Frog hearing and communication
Encyclopedia
Frog
Frog
Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...

s and toad
Toad
A toad is any of a number of species of amphibians in the order Anura characterized by dry, leathery skin , short legs, and snoat-like parotoid glands...

s produce a rich variety of sounds, calls, and songs during their courtship and mating rituals. The callers, usually males, make stereotyped sounds in order to advertise their location, their mating readiness and their willingness to defend their territory; listeners respond to the calls by return calling, by approach, and by going silent. These responses have been shown to be important for species recognition, mate assessment, and localization. Beginning with the pioneering experiments of Robert Capranica in the 1960s using playback techniques with normal and synthetic calls, behavioral biologists and neurobiologists have teamed up to use frogs and toads as a model system for understanding the auditory function and evolution. It is now considered an important example of the neural basis of animal behavior
Neuroethology
Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system...

, because of the simplicity of the sounds, the relative ease with which neurophysiological recordings can be made from the auditory nerve, and the reliability of localization behavior. Acoustic communication is essential for the frog's survival in both territorial defense and in localization and attraction of mates. Sounds from frogs travel through the air, through water, and through the substrate
Substrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...

. The neural basis of communication and audition gives insights into the science of sound applied to human communication
Human communication
Human communication, or Anthroposemiotics, is the field dedicated to understanding how people communicate:* with themselves: intrapersonal communication** expression: body language* another person: interpersonal communication...

.

Behavioral ecology

Frogs are more often heard than seen, and other frogs (and researchers) rely on their calls to identify them. Depending on the region that the frog lives in, certain times of the year are better for breeding than others, and frogs may live away from the best breeding grounds when it is not the species’ mating season. During the breeding season, they congregate to the best breeding site and compete for call time and recognition. Species that have a narrow mating season due to ponds that dry up have the most vigorous calls.

Male-male competition

In many frog species only males call. Each species has a distinct call, though even among the same species, different dialects are found in different regions. Although humans cannot detect the differences in dialects, frog distinguish between regional dialects. For example, male bullfrogs can recognize the calls of their direct territorial neighbors. By ignoring the calls of these neighbors they save energy, and only vocalize aggressively with to an intruder’s call. In this way, calls establish territories, but they also attract females.
Males may have a solitary call for times when there is no competition that uses less energy. During other times, when a frog must compete with hundreds or thousands of other frogs to be heard, together they perform a chorus call where each frog calls in turn, successively. The most important feature of the chorus is the shared pattern. Through this pattern, few individuals calls are drowned out.
One frog’s call may be dominant and trigger the calls of the responding frogs in symphony. Interestingly, calling is linked to physical size and females may be attracted to more vigorous calls.
Frogs in the same region chorus within their species and between different species. Frogs of the same species will retune their frequency so it is distinct from other frogs of the same species. Different species of frogs living in the same region have more dramatically different call frequencies. The frequency and durations of different species' calls vary similarly to the preference of that species' females. The neural circuity of females of different species varies.

Male-female interactions

Like the males, females can distinguish the minute differences between individual frogs. However, males and females are attuned to different parts of the advertisement call. For example, males of the onomatopoeically named species Coqui
Coquí
The Common Coquí or Coquí is a frog native to Puerto Rico belonging to the Eleutherodactylus genus of the Leptodactylidae family. The species is named for the loud sound the males make at night. This sound serves two purposes...

are more attuned to the low frequency co part of the call, whereas females are more attuned to the high frequency qui. In fact, the order of the parts does not matter. Similarly, for females of the Tungara
Tungara Frog
The Túngara Frog is a species of frog in the Leiuperidae family...

species, the female basilar papilla is biased towards a lower-than-average “chuck” portion of a male call. Experiments that measure the vocal responses and approaches shows these attenuations.

Mode of sound communication

Calls are often sent through the air, but other mediums have been discovered. Some species call while they are under water and the sound travels through the water. This is adaptive in a region with many species competing for air time. Narins has found female frog species that use solid surfaces, such as blades of grass and logs, upon which they tap rhythmically to attract mates. Also, Feng has found that some species of frogs use lt

Sound production

The smallest frogs must consume lots of energy to produce calls. In addition, vocalizing muscles can make up 15% of a male spring peeper’s body mass, while the same muscles are only 3% of females. Frogs produce sound from the air sac below their mouth that from the outside, is seen to inflate and deflate. Air from the lungs is channeled to the air sac, which resonates to make the sound louder. The larynx is larger and more developed in males, though not significantly different from females.

Sound localization

Biologists believed that frogs ears are placed too close together to localize sound
Sound localization
Sound localization refers to a listener's ability to identify the location or origin of a detected sound in direction and distance. It may also refer to the methods in acoustical engineering to simulate the placement of an auditory cue in a virtual 3D space .The sound localization mechanisms of the...

 accurately. Frogs cannot hear short, high frequency sounds. Sound is localized by the time difference when the sound reaches each ear. The “vibration spot” near the lungs vibrates in response to sound, and may be used as an additional measure to localize from.

Applications of frog neuroethology

Dr. Feng’s work applies the neuroethology
Neuroethology
Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system...

 of frog communication to medicine. A recent project on hearing aids is based on how female frogs find their mates. Females must recognize the male they choose by his call. By localizing where his call is coming from she can find him. An additional challenge is that she is localizing his call while listening to the many other frogs in the chorus, and to the noise of the stream and insects. The breeding pond is a very noisy place, and females must distinguish a male’s calls from the other noise. How they recognize the sound pattern of the male they are pursuing from the surrounding noise is similar to how intelligent hearing aids help people hear certain sounds and cancel out others. The underlying neural mechanisms are fast neural oscillations
Neural oscillations
Neural oscillation is rhythmic or repetitive neural activity in the central nervous system. Neural tissue can generate oscillatory activity in many ways, driven either by mechanisms localized within individual neurons or by interactions between neurons...

, and synaptic inhibition to cancel out noise. The timing and frequency of the sound also play a part in frog communication and may be used in Feng’s work. He also studies bat echolocation to create intelligent hearing aids. He is also working on cochlear implants.

External links

Neuroethology course link
  • http://instruct1.cit.cornell.edu/courses/bionb424/


Feng
  • http://www.life.uiuc.edu/neuroscience/people/showpeople.php?person=faculty/afeng1
  • http://www.beckman.uiuc.edu/directory/index.php?qry=BY_NETID&type=BIO&filter=afeng1
  • http://www.sciencemuseum.org.uk/antenna/ultrasonicfrogs/


Narins
  • http://www.physci.ucla.edu/Faculty/Narins/research/research.html
  • http://www.acoustics.org/press/swa9501.html
  • http://www.physci.ucla.edu/Faculty/Narins/publications/pdfs/Sun%20and%20Narins%20BC.pdf
  • http://www.physci.ucla.edu/Faculty/Narins/publications/pdfs/NarinsJCP2.pdf
  • http://www.physci.ucla.edu/Faculty/Narins/publications/publications.html


Sound library
  • http://www.animalbehaviorarchive.org/assetSearch.do?method=searchCQL&query=%22Rana%22+and+%22catesbeiana%22&firstRecord=1&maximumRecords=9&totalResults=37&view=list&sortKeys=audioQual,ascending=false
  • http://www.animalbehaviorarchive.org/assetSearchInterim.do
The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
x
OK