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Animal echolocation
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Echolocation, also called Biosonar, is the biological sonar used by several animals such as dolphins, shrews, most bats, and most whales. The term was coined by Donald Griffin, who was the first to conclusively demonstrate its existence in bats. Two bird groups also employ this system for navigating through caves, the so called cave swiftlets in the genus Aerodramus (formerly Collocalia) and the unrelated Oilbird Steatornis caripensis.
Echolocating animals emit calls out to the environment, and listen to the echoes of those calls that return from various objects in the environment. They use these echoes to locate, range, and identify the objects. Echolocation is used for navigation and for foraging (or hunting) in various environments.
Basic principleEcholocation works like active sonar, using sounds made by an animal. Ranging is done by measuring the time delay between the animal's own sound emission and any echoes that return from the environment. Unlike some sonar that relies on an extremely narrow beam to localize a target, animal echolocation relies on multiple receivers. Echolocating animals have two ears positioned slightly apart. The echoes returning to the two ears arrive at different times and at different loudness levels, depending on the position of the object generating the echoes. The time and loudness differences are used by the animals to perceive direction. With echolocation the bat or other animal can see not only where it's going but can also see how big another animal is, what kind of animal it is, and other features as well.
BatsMicrobats use echolocation to navigate and forage, often in total darkness. They generally emerge from their roosts in caves or attics at dusk and forage for insects into the night. Their use of echolocation allows them to occupy a niche where there are often many insects (that come out at night since there are fewer predators then) and where there is less competition for food, and where there are fewer other species that may prey on the bats themselves.
Microbats generate ultrasound via the larynx and emit the sound through the nose or, much more commonly, the open mouth. Microbat
Individual bat species echolocate within specific frequency ranges that suit their environment and prey types. This has sometimes been used by researchers to identify bats flying in an area simply by recording their calls with ultrasonic recorders known as 'bat detectors'. However echolocation calls are not species specific and some bats overlap in the type of calls they use so recordings of echolocation calls cannot be used to identify all bats. In recent years researchers in several countries have developed 'bat call libraries' that contain recordings of local bat species that have been identified known as 'reference calls' to assist with identification.
Since the 1970s there has been an ongoing controversy among researchers as to whether bats use a form of processing known from radar termed coherent cross-correlation. Coherence means that the phase of the echolocation signals is used by the bats, while cross-correlation just implies that the outgoing signal is compared with the returning echoes in a running process. Today most - but not all - researchers believe that they use cross-correlation, but in an incoherent form, termed a filter bank receiver.
When searching for prey they produce sounds at a low rate (10-20/sec). During the search phase the sound emission is coupled to respiration, which is again coupled to the wingbeat. It is speculated that this coupling conserves energy. After detecting a potential prey item, microbats increase the rate of pulses, ending with the terminal buzz, at rates as high as 200/sec. During approach to a detected target, the duration of the sounds is gradually decreasing, as is the energy of the sound.
Bat echolocation: Neural mechanisms in the brainBecause bats use echolocation to orient themselves and to locate objects, their auditory systems are adapted for this purpose, highly specialized for sensing and interpreting the stereotyped echolocation calls characteristic of their own species. This specialization is evident from the inner ear up to the highest levels of information processing in the auditory cortex.
Inner ear and primary sensory neurons
Both CF and FM bats have specialized inner ears which allow them to hear sounds in the ultrasonic range, far outside the range of human hearing. Although in most other aspects, the bat’s auditory organs are similar to those of most other mammals, certain bats with a constant frequency component to their call (known as CF bats) do have a few additional adaptations for detecting the predominant frequency (and harmonics) of the CF vocalization. These include a narrow frequency “tuning” of the inner ear organs, with an especially large area responding to the frequency of the bat’s returning echoes (Neuweiler 2003).
