LAWRIE-DARK-CUTTING BEEF 721

in the milling of different grades of flour. This is seen in the data for the third series of Manitoba
flours shown in Table I11 which were milled to conform to accepted standards for the three
extraction rates. As the rate increased from 70 to 80 to 85% the content of vitamin B, changed
from 21 to 63 to 84% of the wholemeal value but the content of vitamin B, changed only from
16 to 26 to 43% of the wholemeal value. The pantothenic acid content, as previously r e p ~ r t e d , ~
changed from 49 to 58 to 76% of the wholemeal value.
Acknowledgments
The authors wish to thank Mr. H. N. Ridyard for the vitamin B, determinations and Miss
N. Davies for technical assistance.

Research Association of British Flour-Millers

Cereals Research Station
St. Albans
Received 20 February, 1958

References
1 ' Report of the Panel on Composition and Nutritive Parrish, W. P., Loy, H. W., & K h e , 0. L., J , Ass.
Value of Flour ', 1956 (London : H.M.S.O.) off. agn'c. Chem. Wash., 1955.38,'506
2 Hinton, J. J. C., Peers, F. G.. & Shaw, B., Naftrve, OAss. of Vitamin Chemists, Inc., Methods of
Land.. 1953, 172, 993 Vitamin Assay', 1951,2nd Edn., p. 221 (New
3 Clegg, K. M., J . Sci. Fd Agvic., 1958, 9, 366 York : Interscience Publishers, Inc.)
4 Atkin, L., Schultz, A. S., Williams, W. L., & Frey. Heathcote, J. G., Hinton, J. J. C., & Shaw, B.,
C. N., Industr. Engng Chem. (Anal.Edn),1943.15, Proc. YO^. SOC.,[B],1952, 139, 276
14' lo Rabinowitz, J. C.. & Snell, E. E.. Industr. Engng
40 Kodicek, E.. & Pepper, C. R., J. gen. Microbiol., Chem. (Anal. Edn), 1947, 19, 277
1948, 2, 306 l1 Parrish, W. P., Loy, H. W., & Kline, 0. L., J. Ass.
5 Chiao, J. S., & Peterson, W. H., Arch. Biochem. off. agric. Chem. Wash., 1956,39, 157
Biopizys., 1956, 64, 115 l a Hopkins, R. H., & Pennington, R. J., Biochem. J . .
Hinton. J. J. C., PYOC.
rcy. Soc., [B],1947.134, 418 1947. 41, 1 1 0

PHYSIOLOGICAL STRESS IN RELATION TO DARK-CUTTING

BEEF
B y R. A. LAWRIE
Introduction
Breed, sex, age and plane of nutrition are acknowledged basic determinants of meat quality
but it is also influenced by numerous secondary factors. Pathological conditions in the animal
can be important in this context, and may be of such a nature as not only to impair quality,
but even to make consumption of the carcass dangerous. Competent veterinary control a t
the abattoir ensures that grossly pathological carcasses do not reach the public. Nevertheless,
animals are subject to certain other maladies, of physiological rather than pathological nature,
which can give rise to meat of inferior quality without justifying condemnation of the carcasses.
' Dark-cutting ' beef is attributable to a disorder of this kind. Although the condition has been
long recognized-recorded observations date from a t least as far back as 1774-it has not been
possible to apprehend its nature until relatively recent times.
The significance of high ultimate pH
When muscles from beef carcasses are cut, the surfaces thus exposed have initially the
purplish colour of the reduced muscle pigment, myoglobin. As the myoglobin becomes oxy-
genated on exposure, the cut surfaces gradually assume the bright red appearance of fresh
meat. On the other hand, the surfaces of muscle from dark-cutting beef, as the name implies,
remain purplish after even prolonged exposure to the atmosphere.
J. Sci. Food Agric., 9, November, 1958
722 LAWRIE-DARK-CUTTING BEEF
In a survey published in 1944,Hall et al. indicated that dark-cutting beef has an unusually
rapid rate of consumption of oxygen, practically no glycogen (immediately after slaughter)
and an ultimate pH* considerably higher than 5.4-5.6 (the value typical of normal beef muscle).
These factors could cause dark colour on cut meat surfaces in several ways.
The depth of the layer of oxymyoglobin on the surface of meat is determined by the rate of
oxygen utilization of certain tissue enzyme systems and by the rate of diffusion of oxygen inwards
from the atmosphere ; these two processes are themselves determined by a variety of factorsa
If the rate of oxygen utilization increases while the rate of diffusion inwards remains constant,
the depth of the oxymyoglobin layer on the surface will diminish and may become so small
that the purplish colour of reduced myoglobin predominates. Cytochrome oxidase is probably
the principal enzyme responsible for oxygen uptake in biological tissues. Its activity (as
prepared from horse heart) is dependent on pH (Table I).5 If the ultimate pH of the meat is
31

