Voluntary intake and digestibility in horses: effect of forage quality with emphasis on individual variability

Food intake is a key biological process in animals, as it determines the energy and nutrients available for the physiological and behavioural processes. In herbivores, the abundance, structure and quality of plant resources are known to influence intake strongly. In ruminants, as the forage quality declines, digestibility and total intake decline. Equids are believed to be adapted to consume high-fibre low-quality forages. As hindgut fermenters, it has been suggested that their response to a reduction in food quality is to increase intake to maintain rates of energy and nutrient absorption. All reviews of horse nutrition show that digestibility declines with forage quality; for intake, however, most studies have found no significant relationship with forage quality, and it has even been suggested that horses may eat less with declining forage quality similarly to ruminants. A weakness of these reviews is to combine data from different studies in meta-analyses without allowing the differences between animals and diets to be controlled for. In this study, we analysed a set of 45 trials where intake and digestibility were measured in 21 saddle horses. The dataset was analysed both at the group (to allow comparisons with the literature) and at the individual levels (to control for individual variability). As expected, dry matter digestibility declined with forage quality in both analyses. Intake declined slightly with increasing fibre contents at the group level, and there were no effects of crude protein or dry matter digestibility on intake. Overall, the analysis for individual horses showed a different pattern: intake increased as digestibility and crude protein declined, and increased with increasing fibre. Our analysis at the group level confirms previous reviews and shows that forage quality explains little of the variance in food intake in horses. For the first time, using mixed models, we show that the variable 'individual' clarifies the picture, as the horses showed different responses to a decrease in forage quality: some compensated for the low nutritional value of the forages by increasing intake, few others responded by decreasing intake with declining forage quality, but not enough to cause any deficit in their energy and protein supplies. On the whole, all the animals managed to meet their maintenance requirements. The individual variability may be a by-product of artificial selection for performance in competition in saddle horses.     

哺乳动物的消化策略（英文）

理解动物的营养生态位是充分理解其整个生态学的基础,对于害兽控制和物种保护也很重要,食肉动物的小肠很发达,这可能与对食物的高消化能力有关;杂食性动物有更复杂的胃肠器官,其后端有可进行发酵的盲肠,消化物的平均滞留时间（mean retention times,MRTs)更长;最长的平均滞留时间见于食草动物,其消化道内高密度的微生物种群对不同滞留区内的消化物进行发酵,但是,并不是所有的食草动物都能够最大程度地消化植物纤维,只有反刍动物、骆驼和个体较大的后肠发酵动物（hindgut fermenter)能够具有这种能力,对比而言,许多其它的食草动物,如前肠发酵的有袋类和小型的后肠发酵动物如兔子、田鼠和负鼠等,它们具备可以使植物纤维消化效率最大的消化系统,可以在食物中的纤维素含量非常高的情况下仍能处理大量的食物。这些不同的消化策略使哺乳动物具有广幅的营养生态位。     

Responses of pocket gophers (Thomomys bottae) to changes in diet quality

Summary We examined digestibility of dry matter, nutrients, and fiber, and food intake, metabolic fecal losses, weight change, and gut size of pocket gophers (Thomomys bottae) in relation to diet quality in the laboratory. Pocket gophers were maintained for 15–20 days on one of seven diets which contained from 18% to 56% neutral detergent fiber (NDF). NDF content of the diet was an excellent predictor of diet quality. Digestibility of dry matter, NDF, and nitrogen all decreased with increasing NDF content of the diet. In general, pocket gophers compensated for low diet quality by increasing dry matter intake, but those given high quality forage before the lowest quality diet reduced their intake. Thus, the response of pocket gophers to low quality diets may depend on their body condition. Because increased food intake resulted in increased total metabolic fecal losses and metabolic fecal nitrogen losses, decreasing food intake on low-quality diets may be advantageous. A further response of pocket gophers to decreased food quality was an increase in size of cecum and large intestine, suggesting that fermentation of cell walls became increasingly important as diet quality decreased.     

