ORIGINAL ARTICLE T.H.

Friend-Stress

Stress: What Is It and How and the son unexpectedly walked into the room. There is also evidence that
psychological stressors such as being confronted with conspecifics or species

Can It Be Quantified? enemies may be more potent in eliciting physiological reactions indicative of
stress than physical stressors such as electrical shock applied to the feet of an
animal (Peters, 1977).
According to Hans Selye (1973), who is generally considered to be the
founder of human stress research, stress is not something to be avoided and in
T.H. Friend* fact, cannot be avoided. No matter what an animal does, there arises a demand
to provide the necessary energy to resist and adapt to the changing environment.
Selye continues to point out that complete freedom from stress is death. Con-
trary to public opinion, we must not- and indeed cannot- avoid stress.

Abstract Current methods of measuring or quantifying stress of animals involve

measuring its manifestations. Once a stressor is perceived, the animal reacts
An animal may be considered to be in a state of stress if abnormal behavior or either behaviorally, physiologically, or most often, in combination, to enable it to
extreme adjustments in its behavior or physiology are necessary in order to cope resist and perhaps alleviate the challenge. The use of behavioral observations,
with adverse aspects of its environment. Methods used to determine if an animal is physiological measures and combinations of both as measures of stress will be
stressed can be either behavioral or physiological. Behavioral methods may be discussed as they apply to farm animals.
highly erroneous due to their subjective nature since alterations in behavior do not
necessarily prove that an animal is stressed. There is no single measure of stress Behavioral Measures of Stress
that can be used in all situations at this time. Every measure must be critically
evaluated to ensure that it is valid for each application. An integrated approach us- A stressful condition obviously exists, for example, when overt fighting oc-
ing both behavioral observations and physiological measures is necessary. curs or when an animal is frantically trying to break out of a cage. Measures of
chronic or long-term stress, the behavioral signs of which can be very subtle, are
far more controversial. What has been presented in the popular press as
measures of stress range from subjective, anthropomorphic evaluations with no
Introduction foundation to well substantiated, scientifically acceptable measures. Even the
best measures, however, must be critically examined for each application.
Stress is a phenomenon which everyone acknowledges, but few can agree on Several of the most frequently used methods are discussed below.
a definition. Fraser eta/., (1975) discussed various definitions and concepts of
stress in a nine page paper in the British Veterinary journal. These authors propos- Occurrence of a Specific Abnormal Behavior
ed that an animal is in a state of stress if it is required to make abnormal or ex- The occurrence of a specific "abnormal" behavior such as tailbiting in
treme adjustments in its physiology or behavior in order to cope with adverse groups of swine kept in confinement, sterotyped movements (e.g., incessant pac-
aspects of its environment or management. The behavioral and physiological ing in zoo animals or stall weaving in horses), or other displacement or vacuum
responses to a challenge are the effects of stress. activities, are frequently presented as possible indicators of stress. There are,
It should also be recognized that animals can make both physiological and however, serious problems with their use. The behavior of an animal is very com-
behavioral adjustments in the presence of a stressful situation which can plex and even a specific behavior can be attributed to many different factors.
eliminate that event from being a stressor (Moberg, 1976). In fact, morphine-like The tailbiting syndrome in swine is a very good example of the complexity of
compounds (endorphins), which promote emotional detachment from the experi- an apparently simple behavior. In acute tailbiting many of the pigs in a pen may
ence of suffering, have recently been identified in the brain (Goldstein, 1976). be afflicted. In extreme cases, the attacking pig or pigs may continue to consume
There is speculation that they may aid in tolerating stressful conditions. the victim, resulting in crippling and perhaps death. Frequently there is only one
It is immaterial whether the stressor is pleasant or unpleasant. All that tailbiter and if the attacker is not removed from the group, it may be the only pig
counts is the intensity (Selye, 1973). Parents who are suddenly told that their only with an intact tail. The incidence of tailbiting can be increased by potential
son died in battle would suffer the same stress if the news turned out to be false stressors such as artificially high levels of ammonia gas (Van Putten, 1969).
Hence, people often perceive it as a sign of stress, but generally the cause is more
complex. Pigs are "poor sports." If chased from a feeder by a more dominant pig,
* Dr. Friend is Assistant Professor in the Department of Animal Science, Texas A&M University, Col-
lege Station, Texas 77843. This paper was prepared for and presented at The Institute for the Study of they will frequently retaliate by biting the tail of the defenseless dominant pig
Animal Problems Symposium on Pain, Stress and Suffering: Definition, Quantitation and Application to while its head is in a feeder. The tailbiter is frequently one of the smallest or a
Animal Welfare Issues, Orlando, Florida, November 7, 1979. subordinate in the group, which supports the retaliation theory. Removing the

366 /NT 1 STUD ANIM PROB 1(6] 1980 /NT 1 STUD ANIM PROB 1(6) 1980 367
 ORIGINAL ARTICLE T.H. Friend-Stress

Stress: What Is It and How and the son unexpectedly walked into the room. There is also evidence that
psychological stressors such as being confronted with conspecifics or species

Can It Be Quantified? enemies may be more potent in eliciting physiological reactions indicative of
stress than physical stressors such as electrical shock applied to the feet of an
animal (Peters, 1977).
According to Hans Selye (1973), who is generally considered to be the
founder of human stress research, stress is not something to be avoided and in
T.H. Friend* fact, cannot be avoided. No matter what an animal does, there arises a demand
to provide the necessary energy to resist and adapt to the changing environment.
Selye continues to point out that complete freedom from stress is death. Con-
trary to public opinion, we must not- and indeed cannot- avoid stress.

