The effect of size and density on the mean retention time of particles in reindeer (Rangifer tarandus).

Particle passage from the reticulorumen (RR) depends on particle density and size. A classic way of assessing these effects is the use of plastic markers of varying density and size that are recovered in the faeces. Here, we report results of an experiment where four fistulated reindeer (Rangifer tarandus, 96 ± 12 kg) were fed two different diets (browse, voluntary dry matter intake [DMI] 70 ± 10 g/kg0.75/d; or a pelleted diet, DMI 124 ± 52 g/kg0.75/d) and dosed via fistula with 8 different particle types combining densities of 1.03, 1.22 and 1.44 g/ml and sizes of 1, 10 and 20 mm. Generally, particles that passed the digestive tract intact (not ruminated) did so relatively early after marker dosing, and therefore had shorter mean retention times (MRT) than ruminated particles. On the higher intake, the overall mean retention time (MRT) of particles was shorter, but this was not an effect of shorter MRT for either intact or ruminated particles, but due to a higher proportion of intact particles at the higher intake. This supports the concept that ruminants do not adjust chewing behaviour depending on intake, but that a lower proportion of digesta is submitted to rumination due to pressure-driven escape from the forestomach at higher gut fills. Compared to cattle (Bos primigenius taurus), muskoxen (Ovibos moschatus) and moose (Alces alces) that had received the same markers, reindeer had a lower proportion of 1 mm particles that passed intact. Our results support the concept that the critical size threshold for particles leaving the ruminant forestomach is dependent on body size. While the results likely do not represent findings peculiar for reindeer, they indicate fundamental mechanisms operating in the forestomach of ruminants.

Comparative study of feeding and rumination behaviour of goats and sheep fed mixed grass hay of different chop length.

Rumination is reported to be more pronounced in sheep compared to goats. This study compared the feeding and rumination behaviour of small ruminants and consisted of two experiments (E1 and E2). In E1, four sheep and four goats were offered low-quality hay (NDFom: 692 g/kg dry matter [DM]), processed to two chop lengths (long hay [LH]: 35 mm; short hay [SH]: 7 mm) in a 2 × 2 factorial (2 species × 2 chop lengths), cross-over design. In E2, the same animals were offered moderate-quality hay (NDFom: 636 g/kg DM) processed as LH and SH. Hay was offered for ad libitum consumption. Feeding and rumination behaviour was evaluated using video recordings. Aspects of rumination like chewing frequency were evaluated for 30 min per day. Faecal samples were analysed for faecal-N and particle size. There was no species effect on feed intake and organic matter digestibility (faecal N as proxy); however, goats consumed more LH than SH in E1 and E2. There was an effect of species on rumination:eating duration (R:E) ratio (higher in sheep) in E1 but not in E2, where there was a tendency for a species effect on rumination duration. In E1 and E2, sheep had a higher R:E ratio for SH than for LH. For rumination behaviour, there was a species effect for number of daily boli, chewing frequency and chews per day (more in sheep) in E1 and E2. No effect of species was found for faecal particle size. Despite much concordance, feed comminution behaviour differed in some aspects between sheep and goats. In an evolutionary context, a shift of significance of rumination could be triggered by a higher amount of abrasives in natural diets of sheep, rendering a shift of chewing towards ruminally prewashed material a rewarding strategy.

Teeth and the gastrointestinal tract in mammals: when 1 + 1 = 3.

Both teeth and the digestive tract show adaptations that are commonly interpreted in the context of trophic guilds-faunivory, herbivory and omnivory. Teeth prepare food for the digestive tract, and dental evolution focuses on increasing durability and functionality; in particular, size reduction of plant particles is an important preparation for microbial fermentative digestion. In narratives of digestive adaptations, microbes are typically considered as service providers, facilitating digestion. That the majority of 'herbivorous' (and possibly 'omnivorous') mammals display adaptations to maximize microbes' use as prey-by harvesting the microbes multiplying in their guts-is less emphasized and not reflected in trophic labels. Harvesting of microbes occurs either via coprophagy after separation from indigestible material by a separation mechanism in the hindgut, or from a forestomach by a 'washing mechanism' that selectively removes fines, including microbes, to the lower digestive tract. The evolution of this washing mechanism as part of the microbe farming niche opened the opportunity for the evolution of another mechanism that links teeth and guts in an innovative way-the sorting and cleaning of not-yet-sufficiently-size-reduced food that is then re-submitted to repeated mastication (rumination), leading to unprecedented chewing and digestive efficiency. This article is part of the theme issue 'Food processing and nutritional assimilation in animals'.

