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Physiological and Metabolic

Aspects of Feed Intake Control
Introduction
The great majority of farm animals are fed ad libitum, that is, with free access to food for
most of the time. To a considerable extent intake is driven by nutrient requirements, and an
approximate estimate of how much food an animal will eat can be calculated from what
quantities of energy, protein and nutrients it requires for maintenance and production.
However, there are numerous factors which can interfere with the concordance between
requirements and intake.
It is generally considered that intake is controlled by a series of negative feedback signals
from the digestive tract, liver and other organs in response to the presence of nutrients.
In addition, animals learn the metabolic consequences of eating foods with particular sensory
properties (appearance, flavour, texture) and can then use ‘feedforward’ to choose
preferentially or avoid foods which they have experienced previously (Fig. 15.1.)

Fig. 15.1. Satiety cascade (after Blundell and Halford, 1994).

Of course, the whole system is coordinated by the CNS in a diverse set of pathways that
includes the hindbrain and the hypothalamus as important components.
1. Mechanisms of Intake Control
Central nervous system
The brain collects information from the special senses and receptors in the digestive tract wall
and metabolizing tissues. This information is integrated and used to determine what food to
eat and whether feeding should start or stop. The anatomical sites of the brain involved in
intake control are : The ventromedial hypothalamus (depletion center), the lateral
hypothalamus (repletion center), and the paraventricular nuclei of the hypothalamus, just
ventral to the ventromedial nuclei, which are particularly sensitive to the effects of transmitter
chemicals, including noradrenaline and neural peptide Y (NPY).
While the hypothalamus and surrounding parts of the forebrain are likely to be the seat of
intake control, there are centres in the hindbrain, such as the nucleus of the solitary tract,
which receive information from receptors in the visceral organs such as the stomach and liver.
Also in this area are neurons directly sensitive to shortage (but not to excess) of energy
yielding substrates.
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Transmitters stimulate abdominal receptors which transmit information to the CNS via the
vagal pathway.
Gastrointestinal receptors
The ingestion of food causes changes in the degree of fill and the chemical composition of
digesta which can be sensed by stretch receptors and chemoreceptors in the wall of the
digestive tract.
Simple-stomached animals
Although it is not usually considered that stomach fill is a predominant factor limiting intake
in pigs or poultry, extremes of distension can certainly limit intake, and stretch of stomach
and intestine wall may contribute to satiety. Chemical composition can also contribute to
satiety (control intake), but it is less important than the physical presence. For example,
duodenal infusion of glucose solutions inhibits feeding in chickens. This information is used
by the brain, along with that from many other abdominal receptors, to determine whether
feeding should stop or proceed.
As stated above, CCK is secreted by the wall of the duodenum in response to the passage of
digesta, particularly fat and protein, and stimulates receptors locally which relay their
information to the CNS where it results in a decrease in food intake.
Ruminants
PHYSICAL LIMITS. Despite the fact that the rumen has such a large capacity, the slow rate
of digestion of forage feeds means that rumen capacity can be limiting to intake (see Chapter
16). There are stretch receptors in the rumen wall, especially in the anterior dorsal part, and
these signal the degree of distension to the brain via the vagus nerves. However, there are
other chemical and metabolic factors involved in intake control in ruminants, as in
simple-stomached animals.
CHEMICAL FACTORS. The stretch receptors in the rumen wall are also sensitive to
chemicals, including the acids produced by rumen fermentation. Figure 15.4 shows the effects
of infusion into the rumen of sodium acetate or propionate, and of distension, on the intake of
hay or silage by lactating cows prepared with rumen fistulas (Anil et al., 1993). Some of the
effect of introduction of salts into the rumen is due to their osmotic properties.

Fig. 15.4. Intake of grass silage by lactating dairy cows with a balloon inflated to different
volumes (–––, level of treatment = litres) for 3 h or infused with sodium acetate (....) or
sodium propionate (----) at different rates (level of treatment = mol 3 h_1) (Anil et al., 1993).
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Liver
The liver is the first site after absorption which is able to monitor the results of eating a meal,
although lipids are absorbed into the lymphatic system and might not, therefore, be able to
exert their full effect on the liver before being utilized by some other tissues. There are
receptors in the liver sensitive to glucose and to other metabolites that are oxidized in the liver
(Forbes, 1988); where increasing in their concentration depress intake; the mechanism is
probably an activation of the sodium pump which changes the transmembrane potential of the
hepatocytes resulting in a change in the rate of impulses generated in afferent fibres of the
autonomic nerves (vagal and sympathetic nerves) terminating in the liver. It was demonstrated
that the blood is not involved in transmission of the metabolic information from liver to the
CNS (Anil and Forbes, 1988).
Ruminant animals rely on liver uptake of propionate and its use for glucose synthesis. Figure
15.5 shows that infusion of sodium propionate into the hepatic portal vein of sheep depresses
food intake relative to a control infusion of sodium chloride, while infusion of propionate into
the general circulation via the jugular vein had no effect (Anil and Forbes, 1980).

