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Feeding Behaviour
Introduction
The majority of farm animals are fed ad libitum; that is, they have food available almost all of the
time. This is true whether they are kept intensively – the supply of food being under direct control of
their keeper – or extensively, where the herbage available varies in quantity and quality according to
the time of year, but is rarely completely unavailable. Despite this freedom of access to food there are
many situations in which animals over- or undereat, and ‘voluntary food intake’ and the factors which
control and influence it are of great importance to agriculturists across the world.
The Significance of Voluntary Food Intake
Animals compete with man for food and there is a need to reduce the amount of grain-based food,
especially in ruminants, which are able to make use of grass, grass products and by-products of other
agricultural and industrial activities; cereals can thus be saved for human consumption and for pigs
and poultry which cannot digest cellulose, unlike the ruminant. Because the bulk of a food as well as
its nutritive value may limit intake, knowledge of the effects of changes in the type of food offered is
essential.
The scientific study of voluntary food intake is important, therefore, and demands a multidisciplinary
approach. The nutritionist, the physiologist, the psychologist, the pharmacologist, the modeller and, in
the agricultural context, the animal and crop scientist all have to be involved in unravelling the
complexities of the subject.
Definitions
Voluntary food intake: the weight eaten by an animal or group of animals during a given period of
time during which they have free access to food.
Food : the materials that animals eat.
Feed: a portion of food offered to an animal (for ruminants especially, feed is often used to mean
food).
Meals: distinct eating periods, which may include short breaks but which are separated by longer
intervals.
Feeding bouts: short within-meal periods of eating.
Hunger ; when an animal starts to eat.
Satiated: when an animal stops eating.
Hunger ratio: measurement from derived of meal size and inter-meal interval (= weight of meal
divided by pre-meal interval).
Satiety ratio = weight of meal divided by post-meal interval.
Appetite: a drive to eat a specific nutrient, rather than to eat food in general.
Palatability: the overall sensory impression the animal receives from its food.
Rate of eating (rate of intake): the weight of food eaten per unit of time (or during a meal).
voluntary intake: the total amount eaten during a given period of time (usually one day).

Prediction of Food Intake

In order to plan a feeding programme or to decide the optimum formulation of a ration to meet the
animals’ requirements under conditions of ad libitum feeding, it is necessary to be able to predict the
level of voluntary intake of a food or foods by farm animals.
There are two general methods for prediction: empirical equations and mathematical modelling, but
the division between them is not absolute. The first involves measurement of several parameters,
including voluntary intake, followed by regression analysis to find the equation which best fits the
data. For example, analysis of live weight (LW), daily gain (LWG), milk yield (MY) and voluntary
intake (I) measurements in a large number of dairy cows would yield an equation of the form:

I = a + bLW + cLWG + dMY

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which could be used to predict intake in other animals of the same type. Note that this equation is a
‘model’ insofar as live weight, live weight gain and milk yield might reasonably be expected to be
causal factors in the determination of food intake.
The second type of prediction method involves incorporation of more basic biological principles and
functions that describe the relationships between factors likely to underlie the control of food intake.
Such methods are likely to be less accurate at prediction but to be more general in their application.
It is clear that voluntary intake is influenced by both animal and food factors. Many prediction
equations are based on one or the other, so it would be surprising if they gave low errors of prediction
under conditions somewhat different from those under which the data used in their construction were
collected. The major effort in predicting intake has been for ruminants in view of the importance of
physical limitations on intake.