Does age matter?-Efficiency of mechanical food break down in Tupaia belangeri at different ages.

The relationship of food comminution and individual age in Tupaia belangeri is investigated. It is hypothesized that with increasing age the performance of the molar dentition decreases due to progressive tooth wear. While this relationship is well-documented for herbivores, age-related test series are largely lacking for insectivorous mammals. 15 individuals of Tupaia belangeri were fed exclusively with mealworms, and their faeces were analyzed for the number and size of chitin particles. The exoskeleton of a mealworm is resistant to digestive fluids in the gastrointestinal tract, and the size of individual chitin particles indicates the effectiveness of mechanical comminution that occurs in the oral cavity during mastication. It is hypothesized that a more precise occlusion of the dentition results in smaller particle size. Although individuals of all ages (juvenile, adult, and senile) were able to effectively process mealworms with their dentition prior to digestion, a larger area of very large chitin particles (98% quantile of all particles in senile animals as compared to in the same quantile in adults) in the feces of senile animals was detected. Even though the particle size of indigestible material is irrelevant for the digestive process, these findings either document somatic senescence in the functionality of the teeth, or alternatively a change in chewing behaviour with age.

Equid Nutritional Physiology and Behavior: An Evolutionary Perspective.

Like other members of the odd-toed ungulates (the perissodactyls), equids once had a higher species diversity in the fossil record than they have today. This is generally explained in comparison to the enormous diversity of bovid ruminants. Theories on putative competitive disadvantages of equids include the use of a single toe as opposed to two toes per leg, the lack of a specific brain cooling (and hence water-saving) mechanism, longer gestation periods that delay reproductive output, and in particular digestive physiology. To date, there is no empirical support for the theory that equids fare better on low-quality forage than ruminants. In contrast to the traditional juxtaposition of hindgut and foregut fermenters, we suggest that it is more insightful to sketch the evolution of equid and ruminant digestive physiology as a case of convergence: both evolved a particularly high chewing efficacy in their respective groups, which facilitates comparatively high feed and hence energy intakes. But because the ruminant system, less based on tooth anatomy but more on a forestomach sorting mechanism, is more effective, equids depend more on high feed intakes than ruminants and may well be more susceptible to feed shortages. Arguably, the most underemphasized characteristic of equids may be that in contrast to many other herbivores including ruminants and coprophageous hindgut fermenters, equids do not use the microbial biomass growing in their gastrointestinal tract. Equids display behavioral and morphophysiological adaptations to high feed intakes, and their cranial anatomy that facilitates the cropping of forage while performing grinding chewing at the same time might be unique. Rather than looking for explanations how equids are better adapted to their present niches than other organisms, considering them remnants of a different morphophysiological solution may be more appropriate.

A pilot investigation on the effect of induced saliva flow on digestive parameters in sheep, and a comparison with cattle.

Sheep with a relatively low methane yield were observed to have shorter fluid and particle mean retention times (MRT). Because the application of pilocarpine, a saliva stimulant, was successful in reducing retention times in ruminants in previous studies, we applied this substance to sheep, expecting a reduction in MRT and methane yield. Three non-pregnant sheep (74 ± 10 kg) were fed a hay-only diet in a 3 × 3 Latin square design with oral doses of 0, 2.5 and 5 mg pilocarpine/kg body weight and day. Measurements included feed and water intake, MRT of liquid and particulate phases in the reticulorumen (RR) and total gastrointestinal tract (GIT), ruminal microbial yield (via urinary purine bases and metabolic faecal nitrogen), total tract methane emission, apparent nutrient digestibility and rumen fluid parameters. Data were investigated for linear and quadratic effects using orthogonal polynomial contrasts. The MRT of liquid and small particles in the RR and total GIT, and the short-chain fatty acid concentration in rumen fluid, linearly declined with increasing pilocarpine dosage, while no quadratic relationship was detected. Intake of feed DM and water, apparent nutrient digestibility, methane yield and microbial yield were not affected by pilocarpine. When combining the sheep data with that of a similar experiment in cattle, we found that the MRT of the liquid phase was positively associated with estimated NDF digestibility and with methane production per digested NDF, but was not associated with microbial yield or the ratio of acetate to propionate. The ratio between MRT of the particulate and the liquid phase was smaller for sheep than that for cattle, and was not affected by treatment. Differences in this ratio might explain why species reacted differently to the saliva-inducing agent, which might help to explain the discrepancy between species in the effect of induced saliva flow on digestive parameters.