Fig. 15.5. Intake of pelleted food by sheep infused with (A) saline (Cont) or sodium
propionate into the mesenteric (Prop) or jugular (Jugul) veins (Anil and Forbes, 1980); (B)
saline (Cont) or sodium propionate into the mesenteric vein without (Prop) or with (Block)
anaesthetic blockade of the splanchnic nerves (Anil and Forbes, 1988).

Infusions into the general circulation

Mayer envisaged that the CNS monitored the concentration of glucose in the blood; when it
was below a certain threshold, feeding started while when it was above a threshold the meal
was terminated. Several pieces of evidence suggested that the receptors responsible were in
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the ventromedial nuclei of the hypothalamus but, while there might be hypothalamic
sensitivity to blood glucose concentration, significant fluctuations in this concentration would
mean that the body had failed in its task of maintaining a relative constancy of nutrient supply
to the brain. Else, there are receptors in intestines and liver that normally prevent over- or
undereating, and the focus of attention has moved away from the concentration of glucose in
the general circulation as a major factor in the control of intake.
Adipose tissue
Metabolites in blood must be either utilized or stored by tissues, such as adipose and muscle,
or else excreted by the kidneys. The relative constancy of body weight suggests that there is a
signal from adipose tissue which is integrated with those other signals already mentioned and
which alters feeding behaviour in a subtle manner to adjust for any changes in body fat
content.this signal is a humoral factor which act by feedback. The nature of the humoral
factor was unknown but it was speculated that it might be a steroid hormone. However, more
recent evidence has implicated insulin, and it now seems clear that the most important
feedback signal from adipose tissue to the CNS is leptin.
Insulin
In the short term, it has been suggested that the rise in insulin secretion during spontaneous
meals is responsible for fed intake termination. However, in long term, insulin has often been
observed to cause an increase in intake, probably in response to the increase in rate of fat
deposition and growth, suggesting that an insulin-mediated control of intake by fatness is not
a primary means of control of intake.

Leptin
Leptin (ob-protein) is a hormone, secreted by adipose cells in proportion to their size, which
circulates in the bloodstream and influences receptors in the brain. It appears to interact with
the insulin and NPY systems and thus to be part of an important control system for food
intake in relation to fatness and metabolism.
…………………………………………………………………………………………………...
Integration of factors
Although it has sometimes been stated that only one factor controls intake in any given set of
circumstances, this does not seem to be a tenable point of view. However, experiments
demonstrated that physical factors (stretch signals) are integrated with the metabolic signals,
and there is evidence that this integration is by simple addition. Figure 15.6 shows that the
effects of distending a balloon in the rumen are additive with those of infusing sodium acetate
and sodium propionate (Mbanya et al., 1993) and there is also evidence for such additivity in
sheep and of glucose and lysine given into the liver of chickens.
The implications are that voluntary intake is not controlled only by physical factors, even in
ruminants, and that it is the sum total of the strengths of signals received by the brain from
many types of receptors in many parts of the body which determines how much an animal
eats.
Thus we have an explanation for the fact that lactating cows have a greater amount of digesta
in the rumen than non-lactating cows even when they are both fed on a forage whose intake
traditionally would be supposed to be limited physically (Forbes, 1986). The fact that the
lactating cow has a much greater demand for nutrients to support lactation causes a more
rapid removal of chemical and metabolic feedback factors from blood, liver and digestive
tract, thus providing a weaker set of signals from metabolic receptors which gives the animal
the opportunity to fill her rumen to a greater extent before the total of all the signals reaches a
satiating level.
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Fig. 15.6. Intake of silage DM by lactating cows with or without infusion of sodium acetate
(9.0 mol 3 h_1, A) or sodium propionate (4.9 mol 3 h_1, P) or distension of a balloon (10 l,
D) in all combinations (Mbanya et al., 1993).

