua saltiana phillipsi), a small antelope inhabiting xeric shrubland habitats in the Eastern 'horn' of Africa, and compared results to literature data. We measured body mass (BM) changes and digestibility in 12 adults kept on different food intake levels to determine, by extrapolation to zero BM change, maintenance energy requirements (MEm) for metabolizable energy (ME). The MEm averaged at 404±20kJMEkgBM(-0.75)d(-1). In addition we conducted 24h-chamber respirometry with seven fed (non-fasted) individuals. Their mean metabolic rate as calculated from oxygen consumption was 403±51kJkgBM(-0.75)d(-1), corroborating the results of the feeding experiments. Selecting the 20 lowest values of the respiration measurement period to estimate resting metabolic rate (RMR) resulted in a mean RMR of 244±39kJkgBM(-0.75)d(-1), which was not significantly lower than the expected basal metabolic rate of 293kJkgBM(-0.75)d(-1). Therefore, resting metabolism was similar to the expected average basal metabolism of a mammal of this size, which suggests a comparatively low metabolic rate in dikdiks. Compared to literature data Phillip's dikdiks have a MEm similar to measurements reported for small domestic ruminants, but considerably lower than those reported for other wild ruminant species inhabiting temperate and cold climates.

Retention of solutes and different-sized particles in the digestive tract of the ostrich (Struthio camelus massaicus), and a comparison with mammals and reptiles.

Ostriches (Struthio camelus) achieve digesta retention times, digesta particle size reduction and digestibilities equal to similar-sized herbivorous mammals, in contrast to some other avian herbivores. The sequence of digestive processes in their gastrointestinal tract, however, is still unexplored. Using two groups of four ostriches (mean body mass 75.1 ± 17.3 kg) kept on fresh alfalfa, we tested the effect of two intake levels (17 and 42 g dry matter kg(-0.75)d(-1)) on the mean retention time (MRT) of a solute and three different-sized (2, 10, 20 mm) particle markers, mean faecal particle size (MPS), and digestibility. Intake level did not affect MRT, but MPS (0.74 vs. 1.52 mm) and dry matter digestibility (81 vs. 78%). The solute marker (MRT 22-26 h) was excreted faster than the particle markers; there was no difference in the MRT of 10 and 20 mm particles (MRT 28-32 h), but 2mm particles were retained longer (MRT 39-40 h). Because the solute marker was not selectively retained, and wet-sieving of gut contents of slaughtered animals did not indicate smaller particles in the caeca, the long MRT of small particles is interpreted as intermittent excretion from the gizzard, potentially due to entrapment in small grit. The marker excretion pattern also showed intermittent peaks for all markers in five of the animals, which indicates non-continuous outflow from the gizzard. When adding our data to literature data on avian herbivores, a dichotomy is evident, with ostrich and hoatzin (Opisthocomus hoazin) displaying long MRTs, high digestibilities, and gut capacities similar to mammalian herbivores, and other avian herbivores such as grouse, geese or emus with shorter MRTs, lower fibre digestibilities and lower gut capacities. In the available data for all avian herbivores where food intake and MRTs were measured, this dichotomy and food intake level, but not body mass, was related to MRT, adding to the evidence that body mass itself may not be sole major determinant of digestive physiology. The most striking difference between mammalian and avian herbivores from the literature is the fundamentally lower methane production measured in the very few studies in birds including ostriches, which appears to be at the level of reptiles, in spite of general food intake levels of a magnitude as in mammals. Further studies in ostriches and other avian herbivores are required to understand the differences in digestive mechanisms between avian and mammalian herbivores.