Is there a difference in ruminal fermentation control between cattle and sheep? A meta-analytical test of a hypothesis on differential particle and fluid retention.

Ruminant species differ in digestive physiology. The species-specific ratio of mean retention time of particles and fluid (MRTparticle/MRTfluid) in the reticulorumen has been interpreted as controlling ruminal fermentation: a higher ratio indicates of a more distinct 'washing' of particulate digesta by liquid. This should increase the harvest of microbes from the reticulorumen, and keep the microbiome in a state of more intense growth; at the same time, this should increase the metabolic losses of faecal nitrogen of microbial origin, leading to lower values for the apparent digestibility of crude protein (aD CP). A systematic difference has been hypothesized between cattle (higher ratio) and sheep (lower ratio), with a lower MRTfluid in cattle due to a higher saliva production. Here, we test these hypotheses in a meta-analysis, using only studies that investigated cattle and sheep simultaneously. The datasets included 12 studies on MRT (of which 11 contained information on feed intake), yielding 102 (or 89) individual data; and 26 studies on protein digestibility (of which 18 contained information on intake), yielding 349 individual data. Cattle had a higher MRTparticle/MRTfluid (2.1) than sheep (1.7), mainly due to longer MRTparticle; only if body mass was included in the model, MRTfluid was significantly shorter in cattle in the larger MRT dataset (and tended to be shorter in the slightly smaller dataset). Cattle had a significantly lower aD CP than sheep, while there was no such difference in overall (dry or organic matter) digestibility. The dataset confirms a shift in fermentation strategy towards microbial production in cattle. While this has been suggested for ruminants in general, cattle appear particularly far on an evolutionary trajectory of maximizing microbial yield from the forestomach. The application of more specific digestive physiology data (like endogenous losses) gained from sheep to cattle should be done bearing these differences in mind.

Relationship between reticular pH parameters and potential on-farm indicators in the early lactation of dairy cows.

Subacute ruminal acidosis (SARA) is an important nutritional disorder affecting animal welfare and economy of milk production. Definitions rely on ruminal pH but due to limitations of its measurement, indicators reflecting low pH are highly desirable. The aim of this study was to investigate the relationship between reticular pH and 18 on-farm indicators in milk, blood, faeces, urine and chewing behaviour in early lactating dairy cows. Ten farms were visited for 3 weeks and in total samples of 100 cows (10 per farm) were taken. The statistics and graphical visualization were performed using Pearson correlation and linear regression models on an animal individual level as well as with linear mixed models. Eight indicators (milk fat, fat-to-protein ratio, rumination time, feed intake time, rumination frequency, rumination boluses, lying time and faecal pH) were statistically significant associated with the daily animal individual reticular pH average. However, none of the models including the potential explanatory variables explained more than 5% of the pH variations. The study confirms the necessity of pH measurement to detect SARA risk animals in early lactation dairy cows.

Individual differences in digesta retention and their relation to chewing in cattle-A pilot investigation.

While information on individual differences in digesta mean retention time (MRT) might be interesting when selecting phenotypes for digestive efficiency, MRT measurements are prohibitively labour-intensive for large-scale application. Therefore, more easily measured proxies of MRT might be helpful. We used the opportunity of an experiment applying saliva stimulant in cattle to investigate the effect of different individual chewing behaviour on fluid and particle MRT with a consistent diet. Four non-lactating cattle (670-850 kg body mass [BM]) were used in a 4 × 4 Latin square design, treated with the saliva stimulant pilocarpine in dosages of 0, 1, 2.5 and 5 mg/kg BM per day. The cattle were fed hay with dry matter intake (DMI) assigned according to their metabolic body weight. MRT in the whole gastrointestinal tract (GIT), the reticulorumen (RR) and the distal tract were measured using Co-EDTA, Cr-mordanted fibre and La-mordanted fibre as markers representing fluid, small particles (2 mm) and large particles (1 cm), respectively. The chewing behaviour was measured via noseband pressure sensor and expressed as chewing frequency (chews per time) and chewing intensity (chews per DMI), both for total chewing (ingestion plus rumination) and rumination chewing alone. The animals differed considerably in chewing behaviour and MRT measures. BM did not show a significant effect on chewing behaviour and MRT measures, though it tended to negatively correlated to total chewing intensity. Chewing intensity exerted a significant negative influence on MRT of fluid and particles in the RR, which was not the case for chewing frequency. Chewing frequency showed a significant relationship with MRT of large particles in the GIT. We suggest that chewing behaviour could influence MRT in two ways: (i) by affecting saliva production via the masticatory-salivary reflex and subsequently, the fluid inflow to the RR; (ii) by contributing to particle size reduction. Should the link between chewing behaviour and MRT be corroborated in larger studies, chewing measures, with their large interindividual variation, could emerge as an easy-to-measure proxy for MRT characteristics.