2. Diet Selection
So far we have considered only those situations in which animals have a single food available
so that they have no opportunity to alter the nutritive value of the food other than by eating
more or less food.
One of the clearest demonstrations of ‘nutritional wisdom’ is the ability of the growing pig to
select between foods with protein contents higher and lower than that required, so as to
provide sufficient protein for growth without overconsumption. They avoid an excess protein
intake as much as they avoid a deficiency, presumably as both make them feel metabolically
uncomfortable and they learn to avoid such discomfort by appropriate diet selection.
Growing broiler chickens demonstrate a clear ability to select a protein intake appropriate to
their requirements and, like growing pigs, choose progressively less protein, relative to
energy, as they get older, to match the declining requirements for protein relative to energy
(Shariatmadari and Forbes, 1993) like shown in figure figure 15.7.
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Fig. 15.7. Proportion of high- and low-protein foods taken by chickens from 27 to 63 days
after hatching (Shariatmadari and Forbes, 1993). ––––: choice between foods containing 225
and 65 g kg_1;– – –: 280 and 65 g kg_1; .....: 280 and 225 g kg_1; –..–: 320 and 280 g kg_1.
Thus, growing pigs and poultry show considerable ‘nutritional wisdom’ and there is evidence
that ruminants also do (e.g. Kyriazakis and Oldham, 1993). However, in addition to
optimizing nutritional balance, they also choose sufficient fibrous food to stabilize rumen
fermentation and prevent rapid changes in fermentation and unstable pH. For example, sheep
changed from long hay to a diet free of long fibre immediately started to eat chopped
polyethylene fibre which they previously ignored (Campion and Leek, 1997).
Specific appetites
Protein supplies many amino acids, and the above-mentioned ability to balance protein and
energy intakes independently is not a clear example of a specific appetite. However, poultry
have been shown to select for lysine, methionine, thiamine (vitamin B1), vitamin B6, ascorbic
acid (vitamin C), calcium and zinc (see Forbes, 1995)0 while ruminants have a
well-developed appetite for sodium. Figure 15.8 shows the ascorbic acid intake chosen by
young broiler chicks by selecting between an unsupplemented food and one supplemented
with 200 mg kg1 ascorbic acid (Kutlu and Forbes, 1993). When kept in heat-stressing
conditions, which are known to increase the metabolic requirement for ascorbic acid, the
chicks voluntarily take in almost twice as much of the vitamin as when in a thermoneutral
environment.
They can distinguish between the two foods only by their colour and learn to associate this
with their metabolic feelings after consuming the foods.
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Fig. 15.8. Intake of ascorbic acid (vitamin C) by young chicks given a choice between two
foods, coloured differently, containing 0 or 250 mg g_1 ascorbic acid. Cool: thermoneutral
zone; Hot: 35°C for 10 h day_1.
*, significantly different from random choice between foods (Kutlu and Forbes, 1993).

Optimizing Voluntary Intake

In practice, marginal deficiencies of a nutrient result in an increase in intake but severe
shortage of an essential nutrient causes intake to decline (Fig. 15.9) (Forbes, 1995), causing a
reduction in rate of growth, milk production or egg production. Alleviation of the deficiency
is followed very quickly by a return to normal levels of intake and production. Also, most
nutrients are toxic when present at very high levels in the diet so that intake and performance
are also reduced when a gross excess of a nutrient is present.
These general effects of diet composition also affect the intake of ruminant animals. However,
with the forage feeds typically available to ruminants, there is another factor which often
imposes a limit to intake before an animal can satisfy its appetite for energy or essential
nutrients, and this is the bulk of the food.
Although the ruminant has a very capacious set of stomachs (up to 150 litres in the cow), the
slow rate of digestion entails each particle of food remaining in the rumen for a very long
time, typically 20–50 h; the more fibrous the forage, the slower its digestion. Rumen fill,
acting through stretch receptors in the rumen wall, is therefore likely to be a dominant factor
in the intake of forages. Any factor which decreases this residence time will increase
voluntary intake; such factors include reducing the particle size of the food itself, increasing
the activity of the rumen microflora by providing additional amounts of nutrients which limit
their metabolism, and replacing some of the forage by rapidly digested concentrate
supplements.
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Fig. 15.9. General relationship between concentration of an essential nutrient in food and rate
of voluntary food intake (Forbes, 1995).

Conclusions
Voluntary food intake is influenced by many factors including negative feedbacks from the
consequences of eating a meal. If the nutrients present are in the same ratio as required, then
food intake will be optimal, i.e. it will meet the animal’s requirements. Choice feeding is
difficult to implement in practice and animals sometimes make nutritionally unwise choices,
the reasons for which are not always clear (Forbes and Kyriazakis, 1995).
To provide for the animals’ needs in farming practice, the composition of the food is usually
changed at regular intervals during growth, lactation or egg-laying, to match the changes in
requirements. With ruminants, a basal forage is usually offered ad libitum, with
supplementary foods of compositions and in amounts judged to balance the diet.
There is no single factor responsible for the control of voluntary food intake. Rather, a
multitude of influences are coordinated by the CNS and the animal eats that amount and
mixture of food that it learns give the minimum of discomfort.