Physiology & Behavior 71 (2000) 203 ± 205

Short communication
Electroencephalographic study of odor responses in the domestic fowl
Tomoko Oosawa, Yoshiyuki Hirano, Keiichi Tonosaki*
Department of Veterinary Physiology, Faculty of Agriculture, Gifu University, 1-1 Yanagido, Gifu, Gifu 501-1193, Japan
Received 28 December 1999; received in revised form 10 May 2000; accepted 12 May 2000

Abstract

Although avian olfaction has been studied behaviorally and anatomically, few electrophysiological (EEG) studies exist. The purpose of
this study is to examine the characteristics of the olfactory function of the fowl with EEG recording. We found that slow waves decreased and
the rapid waves increased in response to a variety of odors. D 2000 Elsevier Science Inc. All rights reserved.

Keywords: Domestic fowl; Electroencephalogram (EEG); Geraniol; Odor; Olfaction

1. Introduction barbital (15 ± 20 mg/kg body weight). The bird was placed
in a head holder and lidocaine hydrochloride paste (0.4%)
Conventional wisdom, based on studies of brain anat-
omy, held that most birds had little or no sense of smell. We
now know, however, that even birds with relatively small
olfactory bulbs are able not only to sense odors, but use
them to find food and perform other functions [1,4 ±6,12 ±
18,21,25,31 ± 35]. Indeed, some anatomical and physiologi-
cal observations have shown that the structure and function
of the olfactory organ in birds are similar to those of other
air-breathing animals [1,5,6,11± 14,18 ±23,26 ± 35].
Olfactory receptor cell responses of birds to odors has
been reported in a number of electrophysiological (EEG)
studies [22 ±24,26,28,33,35]. The bird olfactory response to
amyl acetate is vigorous and stable, like other air-breathing
animals including the dog and box turtle [27]. The purpose
of this study is to examine the characteristics of the fowl
olfactory response to odor stimuli with EEG.

2. Materials and methods

Fourteen mature, female domestic fowls, ranging in

weight from 1.5 to 2.0 kg, were used in the study. The
experiments were performed in a quiet, ventilated room
and the temperature was controlled at 23°C. Each animal
was anesthetized with an i.p. injection of sodium pento-
* Corresponding author. Tel.: +81-58-293-2938; fax: +81-58-293-
2938.
E-mail address: tonosaki@cc.gifu-u.ac.jp (K. Tonosaki).
Fig. 1. (A) Typical example of domestic fowl's EEG response to geraniol.
(B) Frequency histogram of nine fowls' EEGs before and after odor
stimulation with geraniol. The black columns are delta waves, the open
columns are theta waves, the black and white dotted columns are alpha
waves and the white and black dotted columns are beta waves. The
numbers on the vertical axis represent the unit ratio percent ( [(a, b, d or q -
frequency/5 s)/(a + b + d + q -frequency/5 s)]100). The vertical parentheses
bars are mean ‹ SE (n = 9). Asterisk denotes the level of significance
between before and during ( p < 0.01).

0031-9384/00/$ ± see front matter D 2000 Elsevier Science Inc. All rights reserved.
PII: S 0 0 3 1 - 9 3 8 4 ( 0 0 ) 0 0 3 0 9 - 7
204 T. Oosawa et al. / Physiology & Behavior 71 (2000) 203±205

Table 1 response appears to characterize the response to an odor.

EEG response profiles of domestic fowls to odors
An increase in EEG frequency in response to an odor has
Methyl also been reported in studies with cats, rabbits and rats
n-Amyl anthranilic
[2,3,7 ±10].
Fowl no. acetate D-Limonene acid Geraniol Ionone
Table 1 shows that the fowl can respond to almost all
1 + + + + +
of the odors tested. Thus, it is clear that EEG recording,
2 + + + + +
3 + + + + as used in this study, was effective in revealing odor
4 + + + + detection of the domestic fowl. Of course, response
5 ÿ + + ÿ sensitivity across odors may vary among different types
6 + + + + + of birds. Thus, for example, pigeons respond well to
7 + ÿ + + ÿ
floral odors while vultures may respond better to odors
8 ÿ + + + +
9 + ÿ ÿ ÿ ÿ of putrefaction [22,23,26,28,32,33,35]. This present meth-
10 + ÿ + + + od may prove useful in future studies of avian olfaction
11 + ÿ + ÿ + and in examining differential odor sensitivity and other
12 + ÿ ÿ ÿ olfactory functions.
13 + + + ÿ ÿ
14 ÿ + + + +

