Editing Microbat (section)

== Echolocation ==
{{main|Animal echolocation}}
[[Animal echolocation|Echolocation]] is the process where an animal produces a sound of certain wavelength, and then listens to and compares the reflected echoes to the original sound emitted. Bats use echolocation to form images of their surrounding environment and the organisms that inhabit it by eliciting [[Ultrasound|ultrasonic waves]] via their [[larynx]].<ref name=":0">{{Cite journal|last1=Davies|first1=Kalina T. J.|last2=Maryanto|first2=Ibnu|last3=Rossiter|first3=Stephen J.|date=2013-01-01|title=Evolutionary origins of ultrasonic hearing and laryngeal echolocation in bats inferred from morphological analyses of the inner ear|journal=Frontiers in Zoology|volume=10|issue=1|pages=2|doi=10.1186/1742-9994-10-2|issn=1742-9994|pmc=3598973|pmid=23360746 |doi-access=free }}</ref><ref name=":1">{{Cite journal|last1=Reichard|first1=Jonathan D.|last2=Fellows|first2=Spenser R.|last3=Frank|first3=Alexander J.|last4=Kunz|first4=Thomas H.|date=2010-11-01|title=Thermoregulation during Flight: Body Temperature and Sensible Heat Transfer in Free-Ranging Brazilian Free-Tailed Bats (Tadarida brasiliensis)|journal=Physiological and Biochemical Zoology|volume=83|issue=6|pages=885–897|doi=10.1086/657253|pmid=21034204|s2cid=1028970|issn=1522-2152}}</ref> The difference between the ultrasonic waves produced by the bat and what the bat hears provides the bat with information about its environment. Echolocation aids the bat in not only detecting prey, but also in orientation during flight.<ref name=":2">{{Cite book|title=Handbook of Behavioral Neuroscience|last1=Berke|first1=Gerald S.|last2=Long|first2=Jennifer L.|date=2010-01-01|publisher=Elsevier|editor-last=Brudzynski|editor-first=Stefan M.|series=Handbook of Mammalian VocalizationAn Integrative Neuroscience Approach|volume=19|pages=419–426|doi=10.1016/B978-0-12-374593-4.00038-3|chapter=Functions of the larynx and production of sounds|isbn=9780123745934}}</ref>

=== Production of ultrasonic waves ===
Most microbats generate ultrasound with their larynx and emit the sound through their nose or mouth.<ref>{{Cite journal|last1=Hedenström|first1=Anders|last2=Johansson|first2=L. Christoffer|date=2015-03-01|title=Bat flight: aerodynamics, kinematics and flight morphology|journal=Journal of Experimental Biology|volume=218|issue=5|pages=653–663|doi=10.1242/jeb.031203|issn=0022-0949|pmid=25740899|doi-access=free|bibcode=2015JExpB.218..653H }}</ref> Sound productions are generated from the [[vocal folds]] in mammals due to the elastic membranes that compose these folds. Vocalization requires these elastic membranes because they act as a source to transform airflow into acoustic pressure waves. Energy is supplied to the elastic membranes from the lungs, and results in the production of sound. The larynx houses the [[Vocal folds|vocal cords]] and forms the passageway for the expiratory air that will produce sound.<ref>{{Cite journal|last1=Holbrook|first1=K A|last2=Odland|first2=G F|date=1978-05-01|title=A collagen and elastic network in the wing of the bat.|journal=Journal of Anatomy|volume=126|issue=Pt 1|pages=21–36|issn=0021-8782|pmc=1235709|pmid=649500}}</ref> Microbat {{Audio|Ultrasonic_bat_calls.ogg|calls}} range in frequency from 14,000 to over 100,000 [[hertz]], well beyond the range of the human ear (typical human hearing range is considered to be from 20 to 20,000&nbsp;Hz). The emitted vocalizations form a broad beam of sound used to probe the environment, as well as communicate with other bats. At the molecular level, it has been found that [[CPLX1]] is involved in this ultrasonic wave production.<ref>{{Cite journal |last1=Liu |first1=Meiling |last2=Wang |first2=Changliang |last3=Huo |first3=Lifang |last4=Cao |first4=Jie |last5=Mao |first5=Xiuguang |last6=He |first6=Ziqing |last7=Hu |first7=Chuanxia |last8=Sun |first8=Haijian |last9=Deng |first9=Wenjun |last10=He |first10=Weiya |last11=Chen |first11=Yifu |last12=Gu |first12=Meifeng |last13=Liao |first13=Jiayu |last14=Guo |first14=Ning |last15=He |first15=Xiangyang |date=July 2024 |title=Complexin-1 enhances ultrasound neurotransmission in the mammalian auditory pathway |url=https://www.nature.com/articles/s41588-024-01781-z |journal=Nature Genetics |volume=56 |issue=7 |pages=1503–1515 |doi=10.1038/s41588-024-01781-z |pmid=38834904 |issn=1546-1718|url-access=subscription }}</ref>