Food quality and quantity are more important in explaining foraging of an intermediate‐sized mammalian herbivore than predation risk or competition

During times of high activity by predators and competitors, herbivores may be forced to forage in patches of low‐quality food. However, the relative importance in determining where and what herbivores forage still remains unclear, especially for small‐ and intermediate‐sized herbivores. Our objective was to test the relative importance of predator and competitor activity, and forage quality and quantity on the proportion of time spent in a vegetation type and the proportion of time spent foraging by the intermediate‐sized herbivore European hare (Lepus europaeus). We studied red fox (Vulpes vulpes) as a predator species and European rabbit (Oryctolagus cuniculus) as a competitor. We investigated the time spent at a location and foraging time of hare using GPS with accelerometers. Forage quality and quantity were analyzed based on hand‐plucked samples of a selection of the locally most important plant species in the diet of hare. Predator activity and competitor activity were investigated using a network of camera traps. Hares spent a higher proportion of time in vegetation types that contained a higher percentage of fibers (i.e., NDF). Besides, hares spent a higher proportion of time in vegetation types that contained relatively low food quantity and quality of forage (i.e., high percentage of fibers) during days that foxes (Vulpes vulpes) were more active. Also during days that rabbits (Oryctolagus cuniculus) were more active, hares spent a higher proportion of time foraging in vegetation types that contained a relatively low quality of forage. Although predation risk affected space use and foraging behavior, and competition affected foraging behavior, our study shows that food quality and quantity more strongly affected space use and foraging behavior than predation risk or competition. It seems that we need to reconsider the relative importance of the landscape of food in a world of fear and competition.     

Modelling the nutritional ecology of ungulate herbivores: evolution of body size and competitive interactions

Summary A simulation model is used to quantify relationships between diet quality, digestive processes and body weight in ungulate herbivores. Retention time of food in the digestive tract is shown by regression to scale with W^0.27, and to be longer in ruminants than in hindgut fermenters. Allometric relationships between whole gut mean retention time (MRT, h) and weight (W) were: MRT=9.4 W^0.255 (r ²=0.80) for hindgut fermenters and MRT=15.3 W^0.251 (r ²=0.76) in ruminants. Longer retention of ingesta by large-bodied ruminants and hindgut fermenters increases digestive efficiency relative to small animals and permits them to survive on lower-quality foods. Compared with ruminants, hindgut fermenters' faster throughput is an advantage which outweighs their lower digestive efficiency, particularly on poor quality foods, provided that food resources are not limiting. This suggests that the predominance of ruminants in the middle range of body weights results from their more efficient use of scarce resources under conditions of resource depletion. Considering only physical limitations on intake, the model shows that the allometric coefficient which scales energy intake to body mass is 0.88 in ruminants and 0.82 in hindgut fermenters. The advantages of large body size are countered by disadvantages where food quantity is limited, and we suggest that the upper limit to ungulate body size is determined by the ability to extract nutrients from feeding niches during the nadir of the seasonal cycle of resource quality and abundance.     

Differences between cattle and sheep in their digestion and relative intake of a mature tropical grass hay

The dry-matter digestibility (DMD) of a low quality tropical grass hay was much higher in cattle (49.6%) than in sheep (34.6%). This difference was not due to a difference in relative intake per kg liveweight (W^0.9) nor to a difference in the diet selected. However, neutral-detergent fibre (NDF) was digested better by cattle than by sheep. For each kg of dry matter consumed, cattle digested 415 g NDF and sheep 279 g NDF. This was because 60% more of the hemicellulose and 35% more of the cellulose were digested by cattle than by sheep.Differences between cattle and sheep in the concentrations of urea and sulphate in blood, and the relatively lower excretion of N, P and Ca by the cattle suggested that the rumen micro-organisms of the cattle were utilizing these nutrients better than those of the sheep. This could have increased microbial activity in the rumen and, hence, digestion of fibre.     