Response profiles of EEG responses in fourteen fowls. (+) Odorant

changed EEG activity, (ÿ) odorant did not change EEG activity, blank:
not tested.
was rubbed onto the scalp for EEG recording. After
sufficient anesthetization, some feathers were clipped from
the head and two small holes (diameter about 0.5 mm)
were made on the vertex and over the external occipital
protuberance with a dental drill. Tungsten insulated fine
needle electrodes were inserted into the holes and were
fixed with dental resin.
N-amyl acetate, D-limonene, methyl anthranilic acid,
geraniol and ionone samples were diluted in propylene
glycol to form a 2% concentration. Each dilution sample
was delivered to the nostrils for 5 s. Stimulus presentation
was approximately once every 2 min. Olfactory responses
were recorded while the bird was breathing freely. An EEG
polygraph (Polygraph System, Nihon Kohden, Japan) and
frequency analyzer (Bimutas, Kissay Comtek, Japan) were
used for the recordings.
The EEG activities were recorded on a frequency modu-
lated (FM) magnetic tape and analyzed at later time.
Frequency analyses were performed for the 5-s period
before and during the odor stimulus. All values were
presented as means ‹ SE. Statistical significance was exam-
ined by two-way analysis of variance (ANOVA) and with
post hoc testing by means of Duncan's multiple-range test.
3. Results and discussion
A positive EEG response was observed in amplitude and
frequency of wave form during odor stimulus presentation.
Fig. 1A presents a typical example of EEG response to
geraniol. From the frequency analysis of nine examples,
EEG activity showed a significant decrease in slow waves
and an increase in rapid waves in response to geraniol (Fig.
1B). From these results, the high frequency of EEG
References
[1] Bang BG. Functional anatomy of the olfactory system in 23 orders of
birds. Acta Anat 1971;79:1 ± 76.
[2] Bressler SL, Frereman WJ. Frequency analysis of olfactory system
EEG in cat, rabbit, and rat. Electroencephalogr Clin Neurophysiol
1980;50:19 ± 24.
[3] Bressler SL. Spatial organization of EEGs from olfactory bulb and
cortex. Electroencephalogr Clin Neurophysiol 1984;57:270 ± 6.
[4] Burne TH, Rogers LJ. Relative importance of odour and taste in the
one-trial passive avoidance learning bead task. Physiol Behav 1997;
62:1299 ± 302.
[5] Burne TH, Rogers JJ. Responses to odorants by the domestic chick.
Physiol Behav 1996;60:1441 ± 7.
[6] Ebinger P, Rehkamper G, Schroder H. Forebrain specialization and
the olfactory system in anseriform birds. An architectonic and tracing
study. Cell Tissue Res 1992;268:81 ± 90.
[7] Freeman WJ. EEG analysis gives model of neuronal template-match-
ing mechanism for sensory search with olfactory bulb. Biol Cybern
1979;35:221 ± 34.
[8] Freeman WJ, Schneider W. Changes in spatial patterns of rabbit ol-
factory EEG with conditioning to odor. Psychophysiology 1982;
19:44 ± 56.
[9] Freeman WJ, Viana DI, Prisco G. Relation of olfactory EEG to be-
havior: time series analysis. Behav Neurosci 1986;100:753 ± 63.
[10] Freeman WJ, Grajski KA. Relation of olfactory EEG to behavior:
factor analysis. Behav Neurosci 1987;101:766 ± 77.
[11] Gasser HS. Olfactory nerve fibers. J Gen Physiol 1956;39:473 ± 96.
