GENERAL I ARTICLE

Echolocation
The Strange Ways of Bats

G Marimuthu

Bats are capable of avoiding obstacles that they e n c o u n t e r ,

even in c o m p l e t e darkness. T h i s is because they e m i t
ultrasound (high frequency sound) and analyse the e c h o
produced w h e n t h e sound hits objects on their path. T h i s
article d e s c r i b e s t h e h u n t i n g flight o f bats a n d h o w
echolocation is useful in prey capture. Prey capture without
G Marimuthu has studied the aid of echolocation by s o m e bats is also described.
the behaviour of bats for
almost two decades. His
The March 1996 issue of Resonance introduced us to the
pioneering experiments
have led to an understand-
fascinating world of bats, the only flying mammals of the world.
ing of how bats catch frogs As opposed to traditional views on bats, they are a harmless and
in total darkness. interesting group of animals. Awareness about bats and the need
to conserve them has increased considerably in recent years. A
very interesting feature which was only briefly mentioned in
Resonance Vol. 1, No.3, is the ability of bats to 'see' through their
ears. The microchiropteran bats use a special property called
'echolocation', both to avoid obstacles on their way and to locate
and capture their prey.

Echolocation is a specialized process of orientation used by bats.

Bats emit high frequency sound waves while navigating, and
process the echo that comes back from obstacles. This method
Bats emit high assists prey location and capture.
frequency sound
waves while The Discovery of Echolocation
navigating, and
process the echo lhat During the year 1790, Lazzaro Spallanzani, an Italian naturalist,
comes back from first observed that bats were able to avoid obstacles while flying
obstades. This melhod even in total darkness. He also found that despite the surgical
assists prey location removal of eyes, bats could fly without bumping into obstacles.
and caplure. Later, Charles Jurine, a Swiss zoologist, plugged the ears of bats

40 RESONANCE t May 1996

 GENERAL J ARTICLE

and observed their inability to perform these correct orientations.

Spallanzani repeated these experiments and obtained similar
results. Both of them concluded that bats could 'see' through
their ears! The French naturalist Cuvier disagreed with this
statement. He explained that a sense of touch in the wing
membrane caused the bats to avoid obstacles. In 1920, Hartridge,
a British physiologist put forward the hypothesis that bats emit
ultrasound and listen to the echoes of these sounds.After 18 years,
the American zoologist, Donald R Grifl'm along with Pierce, a Donald R Griffin showed
physicist, used a microphone sensitive to ultrasound and that bats emit ultrasonic
demonstrated that bats do emit trains of ultrasonic pulses while pulse~
flying. They showed that the number of sound pulses increased
as bats approached obstacles on their flight path. They also
noticed that the bat's mouth was always open when the sounds
were emitted. Griffin continued the experiments and found that
closing the mouth of the bat resulted in disorientation. He
established that bats emit sounds through their mouths. I~ was
Griffin who coined the term 'echolocation' in 1938. In 1958, he
published his classic book, 'Listening in the Dark' which
documents many details about the discovery of echolocation.
Echolocation is one of the methods of orientation mainly used by
the microchiropteran or insectivorous bats. While flying, these
bats emit high frequency ultrasound. These sound pulses hit
obstacles like rocks, trees, walls etc. and their echoes are heard by
bats. By analysing these echoes, bats are able to find their way
even deep into underground caves in which there is absolutely no
light.
it was Griffin who
Vocalizations of Bats coined the term
"echolocation' in 1938.
Like other mammals, including humans, bats emit sound through In 1958, he published
the voice box or larynx. Sound is produced when the vocal chords his dassic book,
vibrate as air passes over them. Hence these sounds are called 'lJstening in 1he Dark'
vocalizations. The muscles in the larynx adjust the tension on the which documents
vocal chords. This controls the rate of vibration of the vocal many details about
chords which explains the frequency or pitch of the sound 1he discovelv of
emitted. Some of the characteristics of sound are shown in the echolocation.