The basilar membrane within the cochlea contains the first of these specializations for echo information processing. In bats that use CF signals, the section of membrane that responds to the frequency of returning echoes is much larger than the region of response for any other frequency. For example, in Rhinolophus ferrumequinum, the horseshoe bat, there is a disproportionately lengthened and thickened section of the membrane that responds to sounds around 83 kHz, the constant frequency of the echo produced by the bat’s call. This area of high sensitivity to a specific, narrow range of frequency is known as an “acoustic fovea” (Zupanc 2004).
Further along the auditory pathway, the movement of the basilar membrane results in the stimulation of primary auditory neurons. Many of these neurons are specifically “tuned” (respond most strongly) to the narrow frequency range of returning echoes of CF calls. Because of the large size of the acoustic fovea, the number of neurons responding to this region, and thus to the echo frequency, is especially high (Carew 2001).
Inferior colliculus
In the inferior collicus, a structure in the bat’s midbrain, information from lower in the auditory processing pathway is integrated and sent on to the auditory cortex.
As George Pollak and others showed in a series of papers in 1977, the interneurons in this region have a very high level of sensitivity to time differences, since the time delay between a call and the returning echo tells the bat its distance from the target object. Especially interesting is that while most neurons respond more quickly to stronger stimuli, collicular neurons maintain their timing accuracy even as signal intensity changes.
These interneurons are specialized for time sensitivity in several ways. First, when activated, they generally respond with only one or two action potentials. This short duration of response allows their action potentials to give a very specific indication of the exact moment of the time when the stimulus arrived, and to respond accurately to stimuli that occur close in time to one another. In addition, the neurons have a very low threshold of activation – they respond quickly even to weak stimuli. Finally, for FM signals, each interneuron is tuned to a specific frequency within the sweep, as well as to that same frequency in the following echo.
There is specialization for the CF component of the call at this level as well. The high proportion of neurons responding to the frequency of the acoustic fovea actually increases at this level (Carew 2001, Pollak 1977, Zupanc 2004).
Auditory cortex
The auditory cortex in bats is quite large in comparison with other mammals (Anderson 1995). Various characteristics of sound are processed by different regions of the cortex, each providing different information about the location or movement of a target object.
Most of the existing studies on information processing in the auditory cortex of the bat have been done by Nobuo Suga on the mustached bat, Pteronotus parnellii. This bat’s call has both CF tone and FM sweep components.
Suga and his colleagues have shown that the cortex contains a series of “maps” of auditory information, each of which is organized systematically based on characteristics of sound such as frequency and amplitude. The neurons in these areas respond only to a specific combination of frequency and timing (sound-echo delay), and are known as combination-sensitive neurons.
- FM-FM area: This region of the cortex contains FM-FM combination-sensitive neurons. These cells respond only to the combination of two FM sweeps: a call and its echo. The neurons in the FM-FM region are often referred to as “delay-tuned,” since each responds to a specific time delay between the original call and the echo, in order to find the distance from the target object (the range). Each neuron also shows specificity for one harmonic in the original call and a different harmonic in the echo. The neurons within the FM-FM area of the cortex of Pteronotus are organized into columns, in which the delay time is constant vertically but increases across the horizontal plane. The result is that range is encoded by location on the cortex, and increases systematically across the FM-FM area (Suga et al. 1975, Suga et al. 1979, Neuweiler 2003, Carew 2001).
- CF-CF area: Another kind of combination-sensitive neuron is the CF-CF neuron. These respond best to the combination of a CF call containing two given frequencies – a call at 30 kHz (CF1) and one of its additional harmonics around 60 or 90 kHz (CF2 or CF3) – and the corresponding echoes. Thus, within the CF-CF region, the changes in echo frequency caused by the Doppler shift can be compared to the frequency of the original call to calculate the bat’s velocity relative to its target object. As in the FM-FM area, information is encoded by its location within the map-like organization of the region. The CF-CF area is first split into the distinct CF1-CF2 and CF1-CF3 areas. Within each area, the CF1 frequency is organized on an axis, perpendicular to the CF2 or CF3 frequency axis. In the resulting grid, each neuron codes for a certain combination of frequencies that is indicative of a specific velocity (Suga et al. 1975, Suga et al. 1987, Carew 2001).