Effect of induced saliva flow on fluid retention time, ruminal microbial yield and methane emission in cattle.

Both in vitro and animal studies indicated that a higher dilution rate is related to a more efficient microbial synthesis and a lower methane (CH4 ) yield. The latter could be a consequence of the former, as an increase in microbial cell synthesis offers an alternative hydrogen sink competing with methanogenesis. To test this assumption in live animals, we applied a saliva stimulant, pilocarpine, to modify liquid flow rate in cattle. Four non-lactating cows (750 ± 71 kg) were fed forage only (restricted to constant intake) in a 4 × 4 Latin square design with oral doses of 0, 1, 2.5 and 5mg pilocarpine/kg body weight and day. We quantified feed and water intake, ruminal and total tract mean retention time (MRT) of solute and particle markers, ruminal microbial yield (via urinary purine bases or metabolic faecal nitrogen), CH4 emission, digestibility, chewing behaviour, reticular motility and rumen fluid parameters. The effect of induced saliva flow was evident by visibly increased salivation and water intake. Increasing the pilocarpine dosages resulted in a linearly decreased MRT of fluid and small particles (p < 0.001 and p< 0.05, respectively) and methane yield as related to digested DM (p < 0.05), the latter at a magnitude of 5%. No effect of treatment was found on ruminal microbial yield estimated via purine derivates. Metabolic faecal N as an indicator of microbial growth linearly correlated with pilocarpine dosages (p < 0.05). No significant relationship was found between pilocarpine dosages and large particle MRT, nutrient digestibility, ruminal pH and short-chain fatty acids. In conclusion, different from some in vitro studies, there was little indication of a reciprocal effect of CH4 and microbial biomass production in cows fed a forage-only diet.

The Ruminant sorting mechanism protects teeth from abrasives.

Dental wear due to ingestion of dust and grit has deleterious consequences. Herbivores that could not wash their food hence had to evolve particularly durable teeth, in parallel to the evolution of dental chewing surface complexity to increase chewing efficacy. The rumen sorting mechanism increases chewing efficacy beyond that reached by any other mammal and has been hypothesized to also offer an internal washing mechanism, which would be an outstanding example of an additional advantage by a physiological adaptation, but in vivo evidence is lacking so far. Here, we investigated four cannulated, live cows that received a diet to which sand was added. Silica in swallowed food and feces reflected experimental dietary sand contamination, whereas the regurgitate submitted to rumination remained close to the silica levels of the basal food. This helps explain how ruminants are able to tolerate high levels of dust or grit in their diet, with less high-crowned teeth than nonruminants in the same habitat. Palaeo-reconstructions based on dental morphology and dental wear traces need to take the ruminants' wear-protection mechanism into account. The inadvertent advantage likely contributed to the ruminants' current success in terms of species diversity.

Fermentation of liquid feed with lactic acid bacteria reduces dry matter losses, lysine breakdown, formation of biogenic amines, and phytate-phosphorus.