Abstract Current methods of measuring or quantifying stress of animals involve

measuring its manifestations. Once a stressor is perceived, the animal reacts
An animal may be considered to be in a state of stress if abnormal behavior or either behaviorally, physiologically, or most often, in combination, to enable it to
extreme adjustments in its behavior or physiology are necessary in order to cope resist and perhaps alleviate the challenge. The use of behavioral observations,
with adverse aspects of its environment. Methods used to determine if an animal is physiological measures and combinations of both as measures of stress will be
stressed can be either behavioral or physiological. Behavioral methods may be discussed as they apply to farm animals.
highly erroneous due to their subjective nature since alterations in behavior do not
necessarily prove that an animal is stressed. There is no single measure of stress Behavioral Measures of Stress
that can be used in all situations at this time. Every measure must be critically
evaluated to ensure that it is valid for each application. An integrated approach us- A stressful condition obviously exists, for example, when overt fighting oc-
ing both behavioral observations and physiological measures is necessary. curs or when an animal is frantically trying to break out of a cage. Measures of
chronic or long-term stress, the behavioral signs of which can be very subtle, are
far more controversial. What has been presented in the popular press as
measures of stress range from subjective, anthropomorphic evaluations with no
Introduction foundation to well substantiated, scientifically acceptable measures. Even the
best measures, however, must be critically examined for each application.
Stress is a phenomenon which everyone acknowledges, but few can agree on Several of the most frequently used methods are discussed below.
a definition. Fraser eta/., (1975) discussed various definitions and concepts of
stress in a nine page paper in the British Veterinary journal. These authors propos- Occurrence of a Specific Abnormal Behavior
ed that an animal is in a state of stress if it is required to make abnormal or ex- The occurrence of a specific "abnormal" behavior such as tailbiting in
treme adjustments in its physiology or behavior in order to cope with adverse groups of swine kept in confinement, sterotyped movements (e.g., incessant pac-
aspects of its environment or management. The behavioral and physiological ing in zoo animals or stall weaving in horses), or other displacement or vacuum
responses to a challenge are the effects of stress. activities, are frequently presented as possible indicators of stress. There are,
It should also be recognized that animals can make both physiological and however, serious problems with their use. The behavior of an animal is very com-
behavioral adjustments in the presence of a stressful situation which can plex and even a specific behavior can be attributed to many different factors.
eliminate that event from being a stressor (Moberg, 1976). In fact, morphine-like The tailbiting syndrome in swine is a very good example of the complexity of
compounds (endorphins), which promote emotional detachment from the experi- an apparently simple behavior. In acute tailbiting many of the pigs in a pen may
ence of suffering, have recently been identified in the brain (Goldstein, 1976). be afflicted. In extreme cases, the attacking pig or pigs may continue to consume
There is speculation that they may aid in tolerating stressful conditions. the victim, resulting in crippling and perhaps death. Frequently there is only one
It is immaterial whether the stressor is pleasant or unpleasant. All that tailbiter and if the attacker is not removed from the group, it may be the only pig
counts is the intensity (Selye, 1973). Parents who are suddenly told that their only with an intact tail. The incidence of tailbiting can be increased by potential
son died in battle would suffer the same stress if the news turned out to be false stressors such as artificially high levels of ammonia gas (Van Putten, 1969).
Hence, people often perceive it as a sign of stress, but generally the cause is more
complex. Pigs are "poor sports." If chased from a feeder by a more dominant pig,
* Dr. Friend is Assistant Professor in the Department of Animal Science, Texas A&M University, Col-
lege Station, Texas 77843. This paper was prepared for and presented at The Institute for the Study of they will frequently retaliate by biting the tail of the defenseless dominant pig
Animal Problems Symposium on Pain, Stress and Suffering: Definition, Quantitation and Application to while its head is in a feeder. The tailbiter is frequently one of the smallest or a
Animal Welfare Issues, Orlando, Florida, November 7, 1979. subordinate in the group, which supports the retaliation theory. Removing the

366 /NT 1 STUD ANIM PROB 1(6] 1980 /NT 1 STUD ANIM PROB 1(6) 1980 367
 T.H. Friend-Stress T.H. Friend-Stress Original Article I
Original Article I