abnormally high, as it is in dark-cutting beef, the rate of oxygen uptake by cytochrome oxidase
would rise considerably and, for the above reasons, could contribute to the dark colour.
The depth of oxymyoglobin on the surface of dark-cutting beef is also likely to be diminished
because of the unusual structural features of the latter. This is another consequence of high
pH. The isoelectric point of the principal muscle protein, actomyosin, is at pH 5-6.6 A t
higher pH values the protein is associated with more water and there is a consequent diminution
in the fluid phase of the muscle. The muscle as a whole has, thus, a more ' close ' structure.'
This feature must lower the rate of inward diffusion of oxygen to the intracellular proteins and
thus impose an additional limit on oxymyoglobin formation a t the surface. Meat of high pH
is also likely to appear dark, as Bate-Smiths has pointed out, because its surface will scatter
incident light less than that of normal beef in which the constituent muscle fibres will have
shrunk apart, whereas in the other they will be swollen and tightly packed together.
I t may be mentioned, incidentally, that unpublished observations of Hall & Macintosh in
1932 indicated that no unusual pigment is present in dark-cutting beef ; nor is there necessarily
a greater than normal amount of residual blood in the c a r c a s ~ .Moreover,
~ myoglobin concentra-
tion in dark-cutting beef is within the normal range (unpublished observations : Lawrie, 1957).
These findings have been recently confirmed with dark-cutting beef produced by several
methods.1°
It will be apparent, therefore, that a high ultimate pH is of primary importance in accounting
for the colour (and consistency) of dark-cutting beef. In order to understand the aetiology of
the condition further it is thus necessary to consider the factors which control the quantity of
glycogen in muscles a t death.

Table I
Activity of cytochrome onidase at diflerent PH values
PH Activity
(pl. O,/mg. d r y fat-free
enzyme solids/h. at 39')
5'9 750
6.4 900
6.8 1650
7'3 1900

Various factors affecting ultimate pH in muscle

Fasting, exhausting exercise and training are among factors known to determine the quantity
of glycogen available at death for the formation of lactic acid, and hence the ultimate pH in a
specific muscle. Between different muscles there are characteristic differences in glycogen
content;l0*l1 these are probably functional, however, and will not be discussed here. Struggling
at death can cause the breakdown of considerable quantities of glycogen, but since the circula-
tion stops at about this time, the lactic acid formed is not appreciably lost from the muscle and
thus still contributes to the ultimate pH. This is supported by experiments with rabbits12
and steers's in which struggling at death was prevented by pre-slaughter administration of
* Le., th e pH when post-mortem lactic acid production has ceased.
J. Sci. Food Agric., 9, November, 1958
 LAWRIE-DARK-CUTTING BEEF 723

the muscle-relaxant drug myanesin. The conservation of glycogen achieved by this means
did not affect the ultimate pH.
The influence of fasting in depleting the glycogen reserves of muscle has been known since
the work of Bernard in 1877.14 Recognition of the importance of this factor in relation to the
meat from domestic species is much more recent. In this context, Callow'* l5 indicated that
inadequate feeding could lower reserves of glycogen in the muscles of pigs and Bate-Smith &
Bendall'* showed that fasting for only 48-72 hours lowered the glycogen of rabbit psoas muscles
sufficiently to raise the ultimate pH from the normal 5.9 to 6.5. In contrast, when steers are
fasted a t normal ambient temperatures for periods even up to 28 days, the ultimate pH is
unaffected. Although there is some suggestion that muscle glycogen levels have fallen some-
what after 7 days' fasting, there appears to be little further diminution with continued inanition
(Table II).13
Table I1
Glycogen concentrations* and ultimate p H i n psoas and 1. dorsi muscles of steers after fasting
_-
L. dovsi Psoas
Glycogen, mg.-% Ultimate pH Glycogen, mg.-% Ultimate pH
Controls 1126f 98 5'50 zk 0.04 1082 f 109 5'53 f 0.07
Fasted 7 days 753 5'46 696 5.48
Fasted 14 days 597 5'73 717 5'55
Fasted 28 days 768 5'44 809 5'47
* Samples of muscle have varying initial pH, as more or less glycogen has been broken down at time of
receipt. A reasonable approximation to the i n uiuo resting value for muscle glycogen can be made by calcu-
lating the glycogen concentration of all samples on the basis of a standard initial pH. pH 7.2 may be regarded
as suitable for this purpose.