1. Description of Feeding Behaviour and its Development

Poultry
For the first 2 days after hatching, chicks are not dependent on eating food as they have sufficient
reserves from the yolk sac. In the first few hours they eat little or nothing, although they are exploring
their environment and pecking at small, round objects.
Chickens peck jerkily at food items, pick up the grain in their beak and then lift the head back before
swallowing.
Although broiler chickens eat almost twice as much as layers of the same age, they spend only half as
much time eating and take more meals of shorter duration.
Beak trimming
While beak trimming, sometimes carried out on laying hens to overcome fighting, does not affect daily food
intake or meal number or size, beak-trimmed birds spent more time per day eating, so ate more slowly. Trimmed
hens seemed to be more selective as there was a higher proportion of large particles in the food refusals,
suggesting that they found it more difficult to pick up particles, for which the tip of the beak is required. Because
of this selection the food actually eaten by the trimmed birds had somewhat higher crude protein and mineral
contents than for the untrimmed hens.
Cattle
A characteristic of grazing cattle is that they wrap their tongue around herbage and pull rather than biting
cleanly. They walk slowly forward, moving their head from side to side to take mouthfuls as they go.
The feeding behaviour of housed cattle depends to a large extent on the ratio of animal numbers to length of
trough. If there is sufficient space for all animals to eat at once then this they will do, especially when fresh food
has been given or when they have just returned from milking. If, however, there is insufficient space for all
animals to eat simultaneously then a great deal of manoeuvring takes place, dominant animals displacing timid
ones, and meals tend to be a succession of brief episodes interspersed with movement to other feeding positions.
Those cows lower in the social dominance order are forced to eat at less popular times of day, including late at
night.
Intake reache a peak in mid-lactation, while the number of meals per day reached a nadir around week 15 of
lactation. Figure 2.2 shows an example of a 24-h meal pattern of one cow. In this case, nine meals of silage were
taken and three allocations each of 2 kg concentrates were dispensed. At 230 min, she tries to get concentrates
but is not allowed and diverts her attention to eating silage. Some of the silage meals are protracted and
fragmented (e.g. those at 650–750 min), while others are continuous (e.g. that starting at 840 min).
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Fig. 2.2. An example of the meals of silage and concentrates and the drinks of water taken by a
lactating dairy cow during a 24-h period using the LUCIFIR system (D.A. Jackson, J.M. Forbes and
C.L. Johnson, unpublished results). S, meals of silage; C, meals of concentrates; U, visits to the
concentrate dispenser that go unrewarded as the requisite time since last dispensation has not elapsed;
D, drinks of water; E, visits to the concentrate dispenser that go unrewarded as the daily allowance has
been exhausted.
Sheep
From 2 or 3 weeks of age, lambs spend increasing amounts of time nibbling at food. After weaning
they eat by prehending food with their lips and tongue, pulling it back into the mouth for extensive
chewing.
Sheep can select well as they have a narrow bite. However, they have a blind area about 30 mm in
front of the nose so they can’t see clearly what they are eating! Perhaps they use touch to decide
exactly what to eat.
2. Rate of Eating
Poultry
Growing broilers eat faster than layers of the same age, but it is not clear whether this is because they
are bigger and heavier or have a greater need of nutrients. Rates of eating were significantly faster for
pellets than for the same food in powdered form, and declined during the 30 min after giving food
after a 0–4-h fast.
Cattle
Scientists observed that the rate of eating of a complete food by a lactating cow increased linearly
from 50 g/min in the first few days of lactation to 90 g/min in week 9 of lactation. The rate declined as
a large meal proceeded, from 120 to about 50 g/min due to longer pauses in actual chewing as the
meal progressed. Intake rate at the beginning of the meal is four to five times higher than at the end for
cows eating maize silage.
Presumably in systems where food is limited, (dairy cows fed concentrates in a group), the fastest
eaters will consume more and produce more milk. Any cow not having eaten all her allowance when
her group has finished milking will hold up the whole group, or miss part of her ration which will then
be eaten by the next cow to come into that position in the milking parlour. Hence, wet food is eaten
more quickly than the same food given in the dry form
Sheep
Pregnant ewes fed throughout on a complete, pelleted food ate at an average rate of 15 g/min and this
increased to around 20 g/min during lactation and to over 30 g/min after weaning. Rate of eating is
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fast during the beginning of a large meal (first 30 min), and reduces progressively during the middle
and the end (second and third 30 min). Hence, ewes penned in groups eat with a higher rate than for
those penned individually. Detailed studies of long meals have shown an exponential slowing of rate
of eating ( Fig. 2.9).