More efficient mastication allows increasing intake without compromising digestibility or necessitating a larger gut: Comparative feeding trials in banteng (Bos javanicus) and pygmy hippopotamus (Hexaprotodon liberiensis)

The digestion of plant material in mammalian herbivores basically depends on the chemical and structural composition of the diet, the mean particle size to which the forage is processed, and the ingesta retention time. These different factors can be influenced by the animal, and they can presumably compensate for each other. The pygmy hippopotamus, a non-ruminating foregut fermenter, has longer mean retention times than ruminants; however hippos do not achieve higher (fibre) digestibilities on comparable diets, which could be due to ineffective mastication. We performed feeding trials with six pygmy hippos (Hexaprotodon liberiensis) and six banteng cattle (Bos javanicus) on a grass diet. As predicted, both species achieved similar dry matter, organic matter, crude protein and gross energy digestibilities. However, neutral and acid detergent fibre digestibility was lower in pygmy hippos. Apparently, in these species, fibre digestibility was more influenced by particle size, which was larger in pygmy hippos compared to banteng, than by retention time. In spite of their higher relative food intake, the banteng in this study did not have greater relative gut fills than the hippos. Ruminants traditionally appear intake-limited when compared to equids, because feed particles above a certain size cannot leave the rumen. But when compared to nonruminating foregut fermenters, rumination seems to free foregut fermenters from an intrinsic food intake limitation. The higher energy intakes and metabolic rates in wild cattle compared to hippos could have life-history consequences, such as a higher relative reproductive rate.     

Pyric Herbivory and the Nexus Between Forage,Fire and Native and Introduced Large Grazing Herbivores in Australian Tropical Savannas

Earth’s tropical savannas typically support high biomass of diverse grazing herbivores that depend on a highly fluctuating resource: high-quality forage. An annual wet–dry cycle, fire and herbivory combine to influence forage quality and availability throughout the year. In the savannas of northern Australia, a depauperate suite of large native (marsupial) herbivores (wallaroos [Osphranter spp.] and the agile wallaby [Notamacropus agilis]) compete for resources with non-native large herbivores introduced in the late nineteenth century, particularly bovines (feral and managed cattle [Bos spp.] and feral water buffalo [Bubalus bubalis]) that now dominate the landscape. Anecdotal reports of recent population declines of large macropods and negative impacts of bovines highlight the need to better understand the complex relationship between forage, fire and abundance of native and introduced large herbivores. The pyric herbivory conceptual model, which posits complex feedbacks between fire and herbivory and was developed outside Australia, predicts that native and introduced large herbivores will both respond positively to post-fire forage production in Australian savannas where they co-occur. We used grazing exclosures, forage biomass and nutrient analyses and motion-sensor camera-trapping to evaluate the overall robustness of the pyric herbivory model in the Australian context, specifically whether forage quantity and quality are impacted by herbivory, season and fire activity, and which forage attributes most influence large grazing herbivore abundance. Forage quantity, as measured by live, dead and total herbaceous biomass and proportion of biomass alive, was higher inside herbivore exclosures, even at relatively low densities of herbivores. Forage quality, as measured by fibre content, was not affected by herbivory, however, crude protein content of live herbaceous biomass was greater outside herbivore exclosures. Recent fire was an important predictor of all measures of forage quantity and quality. Recent fire occurrence decreased overall quantity (biomass) but increased quality (decreased fibre content and increased crude protein content); late dry season fires resulted in forage with the highest crude protein content. The predictions of the pyric herbivory conceptual model are consistent with observations of the feeding behaviour of introduced bovines and some large macropods in northern Australian savannas, lending support to the global generality of pyric herbivory in fire-prone grassy biomes.

     

Modelling digestive constraints in non-ruminant and ruminant foregut-fermenting mammals