This study investigated the fermentation of liquid feed for pigs and the effect of lactic acid bacteria (LAB) supplementation on fermentation rate, dry matter losses (DML), formation of biogenic amines, and degradation of phytate-P. The basal substrate in all three in vitro batch experiments consisted of 50% canola meal, 25% wheat, and 25% barley. The mixed substrates were adjusted to a dry matter (DM) content of 28.4% and fermented in 1-liter vessels at 37 °C for 24 h. Experiment 1 focused on changes in pH profiles over time. Treatments were as follows: 1) liquid feed without additive (control) and 2) liquid feed supplemented with a mixture of Lactobacillus plantarum, Pediococcus pentosaceus, and Lactobacillus lactis (adLAB) at 2.0 × 105 CFU/g liquid feed (wet wt.; n = 8). Substrate pH was measured every 2 h. Experiment 2 focused on DML and the impact of fermentation on phytate-P. Treatments were identical to experiment 1 (control and adLAB; n = 8). Measured parameters included concentration of lactic acid, acetic acid, ethanol, and phytate-P, and DML after 24 h of fermentation. Counts of molds, Enterobobacteriaceae, yeasts, and LAB were determined in one combined sample of all replicates. Dry matter losses were lower in LAB-supplemented fermentations (5.89%) compared to the control (11.8%; P < 0.001). Supplementation with LAB reduced the phytate-P content (2.66 g/kg DM) compared to the control (3.07 g/kg DM; P = 0.002). Experiment 3 evaluated DML and the impact of fermentation on formation of biogenic amines. Treatments were as follows: 1) control, 2) adLAB (2.0 × 105 CFU LAB/g liquid feed), 3) adLys (0.60% DM supplemented lysine), and 4) adLAB+Lys (combination of adLAB and adLys; n = 8). The fermentation of adLys resulted in a nearly complete breakdown of supplemented lysine, whereas only 10% of supplemented lysine was lost in adLAB+Lys. Furthermore, all adLys samples tested positive for cadaverine (mean concentration 0.89% DM), whereas no adLAB samples contained cadaverine above the detection limit (P < 0.001). Results indicate that DML is reduced in fermentations supplemented with homofermentative LAB. Fermentation of liquid feed with homofermentative LAB can effectively reduce the degradation of supplemental lysine and has the potential to further improve P availability.

Sand accumulation in the digestive tract of rabbits (Oryctolagus cuniculus) and guinea pigs (Cavia porcellus): The role of the appendix.

We determined location and amount of accumulated sand in the gastrointestinal tract (GIT) of rabbits (Oryctolagus cuniculus) and guinea pigs (Cavia porcellus) fed diets containing external (silicate) abrasives. Computed tomographic abdominal images of rabbits (n = 44) and guinea pigs (n = 16) that each received varying numbers (4-7) of different diets for 14 days each (total n = 311 computed tomographs), and radiographs of dissected GIT and presence of silica in GIT content (n = 46 animals) were evaluated. In rabbits, the majority of accumulated sand was located in the caecal appendix, an elongated, intestinal structure in the left side of the abdomen. The 'wash-back' colonic separation mechanism in rabbits may be partly responsible for a retrograde transport of sand back to the caecum, where dense, small particles accumulate in the appendix. The appendix likely acted as a reservoir of these particles, leading to significant effects not only of the momentary but also of the previous diet on recorded sand volumes in the rabbits. Guinea pigs have no caecal appendix and a colonic separation mechanism not based on a 'wash-back'. Less sand accumulation was found in their GIT without a specific location pattern, and there were less previous diet effects in this species. None of the rabbits or guinea pigs developed clinical signs of obstruction during the study, and the recorded sand volumes represented 1.0 ± 1.2% of the 14-d sand intake in rabbits and 0.2 ± 0.2% in guinea pigs. Accumulation of sand in volumes up to 10 cm3 in the GIT of rabbits does not seem to cause clinical health impairment. Large inter-individual differences in rabbits indicate inter-individual variation in proneness to sand accumulation. The reason for the presence of a sand-trapping caecal appendix in animals that are, due to their burrowing lifestyle and feeding close to the ground, predestined for accidental sand ingestion, remains to be unveiled.

Digestion of bamboo compared to grass and lucerne in a small hindgut fermenting herbivore, the guinea pig (Cavia porcellus).

Bamboo is an enigmatic forage, representing a niche food for pandas and bamboo lemurs. Bamboo might not represent a suitable forage for herbivores relying on fermentative digestion, potentially due to its low fermentability. To test this hypothesis, guinea pigs (n = 36) were used as model species and fed ad libitum with one of three forages (bamboo, lucerne, or timothy grass) in a fresh or dried state, with six individuals per group, for 3 weeks. The nutrient composition and in vitro fermentation profile of bamboo displayed low fermentation potential, i.e. high lignin and silica levels together with a gas production (Hohenheim gas test) at 12 h of only 36% of that of lucerne and grass. Although silica levels were more abundant in the leftovers of (almost) all groups, guinea pigs did not select against lignin on bamboo. Dry matter (DM) intake was highest and DM digestibility lowest on the bamboo forage. Total short-chain fatty acid levels in caecal content were highest for lucerne and lowest for grass and bamboo. Bamboo-fed guinea pigs had a lower body weight gain than the grass and lucerne group. The forage hydration state did not substantially affect digestion, but dry forage led to a numerically higher total wet gut fill. Although guinea pigs increased DM intake on the bamboo diet, the negative effects on fermentation of lignin and silica in bamboo seemed overriding. For herbivores that did not evolutionary adapt, bamboo as an exclusive food resource can be considered as inadequate.

