Sistema nervoso das aves

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10 D . POLIN 
and their role in neural transmission— 
and the list goes on. Symposia have their 
role in producing renewed interest in the 
subjects under review, and the scientists 
have performed their role well, as you will 
read. 
I wish to thank each of the participants 
for their energetic support in making the 
symposium a success. I shall look forward 
to future physiology symposia at Poultry 
Science Meetings. 
From Science shall come Truth, 
From Truth shall come Science, 
From Faith shall come both. 
2. THE NERVOUS SYSTEM OF BIRDS: A REVIEW 
A. VAN TlENHOVEN 
Department of Poultry Science, Cornell University, Ithaca, N. Y. 14850 
Students of the avian nervous system 
are faced with somewhat of a dilemma be­
cause so little information is available on 
the anatomy and physiology of the avian 
system when it is compared with the 
available information for mammalian 
systems. Two strategies of attack are 
obvious. In the one strategy one basically 
"starts from scratch" and tries to obtain 
a bird's eye view of the problems or one 
may start with the knowledge of the 
mammalian system and one compares the 
similarities and differences in function be­
tween these classes and attempts to gen­
eralize from the data obtained. 
In this review an attempt is made to use 
the latter strategy. 
ANATOMY 
Some of the major anatomical differ­
ences between many of the avian brains 
and many of the mammalian brains are: 
1. In birds a cortex defined according to 
Cobb (1960) as "a peripherally 
placed coating (pallium) of cells 
arranged in layers" is absent. Haefel-
finger (1957) and Stingelin (1958) 
have, however, considered the hy-
perstriatum to be analogous with the 
mammalian cortex. There is an area 
in the brain, the hippocampus, which 
borders the surface of the cerebral 
hemisphere and shows layers of cells, 
but this structure is archicortex and 
not neocortex (Cobb, 1960). 
2. The hyperstriatum is found in birds 
only. 
3. The optic lobes are well developed in 
birds. 
4. Generally, the olfactory system is 
poorly developed. Exceptions are: 
(a) those with well-developed olfac­
tory bulbs—kiwi (Apteryx australis) 
(Craigie, 1930), procellariformes 
(Bang, 1965); (b) those with well-
developed olfactory nerves—turkey 
vulture {Cathartes aura), Trinidad 
oil bird (Steatornis catipensis), Lay-
san albatros (Dromedea immutabilis), 
black-footed albatros (£>. nigripes) 
(Bang, 1960); (c) those with well-
developed olfactory tubercles— 
honey guides {Indicator indicator; 
I. minor and / . variegatus) (Stager, 
1967). 
In general, birds with small olfactory 
bulbs live in trees, and those with larger 
olfactory bulbs live in water, marshes, or 
on the ground (Cobb, 1960). Smell may 
be important for some species to locate 
their food, but the evidence is only ob­
servational (Stager, 1967). However, 
Michelsen (1959) using operant condition­
ing showed that pigeons discriminate be-
SYMPOSIUM: PHYSIOLOGICAL RESPONSE AND STRESS 11 
tween odors (secondary butyl acetate, 
iso-octane) and no odor. 
FUNCTIONS 
Hyperstriatum. The results obtained from 
various stimulating or lesion techniques 
indicate that in many respects the hy­
perstriatum performs functions similar 
to those associated with the mammalian 
cortex. This applies also to the results on 
recordings of evoked potentials and of the 
correlation between behavioral sleep and 
electrical activity. 
The hyperstriatum appears to play a 
role in the function of the auditory sys­
tem, and possibly in the visual system, cf. 
chickens and pigeons. Bremer et al. (1939) 
and Cohen and Pitts (1967) explored the 
function of the hyperstriatum and the 
corticoid regions of the pigeon and found 
that stimulation from different areas re­
sulted in different direction of head move­
ments. These head movements were ac­
companied by appropriate body and limb 
movements. Head orientating response to 
sound stimulation was attributed to the 
chicken's hyperstriatum by Adamo and 
Bennett (1967) as a result of lesions in this 
brain area producing more incorrect re­
sponses to a sound source. Adamo and 
King (1967), using the chicken, found 
that auditory stimuli elicited evoked 
potentials on the hyperstriatum which 
could be abolished by local application of 
procaine. Rougeul (1957) noted latent 
responses from the hyperstriatum in re­
sponse to electrical stimulation of the 
optic nerve. Photostimuli-evoked poten­
tials are more widely distributed than 
those evoked by auditory stimulation and 
they are not abolished by local application 
of procaine (Adamo and King, 1967). 
