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.

The content of ruminally fermented OM in the diet affects the fiber requirement of dairy cattle. Physically effective fiber is the fraction of feed that stimulates chewing activity. Chewing, in turn, stimulates saliva secretion. Bicarbonate and phosphate buffers in saliva neutralize acids produced by fermentation of OM in the rumen. The balance between the production of fermentation acid and buffer secretion is a major determinant of ruminal pH. Low ruminal pH may decrease DMI, fiber digestibility, and microbial yield and thus decrease milk production and increase feed costs. Diets should be formulated to maintain adequate mean ruminal pH, and variation in ruminal pH should be minimized by feeding management. The fraction of OM that is fermented in the rumen varies greatly among diets. This variation affects the amount of fermentation acids produced and directly affects the amount of physically effective fiber that is required to maintain adequate ruminal pH. Acid production in the rumen is due primarily to fermentation of carbohydrates, which represent over 65% of the DM in diets of dairy cows and have the most variable ruminal degradation across diets. The non-fiber carbohydrate content of the diet is often used as a proxy for ruminal fermentability, but this measure is inadequate. Ruminal fermentation of both nonfiber carbohydrate and fiber is extremely variable, and this variability is not related to the nonfiber carbohydrate content of the diet. The interaction of ruminally fermented carbohydrate and physically effective fiber must be considered when diets for dairy cattle are evaluated and formulated.

Feed and energy intake of ruminant animals can change dramatically in response to changes in diet composition or metabolic state, and such changes are poorly predicted by traditional models of feed intake regulation. Recent work suggests that temporal patterns of fuel absorption, mobilization, and metabolism affect feed intake in ruminants by altering meal size and frequency. Research with nonruminants suggests that meals can be terminated by signals carried from the liver to the brain via afferents in the vagus nerve and that these signals are affected by hepatic oxidation of fuels and generation of ATP. We find these results consistent with the effects of diet on feed intake of ruminants. Of fuels metabolized by the ruminant liver, propionate is likely a primary satiety signal because its flux to the liver increases greatly during meals. Propionate is utilized for gluconeogenesis or oxidized in the liver and stimulates oxidation of acetyl CoA. Although propionate is extensively metabolized by the ruminant liver, there is little net metabolism of acetate or glucose, which may explain why these fuels do not consistently induce hypophagia in ruminants. Lactate is metabolized in the liver but has less effect on satiety, probably because of greater latency for reaching the liver within meals and because of less hepatic extraction compared with propionate. Hypophagic effects of fatty acid oxidation in the liver are likely from delaying hunger rather than promoting satiety because beta-oxidation is inhibited during meals by propionate. A shortage of glucose precursors and increased fatty acid oxidation in the liver for early lactation cows lead to a lack of tricarboxylic acid (TCA) cycle intermediates, resulting in a buildup of the intracellular acetyl-CoA pool and export of ketone bodies. In this situation, hypophagic effects of propionate are likely enhanced because propionate entry into the liver provides TCA cycle intermediates that allow oxidation of acetyl-CoA. Oxidizing the pool of acetyl-CoA rather than exporting it increases ATP production and likely causes satiety despite the use of propionate for glucose synthesis. A better understanding of metabolic regulation of feed intake will allow diets to be formulated to increase the health and productivity of ruminants.

Even under the intensive concentrate feeding systems of ruminant animal production in the United States, forages continue to represent the single most important feed resource. Cell-wall concentration and digestibility limit the intake potential and energy availability of forage crops in beef and dairy production. Identification of cell-wall characteristics that should be targets of genetic modification is required if plant breeders and molecular biologists are to successfully improve forages for livestock feeding. As the forage plant cell develops, phenolic acids and lignin are deposited in the maturing cell wall in specific structural conformations, and in a strict developmental sequence. Lignin is the key element that limits cell-wall digestibility, but cross-linkage of lignin and wall polysaccharides by ferulic acid bridges may be a prerequisite for lignin to exert its affect. Lignin composition and p-coumaric acid in the wall are less likely to affect digestibility. Voluntary intake of forages is a critical determinant of animal performance and cell-wall concentration is negatively related to intake of ruminants consuming high-forage diets. Cell walls affect intake by contributing to ruminal fill. A simple model of cell-wall digestion and passage in which ruminal fill is a function of rates of digestion and passage, as well as the indigestible fraction of the cell-wall indicates that cell-wall concentration and rate of passage are the most critical parameters determining ruminal fill. Plant factors that affect rate of passage include those that affect particle size reduction by chewing and those that affect particle buoyancy in the rumen. The latter is primarily affected by 1) the ability of the particulate matter to retain gases, which is probably related to plant anatomy and rate of digestion of the plant tissue, and 2) the rate at which the gas is produced, which is affected by the potentially digestible fraction of the particulate matter and the rate of digestion of this fraction. Increasing rate of digestion should increase rate of passage by diminishing the gas produced and increasing density over time. A reduction in the indigestible cell-wall fraction is beneficial because this will decrease fill and increase digestibility. Animal production and economic benefits from reduced cell-wall concentration and increased digestibility are significant. Because of the high cell-wall concentration and large digestible cell-wall fraction of grasses, reduction in cell-wall concentration would probably be of greater value than improving digestibility in these species. Legumes represent the opposite situation and may benefit more from improvements in the digestibility of their cell walls.