Fig. 2.9. Cumulative intake of lucerne by a sheep given access for 6h/day: intake (g DM) = 1975(1–
e(–0.0044.time)) (from Baumont et al., 1989).

Short-term Intake Rate

It has been suggested that foods that are initially eaten more quickly than others will have a higher
daily intake, and Short-term Intake Rate (STIR) has been proposed as a way of predicting intake (a test
for intake potential of foods). STIR was defined for cows as the amount eaten in 4 min after a 4-h fast.
However, many factors are likely to affect STIR, including starvation time, length of ingestion period,
form of presentation of food, fibre concentration, DM content, feeding level, body condition of
animals and physiological state of the animals.
Circadian Rhythms of Feeding
Poultry
Unless the nights are very long, poultry do not eat during the hours of darkness, so there is a very
marked diurnal rhythm of feeding. However, a peak of feeding activity and intake have been observed
during the hour before dusk, presumably to fill the crop as a reservoir of food to last the bird through
the night.
Cattle
A high level of feeding activity is usually observed after fresh food is offered. A smaller peak often
occurs before sunset and another just after midnight.
Under conditions where little competition for feeder space exists, dairy cows eat few meals from
around 01.00–06.00 h. When there are more animals than feeder spaces there is strong competition at
busy times, particularly after milking and when fresh food has been delivered. In cows milked three
times per day, we observe a clear triphasic pattern of eating, peaks of feeding activity occurring during
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the periods after return from milking, including large meals. The more dominant animals ate less at
night (01.00–06.00 h) than those at the bottom of the ‘pecking’ order. Therefore, heifers low in the
social dominance order might not be able to satisfy fully their desire to eat when the level of
competition for feeding space was high. The high incidence of aggressive interactions during the
period immediately after return from milking has implications for the health and nutrition of the
animals.
The fact that lower-yielding cows were lower in the dominance order than high yielders but ate more
at night suggests that high demand for nutrients was not the prime cause of nocturnal eating, although
it is possible that low dominance caused low intake, resulting in lower milk yield. It is more likely that
those lower in the dominance order were prevented from meeting their needs during the day and were
thus forced to satisfy their requirements by eating in the middle of the night.
Sheep
Sheep show similar light-entrained circadian rhythms of feeding to cattle (see Fig. 17.7). Another
factor affecting feeding rhythms is environmental temperature: sheep do not eat in the hottest part of
the day when the heat load is so great that they must seek shelter.
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Fig. 17.7. Feeding patterns of ram lambs in (a) 16 h days, (b) 8 h days and (c) 8 h days with ‘flash’ in middle of the night.
Solid bars, hours of darkness; open bars, hours of daylight/‘flash’ (from Schanbacher and Crouse, 1981).