Cutaneous electrical (tactile) stimuli did 
not elicit responses from the hyperstria­
tum although a response was obtained 
from the deeper neostriatum (Adamo and 
King, 1967). In general, the distribution 
of points from which evoked potentials 
can be recorded seems to be more localized 
in mammals than in birds. 
Bremer et al. (1939) found spontaneous 
activity recorded from the surface of the 
avian brain to be similar to that of the 
rabbit. Durkovic and Cohen (1968) noted 
that steady potential changes recorded 
from the hyperstriatum of pigeons were 
quite similar to those found in mammals. 
Pigeons and chickens, as found by Bures 
et al. (1960) and Ookawa and Gotoh 
(1965a), respectively, show Leao's spread­
ing depression in the hyperstriatum in 
spite of its lack of layers of cells (Cobb, 
1960). In mammals, the Leao's spreading 
depression of spontaneous electrical ac­
tivity as a result of electrical, mechanical, 
or chemical stimuli is limited to the cor­
tical grey matter; in birds the depression 
effect is a "three dimensional" phenom­
enon (Bures et al., 1960) since there is no 
continuous layer of white matter. 
In chicks there is good correlation be­
tween the behavorial sleep of the chick 
and the electro-"cortico"gram (Ookawa 
and Gotoh, 1965b; Peters et al., 1965). 
The observed periods of fast waves are 
probably analogous to the so-called para­
doxical sleep of the mammal. According to 
Klein et al. (1964) paradoxical sleep does 
not occupy more than 0.6% of the total 
sleep in adult pigeons and chickens, as 
compared to a value of 15-20% for the 
mammals investigated. 
Hyperstriatal lesions have no effect on 
body temperatures, food and water in­
take, respiratory and cardiac rates and 
cardiac conditioning (Cohen, 1967). How­
ever, an initial cardio-acceleration and an 
effect on respiration (acceleration not al­
ways accompanied by an increase in 
amplitude) was observed following electro­
stimulation (Cohen and Pitts, 1967). 
Lesions did not affect the pigeon's normal 
12 A. VAN TlENHOVEN 
behavior, such as avoidance, feeding, 
accuracy of pecking at a key (Zeigler, 
1963a), but they did influence discrimina­
tion responses in pigeons (Zeigler, 1963b) 
or reversal learning in white quail (Stett-
ner and Schultz, 1967). According to Tuge 
and Shima (1959) hyperstriatal lesions 
have no effect on conditioned defensive 
responses. The effects noted were transi­
ent; 6 months later no differences were 
detected between controls and lesioned 
birds (Zeigler, 1963b). Thus, hyperstriatal 
lesions show no obvious effects; rather 
sophisticated methods are required to find 
their effects. 
Hypothalamus. This area has many func­
tions bu t for this brief review the subjects 
to be considered are regulation of feed and 
water and one aspect related to adrenal 
function. 
1. Regulation of food intake in mam­
mals occurs in two hypothalamic centers, 
the "satiety center" in the ventromedial 
nucleus and the "feeding center" in the 
lateral hypothalamus (see Mayer and 
Thomas, 1967). The areas which control 
similar responses in the chicken have not 
been adequately described. Feldman et al. 
(1957) reported tha t electrolytic lesions in 
the "anterior medial or posterior hypo­
tha lamus" caused aphagia. Inspection of 
his photographs of these brain lesions in­
dicates that the lesions were in the s t ra tum 
cellulare externum and in themedial 
posterior hypothalamic nucleus. Lep-
kovsky and Yasuda (1966) produced 
hyperphagia and obesity by lesions in 
"wha t may be assumed to be the ventro­
medial area" of the chicken. From dia­
grams given by Akerman et al. (1960) us­
ing the pigeon, electrical stimulation of 
the area ventralis anterior, the s t ra tum 
cellulare externum and septal area caused 
eating and prolonged stimulation overeat­
ing. Goodman and Brown (1966) did not 
elicit eating by stimulation of the septum 
of one pigeon. Some students working in 
my laboratory found tha t goldthioglucose 
injections to the ventromedial nucleus in 
day-old Japanese quail, a t levels close to 
the LD50 failed to induce hyperphagia and 
obesity (Carpenter, Silverstein and Stein, 
1968, unpublished data) . Obviously, addi­
tional research is needed to define more 
accurately neural control of feeding and 
satiety in birds. 