tailbiter will eliminate tailbiting in the group. One would expect another tailbiter to examine the health records of both populations to make a meaningful deci-
to evolve if the activity were caused by stressful conditions, but this is not the sion. There is a good chance of discovering that the population with no stress-
case. Also, in a typical, very large confinement farm the author observed related disease also had a much greater mortality rate.
tailbiting in only one pen. If tailbiting is an indication of stress, what can be said
about the other forty-nine identical pens in which it did not occur? Other poten- Change in Behavior Patterns
tial causes and cures of tailbiting are covered in numerous review articles (e.g., This procedure generally establishes fairly quantitative behavioral patterns
Fritschen, 1978). (e.g., time standing, time resting, or number of vocalizations) as ind~ces of str~s.s
The appearance of displacement or vacuum activities does not necessarily or the overtaxing of resources. This approach was used to determme the mml-
prove that an animal is stressed. There may be signs of boredom or frustration mum number of free stalls (resting sites) required by dairy cattle in confinement
(Wood-Gush et a/., 1975; Hediger, 1950), but this may also indicate that the barns before they became overcrowded (Friend eta/., 1977b). Time spent resting
animal is coping with the situation. People will often do something they would per day was determined using time lapse photography as th_e n~mber_ of ~ree
not normally do to relieve boredom or frustration. Following this line of thought, stalls were reduced at seven day intervals. A significant reduct1on m restmg t1me
we could even rationalize that it is the animal which does not show displacement occurred at .50 and .33 stalls per cow (Table 1). However, we could not say that
activities (or, the person who fails to release his or her emotions) that is most the cattle were stressed based on this change in behavioral patterns alone. These
probably stressed. It can also be argued that some vacuum activities are instinc- data indicated only at what density the cattle were no longer able to rest as often
tive behavior patterns (e.g., nest building in swine) which the animal would per- as they had when space was abundant. . .
form regardless of environmental conditions. All three measures of stress discussed to this point need supportmg ev1dence
before they become valid measures of stress. It must first be established that
Absence of "Normal" Behavior preventing an animal of a particular species from turning around, 0r hav~ng a nor-
Frequently the behavior of an animal in a potentially stressful environment, mal amount of resting time is stressful. This generally means measurmg some
i.e., a cage or in confinement, is compared with that of conspecifics in more change in an animal's physiology indicative of stress.
natural conditions. The prevention of dustbathing, or wing flapping, or simply the
ability to move about in poultry cages is often considered indicative of a stressful TABLE 1 -Effect on reducing number of free stalls on daily free stall
condition. The mere inability of an animal to show behavior that typically occurs utilization per cow.
under a different set of conditions, even though they may be less restrictive, is
not necessarily indicative of a stressful condition.
Free stalls per cow
The Brambell Committee (Brambell, 1965) recommended that an animal
should have freedom to turn around, groom itself, get up, lie down and stretch its
1.0 .83 .67 .50 .33
limbs. There is a problem with blanket recommendations (or legislation) due to
the immense amount of variation across and within a species. The basic behavior-
al repertoire of the species in question, as well as the experience and background
Average time resting
of each individual within the species, must be considered.
in stalls (hr)±SD 14.2±1.5 14.2±1.9 13.2±2.0 10.4±2.0* 6.9±2.0*
For example, a farrowing crate prevents a sow from turning around and
greatly restricts her activity during the four to five weeks she is confined in it.
•Differs from 1.0 free stall/cow (Dunnett's test).
This may be stressful for an animal with a strong drive to travel, such as a wolf. A
pig, however, will normally spend 80% of its time asleep if its environment is sat-
isfactory (Haugse eta/., 1965). Pigs that are accustomed to crates will not show
signs of maladjustment (fighting the crate). When sows are released in the alley Choice Tests
leading to the farrowing crates at our University Swine Center, the sows will sort Why not give the animal a choice between conditions and see which one(s) it
themselves out and willingly enter the crate in which they last farrowed. prefers? This approach does not necessarily discern what is stressful or adverse_ to
Defining normal behavior can also be a problem. Comparing the behavior of an animal's welfare, but merely indicates the animal's preference. The choJCe
domestic animals to that of their wild relatives lacks meaning due to genetic dif- could be due to comfort or familiarity, especially if the second choice is novel.
ferences. Domestication has exerted a strong selective pressure to develop ani- The influence of prior experience in choice tests was shown by Dawkins (1976).
mals that are adapted to domestic conditions. Feral or free ranging domestic She found that when hens were offered a simultaneous choice between a battery
animals would yield more relevant comparisons. If one thinks about this whole cage and an outside run, birds from battery cages initially preferred cages. Hens
approach, it appears analogous to comparing a "primitive" tribe of humans to given continuous access to both did not show evidence of a preference. Operant
apartment dwellers in a major city. One would be making a strictly philosophical conditioning techniques in which the animal indicates its discomfort have the
decision concerning how people should live. It would be necessary, for example, same limitation.

368 /NT 1 STUD ANIM PROB 1(6) 1980 369

/NT 1 STUD ANIM PROB 1(6) 1980
 T.H. Friend-Stress T.H. Friend-Stress Original Article I
Original Article I