Food Restriction and Fasting

Periods without food leave the animal in a different state than when fully fed. It is important to measure fully this
state in order to predict how the animal will respond when fully fed again.
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Poultry
Pecks at food become faster, stronger and less accurate after increasing periods of deprivation (Wood-
Gush and Gower, 1968) but, when food is unavailable for 1 day in 5, there is complete compensation
of intake and growth.
Food restriction is sometimes used to delay puberty in pullets. When pullets were subsequently
restricted to that weight of food eaten voluntarily at 6 weeks of age (45 g/day), the birds ate and
pecked at empty feeders for the same amount of time per day as ad libitum controls spent eating.
Broiler breeder stock are usually restricted to less than one-half of ad libitum intake. Such animals
have a high rate of working for food, even just after their ration has been consumed, and are highly
motivated to eat at all times.
Cattle
The intake of hay by cows increased by 20% as the increase in time of access to food was increased
from 5 to 24 h/day, in contrast to an 80% increase in intake when the food was a concentrate mixture
(Freer and Campling, 1963), suggesting a physical limit to intake of the hay.
Sheep
When sheep were given access to hay cubes for 3, 6, 12 or 24 h/day, intakes were 1502, 1869, 2086
and 2413 g/day. With fasts of 1–6 h, sheep fed on a pelleted 0.5 dried grass:0.5 barley compound food
compensated for the fasting period in the first two or three meals after the food had been replaced and
there was no effect on daily intake.
Abnormal Feeding-related Behaviour
In intensive husbandry it is not uncommon for abnormal feeding-related behaviours to occur. While
these have been ascribed to boredom, it is also possible that such behaviours as tail biting, stereotypy
and feather pecking are expressions of a desire to obtain nutrient(s) not provided in sufficient
quantities by the diet.
Poultry
Feather pecking can occur in all types of poultry, especially if insufficient trough space is provided. This can be
alleviated by giving birds something to search for, e.g. by giving some whole grains in the food or on the floor.
Feather pecking sometimes leads to body pecking and wounding.
Egg eating by hens may be an indication of mineral deficiency in the diet, as it is reduced by providing grit,
although this again may be a question of lack of variety of activities rather than a nutrient deficiency. Birds eat
shavings on which they are bedded if they have insufficient trough space; this can result in gizzard compaction
and eventual death if not forestalled by increasing the availability of food.
Pecking at metal or wood in a stereotypic manner is sometimes observed in birds, again when the environment is
monotonous. It has been noted that the amount of time spent stereotypically pecking was negatively related to
plasma corticosterone levels, and it has been suggested that such activity relieves stress.
Cattle
Calves spend about 1 h/day sucking naturally from their mothers, but can drink milk replacer much more quickly
than this, leading to boredom and sucking of other calves, particularly the navel. The use of dispensers that
deliver the milk slowly to simulate natural suckling is helpful, as is the provision of straw bedding. Many calves
in crates lick their own hair, a symptom of boredom and possibly of nutrient deficiency. This often creates
hairballs in the stomach, which can occasionally become so large that they block the digestive tract.
Restricted-fed heifers sometimes show stereotypies in the post-feeding period, and those individuals spending
less time eating spend more time exhibiting stereotypies. Feeding-related activities such as sniffing or licking the
feeding trough are seen more frequently around the time that stereotypies are exhibited, suggesting that oral
stereotypies are indicative of frustrated feeding behaviour. Although lactating cows are usually fed ad libitum,
they are often in negative energy balance and higher-yielding cows, which typically lose more weight in early
lactation, have been seen to show more stereotypies.

Sheep and goats

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Sheep will pull and eat the wool of others if too tightly penned, but this can usually be prevented by providing
roughage for them to chew. Agile, young goat does can suck their own teats.
Marsden and Wood-Gush (1986) fed individually penned lambs aged 8 weeks either high (174 g/kg DM) or low
(110 g/kg DM) protein foods ad libitum or restricted to 1 kg/day. More time was spent in abnormal activities
(such as biting empty food bins, wood or wool, ‘star-gazing’) by those on the restricted level of food but, when
hay was offered in addition, there was much less abnormal behaviour.
Conclusions
There are strong circadian rhythms of feeding, but these are not inflexible: for example, chicks will eat during
the dark if that is the only time food is available, even though they normally eat nothing at night. Short periods
without food can be compensated for when food becomes available, but longer periods of fasting result in a
significant reduction in daily intake.
References
J.M. Forbes 2007.Voluntary Food Intake and Diet Selection in Farm Animals 2nd Edition (J.M.
Forbes).
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Feeding Behaviour
The term ‘feeding behaviour’ generally refers to meal-taking activity, in which the time (and weight)
of each meal is monitored. Monitoring and analysing feeding behaviour are areas of considerable
scientific activity, but their importance in understanding and predicting food intake and selection over
longer periods is dubious.
Many factors affect feeding behaviour, including the degree of competition for food and
environmental variables – the weather and management system – without having major impacts on
food intake over periods of several days.
The justification for monitoring feeding behaviour is, on the one hand, to allow measurement of daily
intake in situations in which it cannot be measured directly (e.g. group housing, see below) and, on the
other, to satisfy curiosity and with the hope of gaining understanding about the factors controlling
intake over periods of a day or more.