2. Polydipsia (excessive thirst) results 
from electrolytic lesions (made with stain­
less steel electrodes) of the chicken's 
nucleus supraopticus, one of the neuro­
secretory nuclei of the hypothalamus 
(Ralph, 1960), or of the hypothalami o-
hypophyseal tract plus the tuberal nucleus 
and mammilary nucleus (McFarland, 
1959). Posterior pituitary lobectomy does 
the same thing (Shirley and Nalbandov, 
1956), and this is not surprising because 
the antidiuretic hormones are produced in 
hypothalamus and transported to the 
posterior pituitary for storage (Scharrer 
and Scharrer, 1954). Lesions (technique 
not given) of the dorsal hypothalamus 
(exact location not given) of the chicken 
caused adipsia (Lepkovsky and Yasuda, 
1967). 
Akerman et al. (1960) induced drinking, 
during stimulation, by electrically stimu­
lating the nucleus paraventricularis mag-
nocellularis and of the nucleus preopticus 
medialis; whereas poststimulation drink­
ing was obtained after electric stimulation 
of the nucleus preopticus lateralis, the 
s t ra tum cellulare and area lateral to the 
nucleus paraventriculares magnocellu-
laris. Histological and histochemical 
changes are observed in the cells of the 
supraoptic nucleus and the posterior pitui­
tary of the white-crowned-sparrow and 
chicken following dehydration (Farner 
et al., 1964), indicating increased activity 
(Legait, 1959). 
SYMPOSIUM: PHYSIOLOGICAL RESPONSE AND STRESS 13 
3. There are some features which ap­
pear to be unique for at least some species 
of birds regarding adrenal and brain rela­
tionship. The adrenal of the pigeon atro­
phies after hypophysectomy, but will 
hypertrophy in such birds given formalin 
injections (a non-specific stress). This re­
sponse was not prevented by lesions in the 
median eminence. On the contrary, Miller 
(1961) reported that hypertrophy of the 
adrenal occurred following lesions of the 
median eminence and without formalin 
injections. Resko et al. (1964) had found 
that damage to the median eminence de­
creased the concentration of fluorescent 
material (presumably corticosterone) in 
the adrenal venous plasma. These data 
appear to be in conflict with those of 
Frankel et al. (1967b), who produced a de­
crease in corticosterone concentrations of 
adrenal vein plasma and a decrease in 
adrenal weight by lesions of the ventral 
tuberal hypothalamus of intact cockerels. 
Similar lesions in hypophysectomized 
cockerels did not appear to alter these 
values significantly from those obtained in 
the lesioned, intact cockerel (Frankel et 
al., 1967a). Nagra et al. (1963) failed to 
obtain decreased adrenal weight following 
hypophysectomy whereas, Frankel et al. 
(1967b) observed atrophy of the adrenal 
and a loss of differentiation between 
interrenal and chromaffin tissue following 
hypophysectomy. Frankel and co-workers 
in a series of experiments using chickens 
were also able to demonstrate that hypo­
physectomy results in a decrease in cor­
ticosterone concentration in adrenal vein 
effluent plasma (Frankel et al., 1967a,b; 
Resko et al., 1964), and in decreased out­
put of corticosterone produced in vitro by 
the adrenal. 
Intact control cockerels respond to a 
one-hour and 4-hour stress by an out­
pouring of adrenal corticosterone, whereas 
hypophysectomized cockerels responded 
only to a one-hour stress (Frankel et al., 
1967a). Nagra et al. (1963) obtained higher 
steroid concentration in adrenal vein 
blood at 1, 2, 3 and 4 hours, than at 0 
hours after surgery carried out to obtain 
blood. However, when Nagra et al. (1963) 
superimposed cold stress the corticoster­
one level decreased in 2 hours, a situation 
resembling the findings of Frankel et al. 
(1967a) with respect to lack of a response 
after long term stress. Cold stress did not 
increase the response in intact roosters. 
The data obtained by Miller (1961) and 
Frankel et al. (1967a,b) indicate an extra 
hypophyseal source of ACTH. The pineal 
gland may be the source of the extra 
hypophyseal ACTH as may be deduced 
from the following findings: 
a. 5-hydroxytryptamine is produced by 
avian pineal glands (Quay, 1966). 
b. 5-hydroxytryptamine can stimulate 
corticosterone production in vitro of 
adrenals of hypophysectomized rats 
(Gromova et al., 1967). 
Obviously experiments need to be car­
ried out to verify this hypothesis. Con­
siderable attention has been directed to­
ward elucidation of the hypothalamo-
hypophyseal-gonadal system and its stim­
ulation by light. Two receptor systems are 
known to produce gonadotrophin secre­
tion in the drake by light, i.e. a superficial 
system via the retina and optic nerve, and 
a deep receptor which is probably located 
in or near the hypothalamus (see reviews 
by Benoit, 1964a,b). The latter is involved 
in the gonadal response of blinded drakes 
to light stimulation. 