tailbiter will eliminate tailbiting in the group. One would expect another tailbiter to examine the health records of both populations to make a meaningful deci-
to evolve if the activity were caused by stressful conditions, but this is not the sion. There is a good chance of discovering that the population with no stress-
case. Also, in a typical, very large confinement farm the author observed related disease also had a much greater mortality rate.
tailbiting in only one pen. If tailbiting is an indication of stress, what can be said
about the other forty-nine identical pens in which it did not occur? Other poten- Change in Behavior Patterns
tial causes and cures of tailbiting are covered in numerous review articles (e.g., This procedure generally establishes fairly quantitative behavioral patterns
Fritschen, 1978). (e.g., time standing, time resting, or number of vocalizations) as ind~ces of str~s.s
The appearance of displacement or vacuum activities does not necessarily or the overtaxing of resources. This approach was used to determme the mml-
prove that an animal is stressed. There may be signs of boredom or frustration mum number of free stalls (resting sites) required by dairy cattle in confinement
(Wood-Gush et a/., 1975; Hediger, 1950), but this may also indicate that the barns before they became overcrowded (Friend eta/., 1977b). Time spent resting
animal is coping with the situation. People will often do something they would per day was determined using time lapse photography as th_e n~mber_ of ~ree
not normally do to relieve boredom or frustration. Following this line of thought, stalls were reduced at seven day intervals. A significant reduct1on m restmg t1me
we could even rationalize that it is the animal which does not show displacement occurred at .50 and .33 stalls per cow (Table 1). However, we could not say that
activities (or, the person who fails to release his or her emotions) that is most the cattle were stressed based on this change in behavioral patterns alone. These
probably stressed. It can also be argued that some vacuum activities are instinc- data indicated only at what density the cattle were no longer able to rest as often
tive behavior patterns (e.g., nest building in swine) which the animal would per- as they had when space was abundant. . .
form regardless of environmental conditions. All three measures of stress discussed to this point need supportmg ev1dence
before they become valid measures of stress. It must first be established that
Absence of "Normal" Behavior preventing an animal of a particular species from turning around, 0r hav~ng a nor-
Frequently the behavior of an animal in a potentially stressful environment, mal amount of resting time is stressful. This generally means measurmg some
i.e., a cage or in confinement, is compared with that of conspecifics in more change in an animal's physiology indicative of stress.
natural conditions. The prevention of dustbathing, or wing flapping, or simply the
ability to move about in poultry cages is often considered indicative of a stressful TABLE 1 -Effect on reducing number of free stalls on daily free stall
condition. The mere inability of an animal to show behavior that typically occurs utilization per cow.
under a different set of conditions, even though they may be less restrictive, is
not necessarily indicative of a stressful condition.
Free stalls per cow
The Brambell Committee (Brambell, 1965) recommended that an animal
should have freedom to turn around, groom itself, get up, lie down and stretch its
1.0 .83 .67 .50 .33
limbs. There is a problem with blanket recommendations (or legislation) due to
the immense amount of variation across and within a species. The basic behavior-
al repertoire of the species in question, as well as the experience and background
Average time resting
of each individual within the species, must be considered.
in stalls (hr)±SD 14.2±1.5 14.2±1.9 13.2±2.0 10.4±2.0* 6.9±2.0*
For example, a farrowing crate prevents a sow from turning around and
greatly restricts her activity during the four to five weeks she is confined in it.
•Differs from 1.0 free stall/cow (Dunnett's test).
This may be stressful for an animal with a strong drive to travel, such as a wolf. A
pig, however, will normally spend 80% of its time asleep if its environment is sat-
isfactory (Haugse eta/., 1965). Pigs that are accustomed to crates will not show
signs of maladjustment (fighting the crate). When sows are released in the alley Choice Tests
leading to the farrowing crates at our University Swine Center, the sows will sort Why not give the animal a choice between conditions and see which one(s) it
themselves out and willingly enter the crate in which they last farrowed. prefers? This approach does not necessarily discern what is stressful or adverse_ to
Defining normal behavior can also be a problem. Comparing the behavior of an animal's welfare, but merely indicates the animal's preference. The choJCe
domestic animals to that of their wild relatives lacks meaning due to genetic dif- could be due to comfort or familiarity, especially if the second choice is novel.
ferences. Domestication has exerted a strong selective pressure to develop ani- The influence of prior experience in choice tests was shown by Dawkins (1976).
mals that are adapted to domestic conditions. Feral or free ranging domestic She found that when hens were offered a simultaneous choice between a battery
animals would yield more relevant comparisons. If one thinks about this whole cage and an outside run, birds from battery cages initially preferred cages. Hens
approach, it appears analogous to comparing a "primitive" tribe of humans to given continuous access to both did not show evidence of a preference. Operant
apartment dwellers in a major city. One would be making a strictly philosophical conditioning techniques in which the animal indicates its discomfort have the
decision concerning how people should live. It would be necessary, for example, same limitation.

368 /NT 1 STUD ANIM PROB 1(6) 1980 369

/NT 1 STUD ANIM PROB 1(6) 1980
 I T.H. Friend-Stress Original Arrick I T.H. Friend-Stress Original Arrick I

Physiological Measures of Stress may have a lagtime of several seconds. Vasoconstriction stimulated by these
pathways causes an increase in blood pressure and a reduction of body surface
Once a stressor is perceived, physiological changes will occur within the temperature (less blood near the surface of the skin). Along with these changes,
body to help the animal resist stress. The degree and type of physiological increased heart rate, respiration rate, and occurrence of ulcers (due to interfer-
measures of stress involve measuring some type of parameter that indicates that ence with normal digestion) have been used to measure stress. These pathways
the body is responding to a stressor. Many parameters have been used; however react rapidly, making them indicators of short-term stress. However, heart rate,
each. has .only limited application. In order to understand the commonly used blood pressure, and respiration are readily influenced by physical activity, which
physrologrcal measures of stress, we must understand the changes in physiology frequently confounds their use as measures of acute stress.
that occur in a stressed animal. Pathway C is concerned primarily with long-term stress. While stimulation of
Figure 1 is a simplified diagram of the major pathways by which an animal this pathway occurs immediately, a 48 hr period of continuous stimulation is re-
ini.tiates physiological responses that are associated with stress. Neutral pathway quired before full symptoms appear. Due to alteration of gonadotrophins, such
A rnfluences the body almost immediately whereas neural endocrine pathway B measures as conception rates, embryonic mortality, and irregular estrus can be
signs of stress. Due to the suppression of growth hormone, growth rates can also
serve as indicators of stress. People involved in livestock production, where
STRESSOR reproduction and growth rates are extremely important, should be cognizant of
this relationship for economic as well as ethical reasons.
J
BRAIN The hypothalamic-pituitary-adrenal cortical axis is vital in enabling an
animal to cope with stress (Selye, 1973). Adrenalectomized animals cannot sur-
vive even mild physical exertion without glucocorticoid therapy. Several minutes
after a stressor commences, stored glucocorticoids are released from the adrenal
in response to initial ACTH secretion. The stores are quickly depleted and
glucocorticoid secretion returns to low levels until the adrenal undergoes
hyperplasia and hypertrophy.
VASOCONSTRICTION This surge can be measured by obtaining blood samples at close intervals
c ADRENAL MEDULA
releases epinephrine after the onset of stress (Smith eta/., 1972). Approximately 48 hrs after initiation
HEART RATE and norelinephrine
increases of the stress, detectable hyperplasia and hypertrophy of the adrenal occurs
(Selye, 1936; Friend eta/., 1977a). Increased glucocorticoid secretion then con-
VASOCONSTRICTION tinues until either the stress is removed, adaptation occurs, or adrenal exhaustion
HYPOTHALAMUS
HEART RATE occurs and the animal dies (Selye, 1973). Change in adrenal weight has been wide-
increases