Tooth wear, growth and height in rabbits (Oryctolagus cuniculus) fed pelleted or extruded diets with or without added abrasives.

Among the different factors thought to affect dental wear, dietary consistency is possibly the least investigated. To understand tooth wear of herbivorous animals consuming different dietary consistencies with different abrasive potential, we fed 14 rabbits (Oryctolagus cuniculus) exclusively with a timothy grassmeal-based diet in either pelleted or extruded form, or the same diets with an addition of 5% fine sand abrasives (mean size 130 µm). First, we offered the rabbits the pelleted and extruded diets as well as the pelleted control and pelleted abrasive diet in a two-stage preference experiment. Then, the rabbits received each diet for 2 weeks in a randomised serial feeding experiment, where each animal served as its own control. Tooth measurements for wear, growth and height were achieved using a manual calliper, endoscopic examination and CT scans. The analysis of the diets as fed showed almost identical mean particle size, but the extruded diet had a lower density (volume/mass) and softer consistency compared to the pelleted one and was favoured by most rabbits. The rabbits selected against the diet with sand during the preference experiment, possibly because it caused more tooth wear, especially on the teeth most exposed to wear along the upper tooth row (upper P4 and M1). The maxillary teeth also showed evidence of an increased chewing laterality by the end of the experiment. The extruded diet led to a significantly lower cheek teeth height than the pelleted diet, potentially due to the higher chewing effort needed for a similar dry matter intake. The results suggest that dietary hardness alone is a poor predictor of dental wear. The regrowth of the teeth matched wear consistently.

Macrowear effects of external quartz abrasives of different size and concentration in rabbits (Oryctolagus cuniculus).

External quartz abrasives are one of the driving forces of macrowear in herbivorous animals. We tested to what extent different sizes and concentrations influence their effect on tooth wear. We fed seven pelleted diets varying only in quartz concentration (0%, 4%, and 8%) and size (fine silt: ∼4 µm, coarse silt: ∼50 µm, fine sand: ∼130 µm) to rabbits (Oryctolagus cuniculus, n = 16) for 2 weeks each in a randomized serial experiment. Measurements to quantify wear and growth of incisors and the mandibular first cheek tooth, as well as heights of all other cheek teeth, were performed using calipers, endoscopic examination, and computed tomography scans before and after each feeding period. Tooth growth showed a compensatory correlation with wear. Absolute tooth height (ATH) and relative tooth height (RTH); relative to the 0% quartz "control" diet) was generally lower on the higher concentration and the larger size of abrasives. The effect was more pronounced on the maxillary teeth, on specific tooth positions and the right jaw side. When offered the choice between different sizes of abrasives, the rabbits favored the silt diets over the control and the fine sand diet; in a second choice experiment with different diets, they selected a pelleted diet with coarse-grained sand, however. This study confirms the dose- and size-dependent wear effects of external abrasives, and that hypselodont teeth show compensatory growth. The avoidance of wear did not seem a priority for animals with hypselodont teeth, since the rabbits did not avoid diets inducing a certain degree of wear.

Reticular contraction frequency and ruminal gas dome development in goats do not differ between grass and browse diets.

In investigations of differences between ruminant species feeding on browse or grass, it is often unclear whether observed differences are animal- or forage-specific. Ruminant species have been classified as 'moose-type', with little rumen content stratification, or 'cattle-type' with a distinct rumen contents stratification, including a gas layer. To which extent putative differences in forestomach motility are involved in these patterns is unknown. Using sonography, we investigated the frequency of reticular contractions and the stratification of rumen contents in goats fed exclusively on grass hay (n = 6) or dried browse (n = 5) directly after feeding, and after another 6 and 12 h with no access to feed. The frequency of reticular contractions decreased from immediately after feeding (1.8 ± 0.3 min-1 ) to 6 h afterwards (1.2 ± 0.2 min-1 ) and then remained constant, with no difference between diets. A gas dome became more visible over time, but neither its incidence nor its extent differed between diets. The results are in accord with classifying goats as 'cattle-type' in terms of their digestive physiology, and they add to a growing body of evidence that differences in digestive physiology between ruminant species are more due to species characteristics than different kinds of ingested forages.