RESONANCE I May 1996 41

 GENERAL J ARTICLE

box. Most of the species (eg. Indian pygmy bat, free-tailed bat,
tomb bat) emit their echolocation sounds through the mouth. A
few other species (eg. Indian false vampire bat, leaf nosed-bat,
horseshoe bat) produce their vocalizations through the nostrils.
The latter species have grotesque facial ornamentations. This is
known as the noseleaf(Figure 1). It is a shallow, parabolic portion
surrounding the nostrils and a spear shaped, rounded or fleshy
superior portion. The structure of the noseleafvaries from species
to species. The noseleaf serves to narrow and focus the outgoing
beam of sound.

Vocalizations used in echolocation are generally divided into two

figure I The face of the categories.
Indian false vampire bat
Megadenna lyre. It is a 9 Broadband signals These cover a wide range of frequencies,
microchiroptemn a n d car-
from 20 to 140 kHz and have shorter durations of less than 5
n/vorousbnt./t woighsabaut
40 g. While flying it emits milliseconds. They are technically called frequency modu-
ultraseunds through its lated (FM) pulses. Each pulse starts at a high frequency and
noslrils, which help to beam sweeps down to lower frequency within a short duration.
the sound pulses. The huge 9 Narrowbandsignals These have a constant frequency (CF) and
pinnae ore able to collect longer durations of about 100 milliseconds.
the faint noise created while
the prey moves.
F u n c t i o n s of E c h o l o c a t i o n

Even though there are two such distinct kinds of sounds

(Figure2), bats use eitherone or combinations of both depending
on the situation and gather detailed information on their flight
path. The hunting flightof bats is divided into three stages: the
search stage,the approach stage and the terminal stage(Figure3).
During the search stage, bats emit sound pulses with a low
repetition rate of about 10 pulses per second. Actually a correla-
The hunting flight of tion exists between the habitat in which a species regularly
bats is divided into forages and the type of signal itemits at this stage.Usually short
three stages: the C F pulses, with or without an F M rail,arc found in species that
search stage, the forage in open spaces where vegetation and other obstacles arc
approach stage and not found. Bats that hunt close to vegetation or the ground, emit
the terminal stage. pulses which mainly have an F M sweep. Theoretically, the

42 RESONANCE I May 1996

 GENERAL I ARTICLE

amount of information available from a signal is proportional to Figure 2 Sonograms of

its bandwidth. A broadband outgoing sound pulse would cause a different types of echoloca-
tion sounds shown as fre-
greater number of altered frequencies in the returning echoes.
quency in the ordinate and
Bats use such echoes to analyse the features of the target, for
duration in the abscissa
example to differentiate prey from the background clutter and to scales : (A). Steep broad-
differentiate smooth and rough surfaces suitable for landing. bond FM signal starts at o
They can accurately discriminate between targets that are within higherfrequency and ends
10-15 mm of each other. To estimate the target range (distance), in a lower frequency in a
bats analyse the time delay between the emission of sound and its short duration. (B). Steep
FM signal ends with a shal-
return as echo just like a radar detects objects several metres away.
low FM component. (C). A
long CF narrewband signal
A few other species like horseshoe bats which emit narrowband with an initial increasing FM
signals with longer CF component use an alternative strategy of and a decreasing FM tall at
echolocation. They distinguish the moving prey from nonmoving the end. (D). Signal starts as
obstacles by means ofthe Doppler effect(see box and Resonance, a shallow FM with a long CF
component.
Vol.1, No.2, Page 14).
Figure 3 Pattern o f the
The main function of the search stage is t o detect the potential
emission of echolocation
prey among obstacles. The big brown bat Eptesicusfuscus can pulses by bats which emit
detect a sphere having a diameter of 2 cm, at a distance of Sin. The only FM signals (top) and
same bat detects a 0.5 cm sphere at a distance of 3 m. bats which emit a combina-
tion of both CF and FM sig-
The onset of the approach stage represents the first visible nals (bottom) at three dif-
reaction of the bat to the target. This stage begins when the bat ferent stages. (S) - search
stage, (A) - approach stage,
is between 1 or 2 metres away from the target. The bat turns its
(1") - terminal stage.
head and ears towards the target. It also increases the repetition
rate of the echolocation sounds to about 40 pulses per second. In

