Drives and limits to feed intake in ruminants

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Abstract

The control of energy intake is complex, including mechanisms that act independently (e.g. distention, osmotic effects, fuel-sensing) as well as interacting factors that are likely to affect feeding via their effects on hepatic oxidation. Effects of ruminant diets on feed intake vary greatly because of variation in their filling effects, as well as the type and temporal absorption of fuels. Effects of nutrients on endocrine response and gene expression affect energy partitioning, which in turn affects feeding behaviour by altering clearance of fuels from the blood. Dominant mechanisms controlling feed intake change with physiological state, which is highly variable among ruminants, especially through the lactation cycle. Ruminal distention might dominate control of feed intake when ruminants consume low-energy diets or when energy requirements are high, but fuel-sensing by tissues is likely to dominate control of feed intake when fuel supply is in excess of that required. The liver is likely to be a primary sensor of energy status because it is supplied by fuels from the portal drained viscera as well as the general circulation, it metabolises a variety of fuels derived from both the diet and tissues, and a signal related to hepatic oxidation of fuels is conveyed to feeding centres in the brain by hepatic vagal afferents stimulating or inhibiting feeding, depending on its energy status. The effects of somatotropin on export of fuels by milk secretion, effects of insulin on gluconeogenesis, and both on mobilisation and repletion of tissues, determine fuel availability and feed intake over the lactation cycle. Control of feed intake by hepatic energy status, affected by oxidation of fuels, is an appealing conceptual model because it integrates effects of various fuels and physiological states on feeding behaviour.

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Effects of diet on short-term regulation of feed intake by lactating dairy cattle.

M Allen (2000)

Physical and chemical characteristics of dietary ingredients and their interactions can have a large effect on dry matter intake (DMI) of lactating cows. Physical limitations caused by distension of the reticulo-rumen or other compartments of the gastrointestinal tract often limit DMI of high producing cows or cows fed high forage diets. Fermentation acids also limit DMI from a combination of increased osmolality in the reticulo-rumen and specific effects of propionate, although the mechanisms are not clear. The specific physical and chemical characteristics of diets that can affect DMI include fiber content, ease of hydrolysis of starch and fiber, particle size, particle fragility, silage fermentation products, concentration and characteristics of fat, and the amount and ruminal degradation of protein. Site of starch digestion affects the form of metabolic fuel absorbed, which can affect DMI because absorbed propionate appears to be more hypophagic than lactate or absorbed glucose. Dry matter intake is likely determined by integration of signals in brain satiety centers. Difficulty in measurement and extensive interactions among the variables make it challenging to account for dietary effects when predicting DMI. However, a greater understanding of the mechanisms along with evaluation of animal responses to diet changes allows diet adjustments to be made to optimize DMI as well as to optimize allocation of diet ingredients to animals. This paper discusses some of the characteristics of dietary ingredients that should be considered when formulating diets for lactating dairy cows and when allocating feeds to different groups of animals on the farm.

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Rebeca Acín-Pérez, Eric Salazar, Margarita Kamenetsky … (2009)

Mitochondria constantly respond to changes in substrate availability and energy utilization to maintain cellular ATP supplies, and at the same time control reactive oxygen radical (ROS) production. Reversible phosphorylation of mitochondrial proteins has been proposed to play a fundamental role in metabolic homeostasis, but very little is known about the signaling pathways involved. We show here that protein kinase A (PKA) regulates ATP production by phosphorylation of mitochondrial proteins, including subunits of cytochrome c oxidase. The cyclic AMP (cAMP), which activates mitochondrial PKA, does not originate from cytoplasmic sources but is generated within mitochondria by the carbon dioxide/bicarbonate-regulated soluble adenylyl cyclase (sAC) in response to metabolically generated carbon dioxide. We demonstrate for the first time the existence of a CO(2)-HCO(3)(-)-sAC-cAMP-PKA (mito-sAC) signaling cascade wholly contained within mitochondria, which serves as a metabolic sensor modulating ATP generation and ROS production in response to nutrient availability.

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Physical constraints on voluntary intake of forages by ruminants.

M Allen (1996)

Voluntary dry matter intake (VDMI) of forages by ruminants may be limited by distention resulting from restricted flow of digesta through the gastrointestinal tract. An animal's capacity for fill depends on the weight and volume of digesta that causes distention and the flow rate of digesta from the organ in which distention occurs. The reticulorumen is generally regarded as the site in the gastrointestinal tract for which distention limits VDMI with high-fill diets, although evidence suggests that distention of the abomasum may also limit VDMI. Linear decreases in VDMI have been noted with increasing amounts of inert fill inserted into the reticulorumen, but results have not been consistent across several experiments. Reduction in VDMI depends on the extent to which intake is limited by fill before insertion of inert fill; hence animals with high energy requirements consuming relatively low-energy, high-fill diets are affected to the greatest extent. Because NDF generally ferments and passes from the reticulorumen more slowly than other dietary constituents, it has a greater filling effect over time than non-fibrous feed components and has been found to be the best single chemical predictor of VDMI. However, many other factors affect fill, including particle size, chewing frequency and effectiveness, particle fragility, indigestible NDF fraction, rate of fermentation of the potentially digestible NDF, and characteristics of reticular contractions. These factors are only partially accounted for in models that have been developed to predict VDMI. Increased accuracy of prediction of VDMI is expected as models continue to evolve.