The importance of exhausting exercise as a factor in depleting glycogen reserves has also
been recognized for a considerable time. Hoet & Marks" showed that convulsions induced in
rabbits by insulin could completely eliminate muscle glycogen. Although it is considerably
more difficult to affect ruminants in this way," a high ultimate pH, with consequent dark-
cutting character, can be brought about in beef animals, with a very high dose rate of lS
Insulin tetany is not a naturally occurring condition, but intense muscular activity, induced
by more natural means, can lower the glycogen concentration of muscle to varying extents.
Bate-SmithlB* demonstrated that glycogen reserves in the hind limb muscles of rats could
be severely depleted, with a consequent rise in the ultimate pH, if these animals were subjected
to exhausting exercise pre-slaughter. He pointed out, however, that it was the severity of the
exercise rather than its duration which determined the degree of depletion. In fact, if rats
were ' trained ' by being subjected to gradually increasing bouts of exercise, muscle glycogen
levels could actually be built up. Mitchell & Hamiltonz0 reported similar findings for cattle.
The glycogen of pig psoas muscle is particularly susceptible to depletion by mild activity immedi-
ately pre-slaughter;'$ l5 a small but significant elevation of ultimate pH followed a walk of only
a quarter of a mile. It is most difficult, however, to deplete the muscle glycogen of steers by
enforced exercise, Hall et aE.1 quote investigations carried out in 1935 a t the Illinois and New
York Agricultural Experiment Stations in which exercised steers, although hot and exhausted,
nevertheless failed to yield dark-cutting flesh. The normality of the glycogen reserves in steers
which had been exercised to exhaustion immediately prior to slaughter, was confirmed by
Howard & Lawrie.18 The mean glycogen concentration in the longissirnus dorsi muscles was
1187mg.-%, the mean ultimate pH being 5-55. This may be compared with 1126mg.-% and
an ultimate pH of 5-50 for the 1. dorsi of controls (Table 11). The mean ultimate pH of the
psoas muscles from such animals was rather higher than normal (5.80) but there was little effect
elsewhere in the musculature.
The possibility that the stresses of fasting and enforced exercise, although relatively
ineffective separately, might significantly raise the ultimate pH if combined, was investigated
by Howard & Lawrie.1Ov l8 Steers which had been without food for some days, during a pro-
longed train journey from Western Queensland to Brisbane, were used. Data from two of the
muscles studied are summarized in Table 111. While the concentration of glycogen in the
muscles of steers exercised immediately after detraining had been lowered sufficiently to raise
J. Sci. Food Agric., 9, November, 1958
724 LA WRIE-DARK-CUTTIKG BEEF
Table I11
Glycogen concentrations and ultimate +H i n psoas and 1. dorsi muscles of steers after enforced exercise and fasting
L . dorsi- ~ _ Psoas_- _
Glycogen, Ultimate Glycogen, Ultimate
mg.-% pH mg.--% pH
Controls* (fed and rested 14 days after detraining) 957 5'49 1017 5.48
Exercised 13 h.* (after detraining) 628 5'72 352 6.15
Exercised I ) h.t (after detraining and 14 days fasting) 1028 5'55 508 5'55
Exercised 13h. (after 14 days fasting and neopyrithiamin
injection) 480 6-02 207 6-62
* Mean of 3 observations. t Mean of 2 observations.
the ultimate pH, those from steers subjected in addition to a further 14days' fasting contained
sufficient glycogen to reach a normal ultimate pH. This suggested that the disturbances of
prolonged train travel might have been a significant factor in depleting glycogen reserves,
possibly through an effect on the nervous system of the cattle ; and indicated that the com-
bination of fasting with enforced exercise was not effective when detrained steers, although
fasted, were allowed to rest for some days before exercise and slaughter. I t is true that if the
dependence of steer musculature on carbohydrate as a source of energy is increased-this may
be done by pre-slaughter administration of neopyrithiamin which limits energy production from
fat-the combination of fasting with exercise can effectively lower the glycogen reserves of
steer muscles, thus raising their ultimate pH (Table 111). In this context, there are numerous
indications in the literature that ruminants have an unusually high capacity to gain energy
by oxidation of fatty metabolites. Carbohydrate would thus tend to be spared under the
conditions of stress which would deplete glycogen reserves in non-ruminant species. It may
be that glycogen is not physiologically homogeneous in the muscles of steers and that, according
to degree of stress, first the most ' accessible ' glycogen store, next fatty acids, and finally the
remaining glycogen would be utilized.. Nevertheless, it is evident that in natural circum-
stances neither fasting, enforced exercise nor a combination of both bring about a marked
raising of ultimate pH and these factors must, therefore, be largely discounted in explaining
the occurrence of dark-cutting beef.
The influence of ' excitability * on muscle-glycogen concentration
The existence of some other influence controlling muscle-glycogen concentration was
strongly suggested by the finding that certain steers which had been well fed and rested and
would, therefore, have been expected to have ample glycogen in their muscles yielded meat of
high pH.13 Examination of the data from control steers of a reputedly uniform group of 30
showed that there were progressive changes with sequence of slaughter over a period of some
weeks : muscle-glycogen concentrations tended to diminish and ultimate pH values to rise.
In handling the later steers of this group it had been noticed that they were ' excitable ' and
difficult to control. It was possible to identify these animals by their numbers and to eliminate
the holding period itself as a contributory factor, since the steers noticed subsequently to be
excitable were those which had proved relatively intractable when holding commenced. This
excitability was especially marked in one control steer, which exhibited tremor even when a t
rest ; and on sampling at I hour post mortem the psoas and 1. dorsa muscles were found to have
only about 40 mg.-% glycogen and the ultimate pH was in the region of 6.0 to 6.3. It appeared
that certain steers were of an excitable temperament and that in these animals some short-
range muscular tension, not manifested by external movement, reduced the glycogen reserves
in their muscles to a chronically low level.
Conceivably, if tremor or shivering were artificially induced in normal (non-excitable)
steers, an elevation of ultimate pH might result. Two steers were accordingly given massive
doses of tuberculin to produce protein shock and, thereby, tremor.10 One of these failed to
react, but in the muscles of the other, glycogen reserves were considerably depleted, the con-
centrations being only 355 and 395 mg.-%, and the ultimate pH 6.28 and 6.45, in psoas and
1. dorsi respectively. This effect was brought about without visible movement other than
tremor and slight staggering. It is relevant to note that the incidence of dark-cutting beef in
J. Sci. Food Agric., 9, November, 1958
 LAWRIE-DARK-CUTTING BEEF 725