Preliminary evidence for a forestomach washing mechanism in llamas (Lama glama).

Dust and grit are ingested by herbivores in their natural habitats along with the plants that represent their selected diet. Among the functions of the rumen, a washing of ingesta from adhering dust and grit has recently been demonstrated. The putative consequence is a less strenuous wear on ruminant teeth by external abrasives during rumination. The same function should theoretically apply to camelids, but has not been investigated so far. We fed six llamas (Lama glama) a diet of grass hay and a lucerne-based pelleted food in which fine sand had been included at about 8% of ingredients, for ad libitum consumption for 6 weeks. Subsequently, animals were slaughtered and content of the different sections of the gastrointestinal tract was sampled for the analysis of dry matter (DM), total ash, and acid detergent insoluble ash (ADIA, a measure for silica). Additionally, two of the animals were subjected to whole-body computer tomography (CT) after death in the natural sternal resting position. No clinical problems or macroscopic changes in the faeces were observed during the experimental period. The results indicate an accumulation of ADIA in the C3 compartment of the stomach complex, in particular in the posterior portion that is the equivalent of the abomasum in ruminants. By contrast, contents of the C1, from which material is recruited for regurgitation and rumination, were depleted of ADIA, indicating that the contents had largely been washed free of sand. The washing effect is an unavoidable side effect of the flotation- and sedimentation-based sorting mechanisms in the ruminant and the camelid forestomachs. In theory, this should allow ruminants and camelids to live in similar habitats as nonruminant herbivores at lower degrees of hypsodonty.

Increasing food intake affects digesta retention, digestibility and gut fill but not chewing efficiency in domestic rabbits (Oryctolagus cuniculus).

In ruminants, the level of food intake affects net chewing efficiency and hence faecal particle size. For nonruminants, corresponding data are lacking. Here, we report the effect of an increased food intake of a mixed diet in four domestic rabbit does due to lactation, and assess changes in particle size (as determined by wet sieving analysis) along the rabbit digestive tract. During lactation, rabbits achieved a distinctively higher dry matter intake than at maintenance, with a concomitant reduction in mean retention times of solute and particle markers, an increase in dry matter gut fill, a reduction in apparent digestibility of dry matter, and an overall increase in digestible dry matter intake. By contrast, there was no change in faecal mean particle size (mean ± SD: 0.58 ± 0.02 vs. 0.56 ± 0.01 mm). A comparison of diet, stomach content and faecal mean particle size suggested that 98% of particle size reduction occurred due to ingestive mastication and 2% due to digestive processes. Very fine particles passing the finest sieve, putatively not only of dietary but mainly of microbial origin, were particularly concentrated in caecum contents, which corresponds to retention of microbes via a 'wash-back' colonic separation mechanism, to concentrate them in caecotrophs that are re-ingested. This study gives rise to the hypothesis that chewing efficiency on a consistent diet is not impaired by intake level in nonruminant mammals.

Increasing feed intake in domestic goats (Capra hircus): Measured effects on chewing intensity are probably driven by escape of few, large particles from the forestomach.

On the one hand, oral processing - mastication - is considered a relatively inflexible component of mammalian feed acquisition that constrains instantaneous intake rates. On the other hand, experimental data shows that the level of feed intake affects faecal particle size and hence net chewing efficiency in ruminants, with larger particles occurring in the faeces at higher intakes. Here, we report the effect of an increased feed intake during maintenance (L1), late (200% of L1) and peak lactation (300% of L1) of a consistent diet (hay:concentrates 50:50) in eight domestic goats on various measures of digestive physiology including faecal mean particle size (MPS). Increasing intake led to an increased gut fill, a reduction in digesta retention times, and an increase in faecal MPS (from 0.57 to 0.72 mm). However, this was an effect of the large particle fraction (>2 mm) being disproportionately excreted at higher intakes; if MPS was assessed on the basis of particles below the typical escape threshold (≤1 mm), there was no difference between intake levels. These findings suggest that the effect of intake on the calculated net chewing efficiency in ruminants may rather be an effect of increased large particle escape from the forestomach than a reduced chewing intensity per bolus during ingestion or rumination.