~ANTERIOR PITUITARY~
1 GASTROINTESTINAL ACTIVITY
decreases
ly used as an indicator of stress exceeding 48 hrs (Christian, 1961; Selye, 1973), but
this requires euthanizing the animal.
Measuring the adrenals' response in the form of glucocorticoid output after
a massive dose of ACTH appears to be a good alternative to euthanizing an
GONADOTROPHINS (FSH, LH)
decreases
J ADRENOCORTICOTROPIC
HORMONE (ACTH)
animal to determine adrenal function. Figure 2 (Friend et a/., 1979) shows a
increases change in adrenal output of glucocorticoids for cows kept at different free stall
GROWTH HORMONE
tI densities. A crowding condition probably occurred (P .01) when the cows (four
J
REPROpUC!IYE ORGANS
decreases
ADRENAL CORTEX
immediate release of stored
per treatment) had access to .50 or .33 stalls. This procedure shows considerable
promise as a means of measuring long-term stress; however, what occurs during
1nh1b1ted glucocorticoids and hyper- adaptation to long-term stressors requires more research. We are beginning to see
plasia and hypertrophy of
adrenal cortex resulting in trends in unpublished data which indicate that adrenal response decreases to
J
PROTEIN SYNTHESIS
a sustained increase of
glucocorticoids
normal levels after one or two weeks of exposure to certain stressors. It is not yet
known if the return to normal indicates that the animal has adapted and the con-
AND GROWTH I dition is no longer stressful or if the hypothalamic-pituitary-adrenal cortical axis
decreases f
GLUCONEOGENESIS is becoming refractory (Selye's "state of exhaustion," 1936). The severity of the
increases
stressor is probably a controlling factor in adaptation.
INFLAMMATION REACTION
inhibited Since the enlarged adrenal releases increased amounts of glucocorticoids,
many researchers have presented plasma concentrations of glucocorticoids as a
FIGURE 1- Major pathways by which an animal responds physiologically to stressors. measure of stress. However, there is a problem in interpreting plasma concentra-

370 /NT 1 STUD ANIM PROB 1(6) 1980 371

/NT 1 STUD ANIM PROB 1(6) 1980
 I T.H. Friend-Stress Original Arrick I T.H. Friend-Stress Original Arrick I

Physiological Measures of Stress may have a lagtime of several seconds. Vasoconstriction stimulated by these
pathways causes an increase in blood pressure and a reduction of body surface
Once a stressor is perceived, physiological changes will occur within the temperature (less blood near the surface of the skin). Along with these changes,
body to help the animal resist stress. The degree and type of physiological increased heart rate, respiration rate, and occurrence of ulcers (due to interfer-
measures of stress involve measuring some type of parameter that indicates that ence with normal digestion) have been used to measure stress. These pathways
the body is responding to a stressor. Many parameters have been used; however react rapidly, making them indicators of short-term stress. However, heart rate,
each. has .only limited application. In order to understand the commonly used blood pressure, and respiration are readily influenced by physical activity, which
physrologrcal measures of stress, we must understand the changes in physiology frequently confounds their use as measures of acute stress.
that occur in a stressed animal. Pathway C is concerned primarily with long-term stress. While stimulation of
Figure 1 is a simplified diagram of the major pathways by which an animal this pathway occurs immediately, a 48 hr period of continuous stimulation is re-
ini.tiates physiological responses that are associated with stress. Neutral pathway quired before full symptoms appear. Due to alteration of gonadotrophins, such
A rnfluences the body almost immediately whereas neural endocrine pathway B measures as conception rates, embryonic mortality, and irregular estrus can be
signs of stress. Due to the suppression of growth hormone, growth rates can also
serve as indicators of stress. People involved in livestock production, where
STRESSOR reproduction and growth rates are extremely important, should be cognizant of
this relationship for economic as well as ethical reasons.
J
BRAIN The hypothalamic-pituitary-adrenal cortical axis is vital in enabling an
animal to cope with stress (Selye, 1973). Adrenalectomized animals cannot sur-
vive even mild physical exertion without glucocorticoid therapy. Several minutes
after a stressor commences, stored glucocorticoids are released from the adrenal
in response to initial ACTH secretion. The stores are quickly depleted and
glucocorticoid secretion returns to low levels until the adrenal undergoes
hyperplasia and hypertrophy.
VASOCONSTRICTION This surge can be measured by obtaining blood samples at close intervals
c ADRENAL MEDULA
releases epinephrine after the onset of stress (Smith eta/., 1972). Approximately 48 hrs after initiation
HEART RATE and norelinephrine
increases of the stress, detectable hyperplasia and hypertrophy of the adrenal occurs
(Selye, 1936; Friend eta/., 1977a). Increased glucocorticoid secretion then con-
VASOCONSTRICTION tinues until either the stress is removed, adaptation occurs, or adrenal exhaustion
HYPOTHALAMUS
HEART RATE occurs and the animal dies (Selye, 1973). Change in adrenal weight has been wide-
increases