U.S.A. is more marked in the winter months, and that the condition occurs in steers which are
fasted for as little as two days when simultaneously exposed to cold wet c0nditions.l In
Queensland (the location of the steers studied by Howard & Lawrie) ambient temperatures are
such that animals would require to expend relatively little energy in maintaining body tem-
perature and actual shivering would be rare.
Attempts were made to elucidate further the nature of excitability in steers by pre-slaughter
injections, into naturally docile animals, of thyroxine or adrenaline.13 Although there was
evidence that muscle glycogen reserves had been somewhat lowered, and the mean ultimate
pH of the musculature was raised, the effect was not statistically significant. Recently, how-
ever, Hedrick et a1.,21using a considerably larger dose of adrenaline, were able to cause a dark-
cutting condition in beef. It is particularly significant that the injection of adrenaline, to be
effective, had to be administered about 24 hours before slaughter, and that if 75 mg./xoo lb.
live weight of 17-hydroxycorticosterone (hydrocortisone) were given 24 hours before the
adrenaline, the latter no longer caused the dark-cutting condition. These findings suggest
that a large dose of adrenaline gradually exhausts a compensatory mechanism associated with
the production of hormones from the adrenal cortex. Further understanding of the excitability
phenomenon and of its relation to the other factors controlling the carbohydrate reserve of
steer musculature involves a consideration of the general adaptation syndrome.22
Some degree of physiological stress occurs in all animals when they are subjected to con-
ditions more extreme than those to which they are accustomed. Until fairly recently there
was no reason to suppose that the diverse morphological, functional and biochemical phenomena
associated with stress were related. In 1936, however, Selye noted that animals exposed to a
variety of stress-producing factors-emotional excitement, cold, fatigue, anoxia, inanition, etc.
-reacted by a discharge of hormones from the adrenal cortex-irrespective of the nature of
the stress.2s After the initial reaction, which is characterized by release of adrenaline from
the adrenal medulla, maxima in the production of 17-hydroxycorticosterone and xxdeoxy-
corticosterone occur in that order. These substances elicit a large variety of typical responses
in the organism-collectively known as the general adaptation syndrome-of which the
behaviour of muscle and liver glycogen, and of potassium, is characteristic. Adrenaline causes
hyperkalaemia and breakdown of liver and-probably indirectly-muscle glycogen to glucose
and lactic acid respectively ; it also stimulates the anterior pituitary to produce adrenocorti-
cotrophic hormone (ACTH). This hormone, in turn, stimulates the adrenal cortex to produce
17-hydrouycorticosterone, which tends to restore the glycogen balance by gluconeogenesis from
protein, and 11-deoxycorticosterone, which tends to restore normal potassium balance.22 The
evidence of the last few years has indicated that the hypothalamus plays an essential r6le in these
events. HarrisZ4foundthat stimulation of the hypothalamus caused secretion of ACTH by the pit-
uitary, while direct stimulation of the pituitary gland itself was ineffective in this respect. The
hypothalamus is also responsible for direct stimulation of the adrenal medulla.26 It is suspected
that the hypothalamus exerts its influence a t least partly through the production of serotonin.
Its action can be inhibited by the newly discovered drugs reserpine, Frenquel and chlorpromazine,
which exert a general tranquillizing effect on the organismae and have, in fact, been used for
this purpose with striking success in human psychiatry.26
Diseases of adaptation
It may be appreciated that imbalance at various points in the hypothalamus-pituitary-
adrenal complex of reactions (the general adaptation syndrome) could cause so-called diseases
of adaptation.27- 28 The work of Ludvigsen,20 in particular his detailed study of ‘ muscle
degeneration disease ’ in pigs,* has thrown considerable light on the aetiology of such diseases
in domestic stock. Macroscopically, muscle degeneration disease is characterized by a pale
grey colour and moist, exudative structure in the muscles. The changes may affect all or only
some of the musculature. Biochemically, there is a very swift fall in pH in the immediate
post-mortem period, values of 5.3 being reached 40 minutes after death, compared with a pH of
about 6.4at this time in normal muscle, and there is a high blood potassium-of the order of
* ‘ Muscle degeneration disease ’ seems t o correspond with ‘ Acuter Herztod des Schweines in Germany
and ‘Mulberry Heart Disease’ in the United Kingdom.
J. Sci. Food Agric., 9, November, 1958 zz
726 LAWRIE-DARK-CUTTING BEEF
90 mg.-yo in comparison with 1 6 2 0 mg.-yo in normal pigs. The condition apparently arises