=== Laryngeally echolocating microbats ===
[[File:BIOL352-WikiPic-2.jpg|thumb|alt=The skull of a microbat|Ventral view of a Florida Freetail bat (''Tadarida cyanocephala'') skull, highlighting both the stylohyal and tympanic bones. Specimen from the Pacific Lutheran University Natural History collection.]]

Laryngeal echolocation is the dominant form of echolocation in microbats, however, it is not the only way in which microbats can produce ultrasonic waves. Excluding non-echolocating and laryngeally echolocating microbats, other species of microbats and megabats have been shown to produce [[Ultrasound|ultrasonic waves]] by clapping their wings, clicking their tongues, or using their nose.<ref name=":0" /> Laryngeally echolocating bats, in general, produce ultrasonic waves with their larynx that is specialized to produce sounds of short [[wavelength]]. The [[larynx]] is located at the [[Cranium|cranial]] end of the [[trachea]] and is surrounded by [[cricothyroid muscle]]s and [[thyroid cartilage]]. For reference, in [[human]]s, this is the area where the [[Adam's apple]] is located. Phonation of ultrasonic waves is produced through the vibrations of the vocal membranes in the expiratory air. The intensity that these vocal folds vibrate at varies with activity and between bat species.<ref>{{Cite journal|last1=Simmons|first1=Nancy B.|last2=Seymour|first2=Kevin L.|last3=Habersetzer|first3=Jörg|last4=Gunnell|first4=Gregg F.|date=2010-08-19|title=Inferring echolocation in ancient bats|journal=Nature|volume=466|issue=7309|pages=E8|doi=10.1038/nature09219|pmid=20724993|issn=0028-0836|bibcode=2010Natur.466E...8S|doi-access=free}}</ref> A characteristic of laryngeally echolocating microbats that distinguishes them from other echolocating microbats is the articulation of their [[stylohyal bone]] with their [[Tympanic part of the temporal bone|tympanic bone]]. The stylohyal bones are part of the [[hyoid apparatus]] that help support the throat and larynx. The tympanic bone forms the floor of the [[middle ear]]. In addition to the connection between the stylohyal bone and the tympanic bone as being an indicator of laryngeally echolocating microbats, another definitive marker is the presence of a flattened and expanded stylohyal bone at the cranial end.<ref name=":1" />

Microbats that laryngeally echolocate must be able to distinguish between the differences of the pulse that they produce and the returning echo that follows by being able to process and understand the [[Ultrasound|ultrasonic waves]] at a [[neuron]]al level, in order to accurately obtain information about their surrounding environment and orientation in it.<ref name=":0" /> The connection between the stylohyal bone and the tympanic bone enables the bat to neurally register the outgoing and incoming ultrasonic waves produced by the [[larynx]].<ref name=":2" /> Furthermore, the stylohyal bones connect the larynx to the tympanic bones via a [[cartilaginous]] or [[Fibrous joint|fibrous]] connection (depending on the species of bat). Mechanically the importance of this connection is that it supports the larynx by anchoring it to the surrounding [[cricothyroid muscle]]s, as well as draws it closer to the [[nasal cavity]] during [[phonation]]. The stylohyal bones are often reduced in many other mammals, however, they are more prominent in laryngeally echolocating bats and are part of the mammalian hyoid apparatus. The hyoid apparatus functions in breathing, swallowing, and phonation in microbats as well as other mammals. An important feature of the bony connection in laryngeally echolocating microbats is the extended articulation of the ventral portion of the tympanic bones and the proximal end of the stylohyal bone that bends around it to make this connection.<ref name=":0" />