- DSCF area: This large section of the cortex is a map of the acoustic fovea, organized by frequency and by amplitude. Neurons in this region respond to CF signals that have been Doppler shifted (in other words, echoes only) and are within the same narrow frequency range to which the acoustic fovea responds. For Pteronotus, this is around 61 kHz. This area is organized into columns, which are arranged radially based on frequency. Within a column, each neuron responds to a specific combination of frequency and amplitude. Suga’s studies have indicated that this brain region is necessary for frequency discrimination (Suga et al. 1975, Suga et al. 1987, Carew 2001).
To summarize, the systematically organized maps in the auditory cortex respond to various aspects of the echo signal, such as its delay and its velocity. These regions are composed of “combination sensitive” neurons that require at least two specific stimuli to elicit a response. The neurons vary systematically across the maps, which are organized by acoustic features of the sound and can be two dimensional. The different features of the call and its echo are used by the bat to determine important characteristics of their prey.
Toothed whales
Toothed whales (suborder odontoceti), including dolphins, porpoises, river dolphins, orcas and sperm whales, use biosonar because they live in an underwater habitat that has favourable acoustic characteristics and where vision is extremely limited in range due to absorption or turbidity.
Toothed whales emit a focused beam of high-frequency clicks in the direction that their head is pointing. Sounds are generated by passing air from the bony nares through the phonic lips. These sounds are reflected by the dense concave bone of the cranium and an air sac at its base. The focussed beam is modulated by a large fatty organ known as the 'melon'. This acts like an acoustic lens because it is composed of lipids of differing densities. Most toothed whales use clicks in a series, or click train, for echolocation, while the sperm whale may produce clicks individually. Different rates of click production in a click train give rise to the familiar barks, squeals and growls of the bottlenose dolphin. A click train with a repetition rate over 600 per second is called a burst pulse. In bottlenose dolphins, the auditory brain response resolves individual clicks up to 600 per second, but yields a graded response for higher repetition rates.
Some smaller toothed whales may have their tooth arrangement suited to aid in echolocation. The placement of teeth in the jaw of a bottlenose dolphin, as an example, are not symmetrical when seen from a vertical plane, and this asymmetry could possibly be an aid in the dolphin sensing if echoes from its biosonar are coming from one side or the other.
Echoes are received using the lower jaw as the primary reception path, from where they are transmitted to the inner ear via a continuous fat body. Lateral sound may be received though fatty lobes surrounding the ears with a similar acoustic density to bone. Some researchers believe that when they approach the object of interest, they protect themselves against the louder echo by quietening the emitted sound. In bats this is known to happen, but here the hearing sensitivity is also reduced close to a target.
Before the echolocation abilities of "porpoises" were officially discovered, Jacques Yves Cousteau suggested that they might exist. In his first book, The Silent World (1953, pp. 206-207), he reported that his research vessel, the Élie Monier, was heading to the Straits of Gibraltar and noticed a group of porpoises following them. Cousteau changed course a few degrees off the optimal course to the center of the strait, and the porpoises followed for a few minutes, then diverged toward mid-channel again. It was obvious that they knew where the optimal course lay, even if the humans didn't. Cousteau concluded that the cetaceans had something like sonar, which was a relatively new feature on submarines. He was right.
OilbirdsOilbirds and some species of swiftlet are known to use a crude form of biosonar (compared to the capabilities of bats and dolphins). These nocturnal birds emit calls while flying and use the calls to navigate through trees and caves where they live.
Shrews and tenrecsTerrestrial mammals other than bats known to echolocate include two genera (Sorex and Blarina) of shrews and the tenrecs of Madagascar. These include the wandering shrew (Sorex vagrans), the common or Eurasian shrew (Sorex araneus), and the short-tailed shrew (Blarina brevicauda). The shrews emit series of ultrasonic squeaks. In contrast to bats, shrews probably use echolocation to investigate their habitat rather than to pinpoint food.
See also
External links- 150,000 recordings of over 10,000 species, including many echolocation recordings
- links to many bioacoustics resources
- has bat and swiftlet sonar signals
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