when forced and unaccustomed muscular activity occurs during transport-with resultant
excitability. Ludvigsenz0showed that methylthiouracil could induce the condition and that it
could be counteracted by feeding iodocasein. He suggested that by breeding the Danish
Landrace pig (in which the disease is apparently prevalent) for a high weight increment per unit
food intake, an automatic selection of animals with hypothyroid function had been made. He
was able to show that the pituitary of pigs suffering from muscle degeneration disease had an
enhanced content of growth-stimulating hormone and lowered contents of ACTH and of thyroid-
stimulating hormone. (The possibility that, in some circumstances, production of one hormone
in the pituitary could be accomplished at the expense of another was first raised by Selye ;30
Nowell & Chester-Jones3' have recently found a reciprocity between the production of ACTH
and that of gonadotrophins in the pituitary of rats.) When subjected to stress, animals having
this particular hormone imbalance would be relatively unable to counteract the initial hyper-
kalaemia and glycogen breakdown arising during the alarm phase of reaction to stress : the loss
of potassium from muscles would account for the structural changes observed in them, for the
loss of pigmentation and for the high rate of glycogen breakdown. Subsequently, Ludvigsenaz
has shown that the administration of hormones from the adrenal cortex to pigs suffering from
muscle degeneration disease prevents the exhaustion and trauma which otherwise arise when
they are exercised. Examination of the hypothalmus in such pigs indicated a state of severe
hyperactivity which, Ludvigsen concludes, represents an ineffectual attempt by this organ to
stimulate the pituitary to produce ACTH (in which it is deficient) and, thereby, production of
restitutive hormones by the adrenal cortex. Muscle degeneration disease can be regarded as
a typical disease of adaptation in which the superficial manifestations correspond especially to
inadequate production of mineralocorticoids (e.g. I I-deoxycorticosterone), the ionic imbalance
implying an acute condition.
Although exudation of fluid has been observed in beef muscle a t relatively high pH,lO
this is extremely rare and the above considerations suggest that ' dark-cutting beef ' may arise
from a more chronic condition of the adaptive type in which the production of glucocorticoids
(e.g. 17-hydroxycorticosterone) is particularly inadequate. This inadequacy could, presumably,
originate from prolonged over-stimulation by a hypersensitive hypothalamus of the pituitary
and the adrenal cortex, and might be aggravated by genetically determined deficiencies in this
gland. In excitable cattle an enhanced susceptibility to stimuli and impaired restitutive capacity
could thus lead to a chronically low equilibrium level of glycogen in the musculature and to
high ultimate pH, with attendant dark-cutting characteristics.
Since, as mentioned above, the ruminant appears to have a considerable capacity to gain
energy by oxidation of fatty metabolites, it must be inferred that the conditions arising initially
in steers exposed to stress stimuli, and chronically in excitable steers, are such that the metabolism
of fatty acids cannot entirely prevent the utilization of carbohydrate. This is supported by
observation. According to Selye22increases in lactic acid and in ketone bodies in the blood are
among characteristic responses in initial alarm phase of the reaction to stress and also in
the subsequent phase of exhaustion. The production of lactic acid indicates that carbohydrate
breakdown is occumng under a degree of anaerobiosis and the presence of ketone bodies reflects
a relative inability to oxidize fatty acids : the latter requires aerobic conditions.
It would be presumed, correspondingly, that the resistance of normal steers to stress implies
a relative lack of reactivity in the hypothalamus. This could explain, for instance, the con-
siderable resistance of the steer to insulin tetany, since impairment of hypothalamic pathways
has been shown in the rat to bring about insensitivity to insulin.33
Such drugs as reserpine might, therefore, be even more effective in preventing dark-cutting
beef, through their suppression of hypothalamic stimuli, than therapy by the administration
of hormones from the adrenal cortex. There is also some indication that prophylaxis by sex
hormones might be feasible since, according to Nowell & Chester-Jones,31 androgens facilitate
ACTH action on the adrenal cortex and oestrogens inhibit it. Conceivably, powers of glycogen
restitution are greater in males than in females. (In this context, it may be mentioned that the
mean ultimate pH of the musculature from cows is significantly higher than that from steers :
Lawrie, unpublished observations, 1953).
J. Sci. Food Agric., 9, November, 1958
 LAWRIE-DARK-CUTTING BEEF 727