The anatomical evidence for the exis­
tence of a retino-hypothalamic tract by 
which photostimuli can reach the hypo­
thalamus has been a matter of contro­
versy. Blumcke (1961) and Karten (1967) 
have found evidence of degeneration in 
the hypothalamus after removal of the eye 
of chickens and pigeons, respectively. 
14 A. VAN TlENHOVEN 
Cowan et al. (1961) and Oksche (1968) 
failed to find such signs of degeneration. 
The photosexual reflex is a rather slow 
response and there seems to be no a priori 
reason for the need of a direct connection 
of the eye to the hypothalamus for this 
particular response. Karten (1967) has 
pointed out tha t an alternate pathway in­
volves the retina, optic nerve, optic tract, 
the dorsolateral anterior thalamus pars 
lateralis, the nucleus intercalatus, the 
hyperstr iatum accessorium, and the trac-
tus septo mesencephalicus to the hypo­
thalamus. 
The manner in which light causes the 
photosexual reflex in blinded birds has not 
been elucidated. The lack of evidence for 
photoreceptors in the hypothalamus itself 
makes it at tractive to consider the possi­
bility that the pineal gland may be in­
volved. As far as we know, no experi­
ments in which the photosexual response 
has been measured in blinded-pinealec-
tomized birds have been published. How­
ever, indirect evidence may be considered: 
1. In rats (Reiter et al., 1968) and 
hamsters (Reiter et al., 1966) pinea-
lectomy prevents the regression of 
the gonads and secondary sex organs 
which normally occurs after blind­
ing. In the English sparrow Passer 
domesticus the synchronization of 
circadian activity is not affected by 
pinealectomy (Menaker, 1968) but 
the pineal has been found to play a 
role in maintenance of the circadian 
rhythm of activity of English spar­
rows placed under constant condi­
tions (Gaston and Menaker, 1968). 
2. In an interesting experiment, Ka to 
el al. (1967) have found tha t applica­
tion of a red radio-luminous paint 
(maximum emission at 6,000 A.) on 
skin above the pineal body prevented 
the regression of testes of Japanese 
quail when the light exposure was 
changed from continuous light to 8 
hours of light and 16 hours of dark­
ness. Such regression occurred in 
controls and in birds with green 
radio-luminouspaint (maximum 
emission at 5,200 A.). 
If one considers these different lines of 
evidence together we may conclude tha t 
the role of the pineal in the photosexual 
response needs to be further investigated 
but that neither the evidence for nor 
against its role has been convincingly 
proven. 
The role of various brain areas in be­
havior will not be discussed because lack 
of space prevents me from discussing it 
adequately. 
In summary, the evidence reviewed 
shows some impressive similarities be­
tween the function of the hyperstr iatum 
and the mammalian cortex. The normal 
performance of birds, in many respects, 
with large hyperstriatal lesions points 
out that other brain areas must play an 
important role in the day-to-day activities 
of birds. The function of these areas have 
been inadequately explored. 
The importance of the pineal gland 
either as an endocrine organ or as a pos­
sible regulation of pituitary function in 
blinded birds needs further investigation 
but tentatively the evidence available 
suggests that it might be a source for 
regulation of the adrenal cortex in hypo-
physectomized birds for the regulation of 
and gonadotrophin secretion in blinded 
birds. 
ACKNOWLEDGMENTS 
The author wants to extend his sincere 
appreciation to Dr. D. Polin of Norwich 
Pharmacal Company, Norwich, N.Y. for 
his review of the manuscript and valuable 
editorial suggestions. 
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SYMPOSIUM: PHYSIOLOGICAL R E S P O N S E AND STRESS 15 
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All programs are easily accessed and operated by 
the nutritionist, purchasing agent, production 
manager or a secretary. 
Copies of the booklet may be obtained from 
Computerized Technology Department, Monsanto 
Company, 800 North Lindberg Blvd., St. Louis, 
Missouri 63166. 
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183-194. 
W.P.C. NOTES 
At the Western Poultry Congress held at San 
Diego, California, October 31, November 1 and 2, 
the following officers were elected: President—N. 
Coleman, Modesto, California; First Vice-Presi­
dent—D. Wharton, Sacramento, California; Sec­
ond Vice-President—E. M. Demler, Anaheim, Cal-
NEWS AND NOTES 
(Continued from page 8) 
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