~ANTERIOR PITUITARY~
1 GASTROINTESTINAL ACTIVITY
decreases
ly used as an indicator of stress exceeding 48 hrs (Christian, 1961; Selye, 1973), but
this requires euthanizing the animal.
Measuring the adrenals' response in the form of glucocorticoid output after
a massive dose of ACTH appears to be a good alternative to euthanizing an
GONADOTROPHINS (FSH, LH)
decreases
J ADRENOCORTICOTROPIC
HORMONE (ACTH)
animal to determine adrenal function. Figure 2 (Friend et a/., 1979) shows a
increases change in adrenal output of glucocorticoids for cows kept at different free stall
GROWTH HORMONE
tI densities. A crowding condition probably occurred (P .01) when the cows (four
J
REPROpUC!IYE ORGANS
decreases
ADRENAL CORTEX
immediate release of stored
per treatment) had access to .50 or .33 stalls. This procedure shows considerable
promise as a means of measuring long-term stress; however, what occurs during
1nh1b1ted glucocorticoids and hyper- adaptation to long-term stressors requires more research. We are beginning to see
plasia and hypertrophy of
adrenal cortex resulting in trends in unpublished data which indicate that adrenal response decreases to
J
PROTEIN SYNTHESIS
a sustained increase of
glucocorticoids
normal levels after one or two weeks of exposure to certain stressors. It is not yet
known if the return to normal indicates that the animal has adapted and the con-
AND GROWTH I dition is no longer stressful or if the hypothalamic-pituitary-adrenal cortical axis
decreases f
GLUCONEOGENESIS is becoming refractory (Selye's "state of exhaustion," 1936). The severity of the
increases
stressor is probably a controlling factor in adaptation.
INFLAMMATION REACTION
inhibited Since the enlarged adrenal releases increased amounts of glucocorticoids,
many researchers have presented plasma concentrations of glucocorticoids as a
FIGURE 1- Major pathways by which an animal responds physiologically to stressors. measure of stress. However, there is a problem in interpreting plasma concentra-

370 /NT 1 STUD ANIM PROB 1(6) 1980 371

/NT 1 STUD ANIM PROB 1(6) 1980
 T.H. Friend-Stress Original Article I T.H. Friend-Stress Original Article

60 genetic factors can make what would appear idyllic to human beings exceedingly
stressful to another animal and vice versa. People's opinions about various
species are also greatly influenced by experience and education (Kellert, 1980).
'J At present we do not have a simple, all-inclusive technique for determining
::;:
...... 48
"' when an animal is being stressed. The use of behavioral observations alone could
~
tn
easily lead to erroneous conclusions. Techniques based solely on physiological
::
0
changes must be thoroughly evaluated for their applicability to the situation. An
~ 36 integrated approach combining behavioral observations with physiological data
,_.
a:
0
u
is more credible. For example, even though the free stall behavioral data
0
u
:::J
presented earlier quantified a significant change in behavior, it was the concur-
...J

"' 24
<(
rence of the physiological data that gave it credence.
::;:
tn
Physiological parameters, especially hormone determinations, are expensive
<(
...J and time consuming. A practical alternative for many situations would be to use
"-
...J physiological data to establish whether a behavioral observation is indicative of
,_.
<(

stress and then to use that behavior (or change in behavior) as a measure of stress
,_.
0 12
in comparable situations.
There has been very little research directed toward developing an objective
0 method of assessing stress. Most of what is known is drawn from work with other
0 1 2 3 4 objectives. People have become interested in this area only relatively recently,
HOURS AFTER ADMINISTRATION OF ACTH and financial support for such work in farm animals in the United States is still
neg I igible.

FIGURE 2- Adrenal output of glucocorticoids after a massive dose of adrenocortocotropin

