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so-called dances, of the honey bee. It increased from 0.4 second, if the
Stingless bees, which do not use dances feeding place was near the hive, to
for communication, use sound signals to 1.5 seconds if the feeding place was 100 200 300 400 500 600 700
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tica. The latter marked the way from family of bees. The duration of single a flash of colored light typically oc-
the feeding place to the hive with scent sound periods gives the distance of a cupies one half of a cycle and a flash
marks, and the foraging bee guided its feeding place in some species of sting- of a standard reference white light,
hive-mates along these marks to the less bees and in the honey bee. Sting- the other half of the cycle. With a
feeding station (2). With T. (Axesto- less 'bees do not dance. These more suitably adjusted flash rate, the lumi-
trigona) tescorum we observed that the primitive bees guide the hive-mates nance of the two components, colored
returning bees stimulated their hive- personally to the feeding place by and standard white, is assumed to be
mates to "sing." In a short time, sound tracing of odor or by repeated indica- equated when, by varying the lumi-
was made by most of the hive mem- tion of flight direction. This personal nance of the colored component, mini-
bers, who then left the hive in great guiding is symbolized in the dances of mum flicker is perceived. Thus, the
numbers to search for food. The sound the more highly evolved genus Apis perceived magnitude of the flicker is
record of this event, when played back by the waggle run (5), which forms a U-shaped function of the luminance
into the hive later, did not stimulate the main part of the dance in which of the variable component, and the
the singing behavior but did increase both distance and direction of the feed- minimum of the function defines equal
the number of bees leaving the hive. ing place are indicated. luminance (4). In the study reported
With all other Thigona, the playing HARALD ESCH*X here we examined the relation be-
back of the authentic sound increased ILSE ESCH tween this psychophysical point of min-
only the number of bees which ran Strahlenbiologisches Institut, imum flicker and the amplitude of the
to the hive entrance, but not the num- Untiversitdt Miinchen, evoked brain potential.
ber actually leaving. It may be that Munich, Germany We used three observers in all; the
smell is an important factor in the WARWICK E. KERRt data for two of them are presented
communication of these bees. Faculdade de Filosofia, Ciencias e here. Two alternating stimulus com-
In the course of our experiments we Letras, Rio Claro, Sao Paulo, Brazil ponents were presented to the left eye
trained individuals of Bombus atratus in Maxwellian view as a circular patch,
References and Notes
to visit artificial feeding places. No centrally fixated, subtending a visual
foraging bee of Bombus produced a 1. K. v. Frisch, Oesterr. Zool. Z. 1, 1 (1946). angle of 3.6 degrees. A concentric
2. M. Lindauer and W. E. Kerr, Z. Vergleich.
characteristic sound signal upon return- Physiol. 41, 405 (1958). white surround, also in Maxwellian
3. H. Esch, ibid. 45, 1 (1961).
ing to the hive, nor did any bee bring 4. H. Esch, in Verhandl. Deut. Zool. Ges. 26,
view, subtended a visual angle of 31.4
another worker bee to the feeding sta- 302 (1962); Z. Vergleich. Physiol. 48, 534 degrees. Thus, the angles subtended
(1964); A. M. Wenner, Animal Behav. 10, 79
tion. Bombus obviously cannot com- (1962). by the stimuli approached those con-
municate information about feeding 5. H. Esch, Umschau 10, 293 (1962). sidered optimal for flicker photometry
6. This work was supported by the Deutsche
stations. Forschungsgemeinschaft and the Fundacao de (5). The light sources were tungsten-
These results enable us to form a Amparo a Pescisa do Estado de Sao Paulo. ribbon filament bulbs (General Elec-
We thank Prof. von Frisch for supporting the
new hypothesis of evolutionary develop- work and for suggestions concerning this paper. tric), run at 18 amp. The luminance
ment of communication behavior con- * Present address: Department of Biology, Uni- of the surround field was kept con-
versity of Notre Dame, Notre Dame, Indiana.
cerning distance and direction of food t Present address: Departam6nto de Gen6tica, stant and equal to the luminance of
Faculdade de Medicina de Ribeirao Preto, Es-
sources. Sound is an element of com- tado de Sao Paulo, Brazil. the standard white component of the
munication which is widespread in the 12 April 1965 H flickering stimulus, namely 368 mlam.
Cycling of the flash was controlled by
shutters mounted on small stepping
motors (6), driven by Tektronix wave-
form and pulse generators. Each cycle
Evoked Brain Potential Correlates of Psychophysical consisted of a flash of white light fol-
lowed immediately by a flash of spec-
Responses: Heterochromatic Flicker Photometry
tral light. Evoked brain potentials were
Abstract. The relation between the amplitude of evoked brain potentials in man amplified by Tektronix Type 122 am-
and the relative luminance of two flicker components of different color was de- plifiers and were summed with a
termined. The function, which is U-shaped, has a minimum which occurs near Mnemotron Computer of Average
the point of equal luminance as judged by the psychophysical method of flicker Transients. The computer was triggered
photometry. by the output of the same wave-form
generator which drives the pulse gen-
The data from many studies of hu- tential amplitude-luminance function to erators controlling the shutters. Thus,
man brain potentials evoked by visual flickering stimuli (2). Only one previ- the computer analysis was accurately
stimuli show that various parameters ous study has attempted to directly time-locked with the presentation of
of the response are correlated with the compare simultaneously obtained psy- the light flashes. A 62.5-msec epoch
physical parameters of the light stimu- chophysical estimates with evoked brain was used with a sampling rate of 1600
lus. For example, numerous studies potential measures of responses to per second. The frequency of stimula-
have shown that the amplitude of the flickering visual stimuli (3). tion was 16 cy/sec.
evoked brain potential is a function of We have investigated the relation Three simultaneous recordings were
the luminance of the stimulus (1). between the evoked brain potential and obtained from the scalp of each ob-
Some investigators have attempted to luminance, as measured by a version server: (i) bipolar, between the inion
ascertain the relation between the of the psychophysical method of and a point 5 cm directly above on
electroretinogram amplitude-luminance heterochromatic flicker photometry. the midline; (ii) monopolar, between
function and the evoked brain po- With this method of visual photometry, the inion electrode and a reference
16 JULY 1965 321
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