[12] Gentle MJ. Sensory involvement in the control of food intake in
poultry. Proc Nutr Soc 1985;44:313 ± 21.
[13] Harriman AE, Berger RH. Olfactory acuity in the common raven.
Physiol Behav 1986;36:257 ± 62.
[14] Ioale P, Papi F. Olfactory bulb size, odor discrimination and magnetic
insensitivity in hummingbirds. Physiol Behav 1989;45:995 ± 9.
[15] Jones RB, Roper TJ. Olfaction in the domestic fowl: a critical review.
Physiol Behav 1997;62:1009 ± 18.
[16] Jones RB, Gentle MJ. Olfaction and behavioral modification in do-
mestic chicks (Gallus domesticus). Physiol Behav 1985;34:917 ± 24.
[17] Mabayo RT, Okumura J, Hirao A, Sugita S, Sugahara K, Furuse M.
The role of olfaction in oil preference in the chicken. Physiol Behav
1996;59:1185 ± 8.
[18] Malakoff D. Following the scent of avian olfaction. Science 1999;
286:704 ± 5.
[19] Porter RH, Hepper PG, Bouchat C, Picard M. A simple for testing
odor detection and discrimination in chicks. Physiol Behav 1999;67:
459 ± 62.
 T. Oosawa et al. / Physiology & Behavior 71 (2000) 203±205
[20] Porter RH, Picard M. Effects of early odor exposure in domestic
chicks. Reprod Nutr Dev 1998;38:441 ± 8.
[21] Rieke GK, Wenzel BM. Forebrain projections of the pigeon olfactory
bulb. J Morphol 1978;158:41 ± 55.
[22] Shibuya Y, Tucker D. Single unit responses of olfactory receptors in
vulture. In: Hayashi T, editor. Olfaction and taste II. Oxford: Perga-
mon, 1965. pp. 219 ± 33.
[23] Shibuya T, Iijima M, Tonosaki K. Responses of the olfactory nerve in
the seagull (Larus crassirofris). Zool Mag 1970;79:237 ± 9.
[24] Shibuya T, Tonosaki K. Electrical responses of single olfactory recep-
tor cells in some vertebrates. In: Schneider D, editor. Olfaction and
taste IV. Stuttgart: Wissenshaftliche Verlagsgesellshaft, 1972. pp.
102 ± 8.
[25] Stattelman AJ, Talbot RB, Coulter DB. Olfactory thresholds of pi-
geons (Columba livia), quail (Colinus verginianus) and chickens (Gal-
lus gallus). Comp Biochem Physiol, Part A: Mol Integr Physiol
1975;50:807 ± 9.
[26] Tonosaki K, Shibuya T. Olfactory receptor cell responses of pigeon to
some odors. Comp Biochem Physiol, Part A: Mol Integr Physiol
1985;81:329 ± 33.
205
[27] Tonosaki K, Tucker D. Olfactory receptor cell responses of dog and
box turtle to aliphatic n-acetates and aliphatic n-fatty acids. Behav
Neural Biol 1982;35:187 ± 99.
[28] Tucker D. Electrophysiological evidence for olfactory function in
bird. Nature 1965;207:34 ± 6.
[29] Walker JC. An operant procedure for testing olfactory capacities in
restrained pigeons. Physiol Behav 1983;30:165 ± 8.
[30] Walker JC, Walker DB, Tambiah CR, Gilmore KS. Olfactory and
nonolfactory odor detection in pigeons: elucidation by a cardiac ac-
celeration paradigm. Physiol Behav 1986;38:575 ± 80.
[31] Wenzel BM, Sieck MH. Olfaction. Annu Rev Physiol 1966; 28:
381 ± 434.
[32] Wenzel BM, Salzman A. Olfactory bulb ablation or nerve section and
behavior of pigeons in nonolfactory learning. Exp Neurol 1968;
22:472 ± 9.
[33] Wemzel BM. Olfactory prowess of the kiwi. Nature 1968;14:1133 ± 4.
[34] Wenzel BM. Olfactory sensation in the kiwi and other birds. Ann NY
Acad Sci 1971;188:183 ± 93.
[35] Wenzel BM, Sieck MH. Olfactory perception and bulbar electrical
activity in several avian species. Physiol Behav 1972;9:287 ± 93.