RESONANCE I May 1996 43

 GENERAL J ARTICLE

Characteristics of Sound

Sound is a series of vibrations in air or water for the loudness of various sounds. The echolocation
example, picked up by the ears and interpreted sound of bats is about 110 dB at 10 cm in front of
as a sensation by the brain. A few characteristics a bat's mouth. This is slightly more intense than
of sound are relevant to echolocation. The fre- the sound from a milk cooker, a common vessel
quency or pitch of the sound of bats is measured in the kitchen.
in kilohertz, abbreviated as kHz. One kHz is one
thousand cycles per second or 1000 Hertz. Hu- Theoretically a bat receives the echo of its sounds
mans can hear up to 20 kHz. Sounds having a within 500 milliseconds (1000 milliseconds = 1
higher frequency than this are called ultrasound. sec ). The obstacles could be away at a maximum
The echolocation calls of bats are inaudible to distance of about 85 m. The bat detects the
humans and hence called ultrasonic. Since high distance of the target by measuring the time
frequency sounds are more rapidly absorbed by interval between the emitted signal and its echo.
the atmosphere, the echolocatory system works The range of frequencies of the echolocation
within a limited distance. The intensilyof sound is pulses is the bandwidth of the signal. The power
measured in decibels, abbreviated as dB. This spectrum explains the distribution of energy on
unit is related to the ratio of the sound intensity to the frequencies of the signal (see Figure 4). A bat
a standard, which is taken to be the threshold collects detailed information about targets b y
sound intensity detectable by the human ear. comparing the power spectra of the emitted
Table I provides the decibel scale to measure sound and its echo.

Table 1. The decibel scale

d8 Examples dB Examples

10 Rustling leaves 80 Vacuum cleaner

20 Whisper 90 Classroom in a school
30 100
40 Voices in city night 110 A road drill
SO Normal speech 120
60 A busy super market 130 Jet aircraft take off
7O 140 Painful sounds

The hearing sensitivily addition to these changes, a qualitative change in the pulse
of bats is much pattern also occurs. In species (eg.Myotis rnyotis) which emit only
higher than FM pulses, the slope of the FM sweep becomes steeper, the
other mammals. duration of the pulse becomes shorter but the bandwidth of the

44 RESONANCE I May 1996

 GENERAL I ARTICLE

Figure 4 Thespech~rom
of the echolocation coil (A)
and its echo (B). The spec-
hal difference between the
pulse and the echo provides
detailed information about
the target structur~

signal remains the same. In a few other species like Nyctalus

noctu/a which emit only CF pulses during the search stage, an
abrupt switch to emitting brief FM pulses occurs. The CF
component is dropped. Horseshoe bats which use long CF-FM The big brown bat
pulses during the search stage do not drop the CF component at Eptesicus [uscus can
the approach stage. Their pulses become shorter with an increase detect a sphere
in bandwidth of the FM component. having a diometer of
2 cm, at a distance
Thus a shift towards emission of FM pulses is discernible atthe of 5m. The same
approach stage. Since the information content is greater in the bat detects o 0.5 a n
broadcast (FM) signals, this shift is useful to decide whether to sphere at a
catch a prey or to avoid an obstacle or to land on a roosting site. distance of 3 m.

RESONANCE I May 1996 45

 GENERAL J ARTICLE

Doppler Shift

Our ears hear a changed sound when we listen moves, the greater the change in frequency.
to a sound source which moves rapidly towards This effect of motion on the frequency of sounds
or away from us, eg. a car passing us with its was first pointed out by an Austrian scientist
horn blowing. We experience a sudden drop in Christian Doppler and it is named after him as
frequency as the car passes away from us. Even Doppler shift.
though we hear a sudden change in frequency,
the horn actually sends out sound waves at a The long constant frequency signal of the echolo-
regular interval. If we stand ahead of the car cation sounds emitted by a few species of bats is
( person 'A" in Figure 5 ) our ears receive more used for measuring the Doppler shift but is not
than the normal number of sound waves and we suitable for target description. Bats are able to
hear a higher frequency than the real tone of the analyse the shifts that occur in the echo fre-
horn. After the car passes, our ears receive fewer quency produced by a flying insect. They use this
sound waves ( person 'B' in Figure 5 ) so that the method to detect the insect prey from the large
frequency becomes lower, with a sudden drop at amount of echo clutter produced by the dense
the moment the car passes us. The faster the car foliage or other background objects.