It has been suggested that large foregut-fermenting marsupial herbivores, the kangaroos and their relatives, may be less constrained by food intake limitations as compared with ruminants, due mainly to differences in their digestive morphology and management of ingesta particles through the gut. In particular, as the quality of forage declines with increasing contents of plant fibre (cellulose, hemicelluloses and lignin; measured as neutral-detergent fibre, NDF), the tubiform foregut of kangaroos may allow these animals to maintain food intakes more so than ruminants like sheep, which appear to be limited by fibrous bulk filling the foregut and truncating further ingestion. Using available data on dry matter intake (DMI, g kg(-0.75) d(-1)), ingesta mean retention time (MRT, h), and apparent digestibility, we modelled digestible dry matter intake (DDMI) and digestible energy intake (DEI) by ruminant sheep (Ovis aries) and by the largest marsupial herbivore, the red kangaroo (Macropus rufus). Sheep achieved higher MRTs on similar DMIs, and hence sheep achieved higher DDMIs for any given level of DMI as compared with kangaroos. Interestingly, MRT declined in response to increasing DMI in a similar pattern for both species, and the association between DMI and plant NDF contents did not support the hypothesis that kangaroos are less affected by increasing fibre relative to sheep. However, when DEI was modelled according to DDMIs and dietary energy contents, we show that the kangaroos could meet their daily maintenance energy requirements (MER) at lower levels of DMI and on diets with higher fibre contents compared with sheep, due largely to the kangaroos' lower absolute maintenance and basal energy metabolisms compared with eutherians. These results suggest that differences in the metabolic set-point of different species can have profound effects on their nutritional niche, even when their digestive constraints are similar, as was the case for these ruminant and non-ruminant foregut fermenters.     

Effects of sward structure on herbivore foraging behaviour in a South African savanna: An investigation of the forage maturation hypothesis

Abstract: The grass layer of savannas is characterized by strong temporal and spatial heterogeneity in the quantity and quality of forage. Besides this, there is strong variation in other aspects of sward structure, here defined as sward chemistry, morphology, architecture and species composition. The forage maturation hypothesis (FMH) is based on the temporal dynamics of forage quantity and quality of grasslands. Recently this hypothesis has been used to explain the foraging behaviour of large herbivores as an optimal solution for balancing forage ingestion and forage digestion, leading to a maximization of daily rates of forage intake in patches of intermediate forage quantity. However, so far studies using this hypothesis have been constrained to mono‐specific forage resources or have ignored inter‐patch variation in sward structure. We studied the foraging behaviour of cattle in a South African savanna. We explicitly addressed forage quantity and quality, and the structure of seven sward types in order to investigate (i) their effects on foraging behaviour; and (ii) the assumptions and predictions of the FMH for foraging behaviour in grasslands with pronounced variation in sward structure. The results indicated that the assumptions of the FMH were met and that forage quantity affected forage intake the most, with sward structure having little or no effect. The predicted maximum rate of daily forage intake agreed with the results of previous studies. However, the forage quantity at which this maximum intake rate was found, was larger than expected based on the results of some studies. It is likely that this discrepancy resulted from the use of artificial leaf‐only grass swards in these studies. The results suggest that the FMH can be used to explain the foraging behaviour of herbivores over a wide range of sward structures.     

The ‘omnivorous badger dilemma’: towards an integration of nutrition with the dietary niche in wild mammals

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Resource partitioning between large herbivores in Hustai National Park, Mongolia

Re-introduced Przewalski horses in Hustai National Park, Mongolia could suffer from food competition with other herbivore species through food resource depletion. Diet composition of the Przewalski horse (Equus ferus przewalskii), red deer (Cervus elaphus) and four livestock species (sheep, goat, cattle and horse) were studied, using micro histological analysis of faecal samples in the summer of 2005 and winter of 2006 – 2007. We expected that herbivores become less selective in food choice in winter regarding to summer, resulting in a larger diet breadth, a larger similarity in diet and a larger dietary overlap in winter, potentially triggering exploitative competition by depletion of shared resources. Vegetation biomass decreased during winter, and the different herbivores species in HNP changed their diet from summer to winter. As expected diet breadth, diet similarity and dietary overlap were significantly larger in winter in comparison to summer. The existence of competition by resource depletion between the different species cannot be ruled out. Vegetation biomass was probably not a limiting factor according to the correlation between annual rainfall and herbivore species biomass, however the forage quality may be limiting, triggering competition.     