Although inadequate stimulation of the adrenal cortex by ACTH may be a feature common
to these diseases of adaptation, it has been pointed out that, more specifically, the symptoms
may be rationalized on the basis of deficiencies in 11-deoxycorticosterone and 17-hydroxy-
corticosterone in pigs and cattle respectively. . This indicates that there may be differences a t
the adrenal gland of these species. Such differences at the histochemical level have in fact
been demonstrated by Ni~ander.~'
Conclusion
Irrespective of the mechanism of reaction to stress, the reason for varying sensitivity in
steers remains unknown. Whereas some of the circumstances predisposing to muscle degenera-
tion disease in pigs can be rationally explained on the basis of selection policy, no similar evidence
has so far been adduced in the case of dark-cutting beef to establish the hereditary or environ-
mental factors which might be involved. It is nevertheless suggestive that, according to
Kidwell,ss there was at one time an apparent genetical link between dark-cutting character and
muscular hypertrophy in the original (then relatively unselected) European stock. This would
accord with the inverse relationship between the production of growth-regulating and adreno-
corticotrophic hormones by the pituitary, to which reference has already been made. It thus
seems reasonable to conclude that effective prevention of dark-cutting beef and of other mani-
festations of adaptive disease in domestic livestock will depend on the concomitant progress
of genetics and neuroendocrinology.
Acknowledgment
This paper has been prepared as part of the programme of the Food Inkestigation Organiza-
tion of the Department of Scientific and Industrial Research.
(Crown Copyright Reserved)
Lower Temperature Research Station
Downing St.
Cambridge
Received a8 February, 1958

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J. Sci. Food Agric., 9, November, 1958