Copyright 2001
By Devorah A. N. Bennu
All Rights Reserved.
This article appeared in the Autumn 2001 newsletter published by the Biology Department at the University of Washington.

The evening was peaceful as the sun settled below the horizon. Suddenly,
the orange and red sky was dotted with flying blue-grey silhouettes
zigzagging above Lake Union. Bats! Some bats dipped low over the water's
surface to drink in a manner similar to swallows, while others followed
erratic courses above, consuming mosquitoes and other insects in flight.
How do bats find and capture insects in darkness? All insect-eating
bats produce sounds -- either with open mouths or through an elaborate
nose-leaf that functions like a megaphone -- that are reflected as
faint echoes from objects in the environment, providing bats an accurate
picture of their environment and allowing them to maneuver in complete
darkness.
We didn't always know that bats rely on sound to move through dark
surroundings. In the late 1800s, Swiss zoologist Charles Jurine's
suggestion that bats could "see" using their ears was rejected as
preposterous by most of his colleagues. However, in 1944, Donald
Griffin, then an undergrad at Harvard University, used special
microphones to show that bats generate ultrasonic sounds above the
range of human hearing to create echoes that reveal the locations
of objects. As a result, Griffin coined the term "echolocation"
to describe this phenomenon.
All bats communicate using a variety of squeaks and squeals that
human ears can hear, but bat species in the taxonomic suborder
Microchiroptera emit high-frequency (high-pitched) sounds for
echolocation. Even though low frequency sounds travel farther, only
small ultrasonic sound waves used for echolocation can provide
detailed information about tiny objects in the environment, such as
flying mosquitoes.
Echolocation sounds are not the same pitch for all bat species. High
frequency echolocation calls produced by bats range from 20-200
kilohertz (kHz) or more (humans cannot hear sounds above 20kHz).
There are differences in the pattern of echolocation calls, as well.
Some species produce a constant frequency (CF) cry, while others
use a frequency modulated (FM) call that descends in pitch. However,
most bat species can choose to use either type of call.
CF and FM sounds reveal different types of information to the bat.
CF is used to detect objects while FM provides distance and other
finely detailed data about the nature of the object. For example,
by comparing the time elapsed between outgoing calls and returning
echoes, bats accurately estimate distance to an object. Deviations
in echo intensity and pitch reveal important details about the
target, such as direction, size, shape and velocity. Most bat species
produce a complicated sequence of CF and FM calls that they modify
apparently depending upon differences in habitat, such as open areas
versus dense vegetation.
Because air absorbs energy contained in sound waves, the usefulness
of high-pitched (short wave) sounds such as those produced by
echolocating bats are limited to a range of 50 feet or less, making
echolocationg bats nearsighted. To partially compensate for this
energy loss and to increase their range, most bats produce high
intensity sounds of up to 120 decibels, which is as loud as a smoke
detector held four inches from your ear. These "shouting" bat species
include the Little Brown Bat, Myotis lucifugus, and the Big
Brown Bat, Eptesicus fuscus, which are common in Seattle
neighborhoods.
How do bats make such loud sounds without deafening themselves? Shortly
before the bat's larynx muscles contract to produce an ultrasonic
call, special muscles in the middle ear contract, separating the three
inner ear bones (the malleus, incus and stapes or "hammer," "anvil" and
"stirrup"), causing momentary deafness. After the call has been made,
the middle ear muscles relax, restoring the bat's hearing in time to receive
echoes from objects that are as close as one meter away.
There still are many unanswered questions about the acoustic abilities
of bats. However, research into echolocation is already yielding
practical benefits for humans. For example, sonar is a primitive form
of echolocation (from a bat's perspective!) that is widely used for
navigation, tracking aircraft, ships, submarines and missiles, and for
forecasting weather.

This graphic reveals how echolocation is
used by a bat
to reveal the location of a
tasty meal, such as a moth.
[Graphic
courtesy of Bob and Rita at bats4kids]

The Little Brown Bat, Myotis lucifugus,
is the most common bat in the Seattle area.
It
typically roosts in attics or barns near water. It is
a "shouting bat."
[Photo copyright by the Missouri
Department of Conservation.]
The author acknowledges Bob and Rita from the website, bats4kids, who graciously provided permission to use their echolocation graphic, and the Missouri Department of Conservation for their excellent photograph of the Little Brown Bat in flight. The author also thanks Dr. Van Tiggelen from the Belgian Museum, who read this article and kindly corrected my misspelling of Donald Griffin's name.
[10 October 2001]