Large herbivores as mobile links between isolated nature reserves through adhesive seed dispersal

Question: Do large herbivores contribute to the dispersal of plant seeds between isolated habitats by epizoochory? Location: Nature reserves in Flanders, Belgium. Methods: Epizoochory was studied by brushing plant seeds from the fur of 201 domesticated large herbivores (Galloway cattle, donkeys and horses), grazing in 27 Flemish nature reserves. Several herbivores were examined after transport between different nature reserves as part of the seasonal grazing system in Flanders, allowing detection of seed dispersal both within and between reserves. The seedling emergence method was used to identify the dispersed plant species. Results: In total, 6385 epizoochorous seeds from 75 species germinated, yet the real seed quantity was underestimated by the seedling emergence method. A wide variety of seed morphology, seed weights and plant heights was represented among the dispersed species, many of which had a transient seed bank. There was a gradual turnover in epizoochorous species composition in the course of the vegetation season, and seed dispersal occurred both within and between different nature reserves. Conclusions: Domesticated large herbivores, as models for wild mammals in the present and the past, are important dispersers of many plant species. Through seasonal grazing, the herbivores function as ‘mobile link organisms’, connecting isolated nature reserves through seed dispersal, possibly influencing vegetation development and long‐term survival of plant populations. As such, large herbivores are important instruments in ecological restoration, especially in fragmented ecosystems.     

Characterisation of particle dynamics and turnover in the gastrointestinal tract of Holstein cows fed forage diets differing in fibre and protein contents

An improved understanding of the role of forage quality on the processes of particle dynamics and turnover is important for the development of healthier and cost-effective feeding strategies that aim at lowering the proportions of concentrates in the diets of cattle. The aim of this study was to evaluate the effects of feeding hays of different qualities on particle dynamics, digestion kinetics and turnover in the gastrointestinal tract (GIT). Three non-lactating, rumen fistulated Holstein cows were fed diets consisting exclusively of hay with either low quality [Group LH; 605 ± 12.4 g/kg neutral detergent fibre (NDF) and 63 ± 4.7 g/kg crude protein (CP)] or good quality (Group GH; 551 ± 20.1 g/kg NDF and 116 ± 3.6 g/kg CP). Data showed that in situ dry matter (DM) disappearance of the soluble fraction was greater for Group GH (p < 0.05). Feeding good quality hay also lowered the proportion of particles >1.18 mm particularly during the eating process (p < 0.05). Changes in the particle size occurring afterwards were greater for Group GH as well (p < 0.05); approximately 30% in the comminution in the particle size occurred postruminally. Feeding hay of good quality lowered DM content of solid rumen digesta (p < 0.05), accelerated (p < 0.05) the turnover rate of DM and NDF in the GIT and increased DM intake (p < 0.05). In conclusion, feeding forages of better quality significantly promoted degradation processes and kinetics in the GIT with positive effects on turnover rate of digesta and feed intake in Holstein cows.     

Competition between grass‐cutting Atta vollenweideri ants (Hymenoptera: Formicidae) and domestic cattle (Artiodactyla: Bovidae) in Argentine rangelands

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Use of trade-off theory to advance understanding of herbivore–parasite interactions

• 1 Trade‐off theory has been extensively used to further our understanding of animal behaviour. In mammalian herbivores, it has been used to advance our understanding of their reproductive, parental care and foraging strategies. Here, we detail how trade‐off theory can be applied to herbivore–parasite interactions, especially in foraging environments.
• 2 Foraging is a common mode of uptake of parasites that represent the most pervasive challenge to mammalian fitness and survival. Hosts are hypothesized to alter their foraging behaviour in the presence of parasites in three ways: (i) hosts avoid foraging in areas that are contaminated with parasites; (ii) hosts select diets that increase their resistance and resilience to parasites; and (iii) hosts select for foods with direct anti‐parasitic properties (self‐medication). We concentrate on the mammalian herbivore literature to detail the recent advances made using trade‐off frameworks to understand the mechanisms behind host–parasite interactions in relation to these three hypotheses.
• 3 In natural systems, animals often face complex foraging decisions including nutrient intake vs. predation risk, nutrient intake vs. sheltering and nutrient intake vs. parasite risk trade‐offs. A trade‐off framework is detailed that can be used to interpret mammal behaviour in complex environments, and may be used to advance the self‐medication hypothesis.
• 4 The use of trade‐off theory has advanced our understanding of the contact process between grazing mammalian hosts and their parasites transmitted via the faecal–oral route. Experimental manipulation of the costs and benefits of a nutrient intake vs. parasite risk trade‐off has shown that environmental conditions (forage quality and quantity) and the physiological state (parasitic and immune status) of a mammalian host can both affect the behavioural decisions of foraging animals.
• 5 Naturally occurring trade‐offs and the potential to manipulate their costs and benefits enables us to identify the abilities and behavioural rules used by mammals when making decisions in complex environments and thus predict animal behaviour.