5
(1.98 IU ACTH per kg body weight .7 ) in cows under different free stall stocking rates (Friend et
Acknowledgements
a/., 1979).
The author appreciates the assistance of G.R. Dellmeier in the organization
and editing of the manuscript.
tions of any hormone. What is present in the blood is a product of secretion and
clearance. Without knowing turnover rate, blood concentrations of any hormone References
are poor indicators of what is actually occurring. For example, Christison and
Johnson (1972) showed that under conditions of prolonged heat exposure, ad~ Brambell, F.W.R. (1965) Chairman. Report of the Technical Committee to inquire
justments are made in turnover rate of glucocorticoids. into the Welfare of Animals kept under Intensive Livestock Husbandry
The effects of glucocorticoids on the body are very complex and are the ob- Systems. H.M.S.O.
ject of considerable study. Figure 1 lists two effects that have been well Christian, J.J. (1961) Phenomena associated with population density, Proc Nat
documented. G lucocorticoids are known to be gluconeogenic; therefore, re- Acad Sci 47:428-449.
searchers can relate changes in plasma glucose to the occurrence of stress (Pear- Christison, G.l. and Johnson, H.D. (1972) Cortisol turnover in heat-stressed cows,
son and Mellor, 1976). The anti-inflammatory effect of glucocorticoids and the j Anim Sci 35:1005-1010.
resulting change in resistance to standardized disease challenges also serve as Dawkins, M. (1976) Towards an objective measure of assessing welfare in
measures of stress (Gross and Colmano, 1969; Gross, 1972). Animals that have domestic fowl, App Anim Ethol 2:245-254.
been transported long distances often show a high incidence of disease. "Ship- Fritschen, B. (1978) Tailbiting in pigs, Confinement 3(2):11.
ping fever" is considered a major problem in the cattle industry. Fraser, D., Ritchie, J.S.D. and Fraser, A. F. (1975) The term "stress" in a veterinary
context, Brit Vet J 131:653-662.
Discussion Friend, T.H., Polan, C.E., Guazdauskas, F. and Heald, C.W. (1977a) Adrenal glu-
cocorticoid response to exogeneous adrenocorticotropin mediated by den-
The common anthropomorphic approach of assessing an animal's welfare sity and social disruption in lactating cows, J Dairy Sci 60:1958-1963.
by attempting to place oneself in an animal's situation as one perceives it lacks Friend, T.H., Polan, C.E. and McGilliard, M.L. (19776) Free stall and feedbunk re-
credibility. Each species lives in a totally different Umwelt (a term to describe the quirements relative to behavior, production and individual feed intake in
world around a living organism as that creature experiences it). Differences in dairy cows, j Dairy Sci 60:108-116.
perception (sight, taste, smell, touch, hearing), sense of time, experience and Friend, T.H. Guazdauskas, F. and Polan, C.E. (1979) Change in adrenal response

372 /NT 1 STUD ANIM PROB 1(6) 1980 /NT 1 STUD AN/M PROB 1(6) 1980 373
 T.H. Friend-Stress Original Article I T.H. Friend-Stress Original Article

60 genetic factors can make what would appear idyllic to human beings exceedingly
stressful to another animal and vice versa. People's opinions about various
species are also greatly influenced by experience and education (Kellert, 1980).
'J At present we do not have a simple, all-inclusive technique for determining
::;:
...... 48
"' when an animal is being stressed. The use of behavioral observations alone could
~
tn
easily lead to erroneous conclusions. Techniques based solely on physiological
::
0
changes must be thoroughly evaluated for their applicability to the situation. An
~ 36 integrated approach combining behavioral observations with physiological data
,_.
a:
0
u
is more credible. For example, even though the free stall behavioral data
0
u
:::J
presented earlier quantified a significant change in behavior, it was the concur-
...J

"' 24
<(
rence of the physiological data that gave it credence.
::;:
tn
Physiological parameters, especially hormone determinations, are expensive
<(
...J and time consuming. A practical alternative for many situations would be to use
"-
...J physiological data to establish whether a behavioral observation is indicative of
,_.
<(

stress and then to use that behavior (or change in behavior) as a measure of stress
,_.
0 12
in comparable situations.
There has been very little research directed toward developing an objective
0 method of assessing stress. Most of what is known is drawn from work with other
0 1 2 3 4 objectives. People have become interested in this area only relatively recently,
HOURS AFTER ADMINISTRATION OF ACTH and financial support for such work in farm animals in the United States is still
neg I igible.

FIGURE 2- Adrenal output of glucocorticoids after a massive dose of adrenocortocotropin

5
(1.98 IU ACTH per kg body weight .7 ) in cows under different free stall stocking rates (Friend et
Acknowledgements
a/., 1979).
The author appreciates the assistance of G.R. Dellmeier in the organization
and editing of the manuscript.
tions of any hormone. What is present in the blood is a product of secretion and
clearance. Without knowing turnover rate, blood concentrations of any hormone References
are poor indicators of what is actually occurring. For example, Christison and
Johnson (1972) showed that under conditions of prolonged heat exposure, ad~ Brambell, F.W.R. (1965) Chairman. Report of the Technical Committee to inquire
justments are made in turnover rate of glucocorticoids. into the Welfare of Animals kept under Intensive Livestock Husbandry
The effects of glucocorticoids on the body are very complex and are the ob- Systems. H.M.S.O.
ject of considerable study. Figure 1 lists two effects that have been well Christian, J.J. (1961) Phenomena associated with population density, Proc Nat
documented. G lucocorticoids are known to be gluconeogenic; therefore, re- Acad Sci 47:428-449.
searchers can relate changes in plasma glucose to the occurrence of stress (Pear- Christison, G.l. and Johnson, H.D. (1972) Cortisol turnover in heat-stressed cows,
son and Mellor, 1976). The anti-inflammatory effect of glucocorticoids and the j Anim Sci 35:1005-1010.
resulting change in resistance to standardized disease challenges also serve as Dawkins, M. (1976) Towards an objective measure of assessing welfare in
measures of stress (Gross and Colmano, 1969; Gross, 1972). Animals that have domestic fowl, App Anim Ethol 2:245-254.
been transported long distances often show a high incidence of disease. "Ship- Fritschen, B. (1978) Tailbiting in pigs, Confinement 3(2):11.
ping fever" is considered a major problem in the cattle industry. Fraser, D., Ritchie, J.S.D. and Fraser, A. F. (1975) The term "stress" in a veterinary
context, Brit Vet J 131:653-662.
Discussion Friend, T.H., Polan, C.E., Guazdauskas, F. and Heald, C.W. (1977a) Adrenal glu-
cocorticoid response to exogeneous adrenocorticotropin mediated by den-
The common anthropomorphic approach of assessing an animal's welfare sity and social disruption in lactating cows, J Dairy Sci 60:1958-1963.
by attempting to place oneself in an animal's situation as one perceives it lacks Friend, T.H., Polan, C.E. and McGilliard, M.L. (19776) Free stall and feedbunk re-
credibility. Each species lives in a totally different Umwelt (a term to describe the quirements relative to behavior, production and individual feed intake in
world around a living organism as that creature experiences it). Differences in dairy cows, j Dairy Sci 60:108-116.
perception (sight, taste, smell, touch, hearing), sense of time, experience and Friend, T.H. Guazdauskas, F. and Polan, C.E. (1979) Change in adrenal response