When bats reach the target within a distance of 50 cm, the

terminal stage begins. A steep increase in the repetition of the
emission of about 100 or even 200 pulses per second occurs. This
increased rate rapidly updates the information and the bat makes
the final decision whether or not to catch the prey. This rapid
increasein the emission of sound pulsesduring the terminal stage
is termed as 'finalbuzz'. In most bat species,the sound pulses
emitted at this stage arc only F M sweeps with three or four
harmonics. These arc of lower duration of within 0.5 millisec-
onds. In batswhich use the Doppler shift,likethe horseshoe bats,
the C F component stillremains but isreduced in duration and is
about 10 milliseconds (compared to 60 milliseconds during
search stage).After detecting the insects,bats capture them by
A few other species using theirwing membranes and transferthem to their mouth.
like horseshoe bats
distinguish the moving The hearing sensitivity of bats is much higher than other
pray from nonmoving mammals. This specializationallows bats to receiveand analyse
obstades by means of faintechoes. W h e n the echoes return to them, they arc received
the Doppler effect by the auditory system similar to other mammalian patterns.

46 RESONANCE I May 1996

 GENERAL J ARTICLE

Figure 5 Each sound wove

storts out as o circle. Since
the horn is moving forword
continuously, the centre of
each circle is a l i m e farther
along the rood than the pre-
v/ous one. This makes the
wove 'crowded" (high fre-
quency) in front (A) and
"stretched out" (low fre-
quency) at the beck (8).

Echolocating bats have prominent external ears. Their pinnae

are specialized to amplify the faint echoes. The mechanical
vibrations of the echoes travel through the ear drum, middle ear
and reach the cochlea of the inner ear. A helical ribbon, known as
the basilar membrane, present in the cochlea contains hair cells.
These are the receptor cells that convert the mechanical vibra-
tions of the echoes into electrical signals and transmit them to the
brain along the auditory nerve. Processing of the echoes takes
place in the brain. The processing includes information such as
the pulse-echo delay and comparison of spectral features of the
original sound and its echoes. From this process a bat gets an
'acoustic picture' of its flight path.

P r e y Capture without E c h o l o c a t i o n

Recent studies show that some species of bats do not use echolo-
cation to detect their prey. These are the false vampire bats in
India, Australia and Africa, long eared bats in North America,
mouse eared bats in Europe, fringe lipped bats in Panama and
slit-faced bats in Africa. The Indian false vampire bats listen
passively to the noise associated with the movement of the prey Recent studies show
(frogs, mice, larger insects, etc.). The fringe-lipped bats use the that some species of
songs of male frogs to locate and capture them. bats do not use
echolocation to
They can distinguish the calls of edible frogs from those of detect 1heir prey.

RESONANCE J May 1996 47

 GENERAL J ARTICLE

The vampire bats of poisonous toads.The vampire bats (livingonly in Central and
Cenkal and South South America) use the breathing noise of the carrieto locate
America use the and to feed upon their blood. All these species of bats produce
breathing noise of the faint echolocation signals but use them only to gather
caltie to locate information about the background. They are hence known as
and to feed upon whispering bats. Echolocation is a unique and fascinating
their blood. characteristic of bats.

Suggested Reading
Address for correspondence
G Marimu~u D R Griffin. Listening in the Dark. Yale University Press, New Haven. 1958.
Department of Animal G Neuweiler. Echolocation and Adaptivity to Ecological Constraints. In:
Behaviour & Physiology, Neuroethology and Behavioural Physiology. (Eds) F Huber and H Markl.
School of Biological Sciences, Springer Verlag, Berlin. 1983. 280-302.
Madurai Kamaraj UnNersily, D Young. Nerve Cells and Animal Behaviour. Cambridge University Press.
Madurai 625 021. India. Cambridge. New York, Sydney. 1989.

\ /
V
Kanizsa triangle ... consists of illusory contours. A normal visual cortex sees a
triangle even though interconnecting lines are missing. Such illusions showthat the
visual cortex must resolve conflicts between different functional areas.

Honeybees can see optical illusions ... Brazilian researchers have found that
bees rewarded with a sugar solution can be taught to "see" Kanizsa triangles.

411 RESONANCE J May 1996