     

Impacts of savanna trees on forage quality for a large African herbivore

Recently, cover of large trees in African savannas has rapidly declined due to elephant pressure, frequent fires and charcoal production. The reduction in large trees could have consequences for large herbivores through a change in forage quality. In Tarangire National Park, in Northern Tanzania, we studied the impact of large savanna trees on forage quality for wildebeest by collecting samples of dominant grass species in open grassland and under and around large Acacia tortilis trees. Grasses growing under trees had a much higher forage quality than grasses from the open field indicated by a more favourable leaf/stem ratio and higher protein and lower fibre concentrations. Analysing the grass leaf data with a linear programming model indicated that large savanna trees could be essential for the survival of wildebeest, the dominant herbivore in Tarangire. Due to the high fibre content and low nutrient and protein concentrations of grasses from the open field, maximum fibre intake is reached before nutrient requirements are satisfied. All requirements can only be satisfied by combining forage from open grassland with either forage from under or around tree canopies. Forage quality was also higher around dead trees than in the open field. So forage quality does not reduce immediately after trees die which explains why negative effects of reduced tree numbers probably go initially unnoticed. In conclusion our results suggest that continued destruction of large trees could affect future numbers of large herbivores in African savannas and better protection of large trees is probably necessary to sustain high animal densities in these ecosystems.     

Effects of forage NDF content and body condition score on forage intake by Holstein-Friesian dairy cows in the dry period

This study used individual weekly results for 160 non-lactating Holstein-Friesian dairy cows in the last 5 weeks of gestation to develop regression equations based on forage NDF content and individual body condition score (BCS) for predicting dry matter (DM) intake. Results were used from treatments in which cows received the same forage and no concentrates throughout the dry period. Ten different conserved forages, either grass silages or mixtures of grass silage and barley straw, were fed in six different experiments and forage NDF ranged from 452 to 689 g/kg DM. On average cows gained 390 g live weight per day, which is less than conceptus growth at this stage - suggesting some mobilisation of maternal tissues to support conceptus growth. BCS remained unchanged at 2.5 over the dry period. DM intake declined from 10.79 kg/day 5 weeks before calving to 9.32 kg/day in the week before calving, with half of this decline occurring in the final week before calving. Intake as a percentage of live weight was moderately predicted (R2 = 0.61 for the entire period) from measures of diet composition (NDF) and cow state (BCS). There were highly significant negative effects of forage NDF and increased BCS on DM intake. The effect of BCS on DM intake was greatly reduced in the week before calving, possibly as a result of a change in metabolic priorities from gaining to losing body reserves.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Steller sea lions Eumetopias jubatus and nutritional stress: evidence from captive studies

• 1 Numbers of Steller sea lions Eumetopias jubatus in the North Pacific have declined. According to the nutritional stress hypothesis, this decline is due to reduced food availability. Data from studies conducted on pinnipeds in the laboratory are used here to test if the nutritional stress hypothesis can explain the decline of Steller sea lions.
• 2 Overall, there is strong evidence for biologically meaningful differences in the nutritional quality of major prey species. Steller sea lions can partly compensate for low‐quality prey by increasing their food consumption.
• 3 There appear to be no detrimental effects of low‐lipid prey on sea lion growth or body composition when sea lions can consume sufficient quantities of prey. However, the ability to increase consumption is physiologically limited, particularly in young animals. Overall, it is more difficult to maintain energy intake on a diet of low‐quality prey than on a normal diet.
• 4 Under conditions of inadequate food intake (either due to decreased prey availability or quality, or increased energy requirements) the overall impacts of nutritional stress are complex, and are dependent upon season, prey quality, age and the duration and intensity of the nutritional stress event.
• 5 Studies on pinnipeds in the laboratory have been instrumental in identifying the conditions under which changes in sea lion prey can result in nutritional stress and the nature of the physiological impacts of nutritional stress events.