372 /NT 1 STUD ANIM PROB 1(6) 1980 /NT 1 STUD AN/M PROB 1(6) 1980 373
I T.H. Friend-StreS> Original Article I REVIEW ARTICLE
from free stall competition, j Dairy Sci 62:768-771.
Goldstein, A. (1976) Opioid peptides (endomorphins) in pituitary and brain,
Science 193:1081-1086.
Problems With Kosher
Gross, W.B. (1972) Effect of social stress on occurrence of Marek's disease in Slaughter
chickens, Am J Vet Res 33:2225-2279.
Gross, W.B. and Colmano, G. (1969) The effect of social isolation on resistance to
some infectious diseases, Poult Sci 48:514-520. Temple Grandin*
Haugse, C.N., Dinussen, W.E., Erickson, D.O., johnson, J.N. and Buchanan, M.L.
(1965) A day in the life of a pig, N Oak Fm Res 23:18-23.
Hediger, H. (1950) Wild Animals in Captivity, Butterworths Scientific Publications,
London, UK, pp 207.
Kellert, S.R. (1980) American attitudes toward and knowledge of animals: an up-
Abstract
date. lnt j Stud Anim Prob 7:87-119. Ritual slaughter to produce kosher meat is rooted in the teachings and
Moberg, G.P. (1976) Effects of environment and management stress on reproduc- writings of the Talmud. However, the preslaughter handling features of modern
tion in the dairy cow, j Dairy Sci 59:1618-1624. systems, particularly the shackling and hoisting of large steers, contravene the
Pearson, R.A. and Mellor, D.J. (1976) Some behavioral and physiological changes basic message of humaneness included in the teachings. The throat-cutting of a
in pregnant goats and sheep during adaptation to laboratory conditions, live, conscious animal is relatively pain-free, provided that certain precautions are
Res Vet Sci 20:215-217. followed, but U.S. kosher plants need to install newly developed conveyor-
Peters, M. (1977) Hypertension and the nature of stress, Science 198:80. restrainer systems to eliminate the abuses of shackling and hoisting. Conveyor-
Selye, H. (1936) A syndrome produced by diverse nocuous agents, Nature 138:32. restrainer systems for large and small animals are discussed.
Selye, H. (1973) The evolution of the stress concept, Am Scientist 61:692-699.
Smith, V.G., Convey, E.M. and Edgerton, C.A. (1972) Bovine serum corticoid
response to milking and exteroceptive stimuli, j Dairy Sci 55:1170-1173. The Religious Ritual
Van Putten, G. (1969) An investigation into tail-biting among fattening pigs, Brit
Vet J 725:511-516. In order for a piece of meat to be kosher, it must be slaughtered and process-
Wood-Gush, D.G.M., Duncan, I.J.H. and Fraser, D. (1975) Social stress and welfare ed according to ritual methods specified in the Talmud. These methods derive
problems in agricultural animals. In The Behavior of Domestic Animals, 3rd from explicit commands contained in the Torah on the types of animals that the
ed. E.S.E. Hafez, ed, Williams and Wilkins Co, Baltimore, MD, pp 182-200. children of Israel may eat and how these animals should be prepared for
consumption. 'Shechitah', the act of killing for food, must be conducted by a
learned, pious Jew, the 'shochet', who is trained in the slaughter ritual.
The shochet slaughters the fully conscious animal with a razor-sharp knife,
which must be twice the width of the throat of the animal to be slaughtered. He
uses a single, smooth deliberate motion, severing the carotids and the trachea.
After each cut, the shochet checks the knife for nicks or imperfections. If the
blade has a nick, then the animal is declared to be 'tref' or not kosher and the
meat is sold on the regular markeL Shochet Rabbi Garb (1977, personal com-
munication) contends that, if the cut is made correctly and the knife has no nicks,
the animal feels little or no pain.

The five rules of kosher slaughter are as follows:

1 . 'Shechiyah' (Delay)- A delay or hesitation of even a moment makes the

*Ms. Grandin is an independent livestock handling consultant and owner of Grandin Livestock Han-
dling Systems, 617 E. Apache Blvd., Tempe, AZ 85281. This is the sixth article in a series of six appear-
ing in the /nt I Stud Anim Prob. Previously published: Livestock Behavior as Related to Handling Facili-
ty Design; Bruises and Carcass Damage; Designs and Specifications for Livestock Handling Equipment
in Slaughter Plants; Mechanical, Electrical and Anesthetic Stunning Methods for Livestock; The Effect
of Stress on Livestock and Meat Quality Prior to and During Slaughter.

374 /NT I STUD ANJM PROB 1(6) 1980 /NT I STUD ANIM PROB 1(6) 1980 375