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=== Breathing=== [[File:Indian softshell turtle (Nilssonia gangetica) Babai River nose-breathing at surface.jpg|thumb|A submerged [[Indian softshell turtle]] nose-breathing at river surface|alt=photo of a river turtle with only its nose above water]] The rigid shell of turtles is not capable of expanding and making room for the lungs, as in other amniotes, so they have had to evolve special adaptations for respiration.<ref name="Cordeiro 2016">{{Cite journal |last1=Cordeiro |first1=Tábata E. F. |last2=Abe |first2=Augusto S. |last3=Klein |first3=Wilfried |date=April 2016 |title=Ventilation and Gas Exchange in Two Turtles: ''Podocnemis unifilis'' and ''Phrynops geoffroanus'' (Testudines: Pleurodira) |journal=Respiratory Physiology & Neurobiology |volume=224 |pages=125–131 |doi=10.1016/j.resp.2014.12.010 |pmid=25534144 |issn=1569-9048 |hdl=11449/158795 |s2cid=37446604 |url=https://repositorio.unesp.br/bitstream/11449/158795/1/WOS000373540800015.pdf |hdl-access=free |access-date=September 25, 2019 |archive-date=July 24, 2021 |archive-url=https://web.archive.org/web/20210724105252/https://repositorio.unesp.br/bitstream/handle/11449/158795/WOS000373540800015.pdf;jsessionid=7500C1BBBBCF7CD5F9A0DE4D94B0AC03?sequence=1 |url-status=live }}</ref><ref name="Lyson 2014">{{Cite journal |last1=Lyson |first1=Tyler R. |last2=Schachner |first2=Emma R. |last3=Botha-Brink |first3=Jennifer |last4=Scheyer |first4=Torsten M. |last5=Lambertz |first5=Markus |last6=Bever |first6=G. S. |last7=Rubidge |first7=Bruce S. |last8=de Queiroz |first8=Kevin |date=7 November 2014 |title=Origin of the Unique Ventilatory Apparatus of Turtles |journal=Nature Communications |volume=5 |page=5211 |doi=10.1038/ncomms6211 |pmid=25376734 |issn=2041-1723 |bibcode=2014NatCo...5.5211L|doi-access=free }}</ref><ref>{{Cite journal |last1=Lee |first1=Stella Y. |last2=Milsom |first2=William K. |year=2016 |title=The Metabolic Cost of Breathing in Red-eared Sliders: An Attempt to Resolve an Old Controversy |journal=Respiratory Physiology & Neurobiology |volume=224 |pages=114–124 |doi=10.1016/j.resp.2015.10.011 |pmid=26524718 |s2cid=5194890 |issn=1569-9048}}</ref> The lungs of turtles are attached directly to the carapace above while below, connective tissue attaches them to the organs.{{sfn|Orenstein|2012|p=41}} They have multiple lateral (side) and medial (middle) chambers (the numbers of which vary between species) and one terminal (end) chamber.<ref>{{Cite journal |last1=Lambertz |first1=Markus |last2=Böhme |first2=Wolfgang |last3=Perry |first3=Steven F. |date=July 2010 |title=The Anatomy of the Respiratory System in ''Platysternon megacephalum'' Gray, 1831 (Testudines: Cryptodira) and Related Species, and its Phylogenetic Implications |journal=Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology |volume=156 |issue=3 |pages=330–336 |doi=10.1016/j.cbpa.2009.12.016 |pmid=20044019 |issn=1095-6433}}</ref> The lungs are ventilated using specific groups of abdominal muscles attached to the organs that pull and push on them.<ref name="Cordeiro 2016"/> Specifically, it is the turtle's large liver that compresses the lungs. Underneath the lungs, in the [[coelomic cavity]], the liver is connected to the right lung by the [[Root of the lung|root]], and the stomach is directly attached to the left lung, and to the liver by a [[mesentery]]. When the liver is pulled down, inhalation begins.<ref name="Lyson 2014"/> Supporting the lungs is a wall or [[septum]], which is thought to prevent them from collapsing.<ref>{{Cite journal |last1=Klein |first1=Wilfried |last2=Codd |first2=Jonathan R. |year=2010 |title=Breathing and Locomotion: Comparative Anatomy, Morphology and Function |journal=Respiratory Physiology & Neurobiology |volume=173 |pages=S26–S32 |doi=10.1016/j.resp.2010.04.019|pmid=20417316 |s2cid=28044326 |issn=1569-9048}}</ref> During exhalation, the contraction of the [[transversus abdominis muscle]] propels the organs into the lungs and expels air. Conversely, during inhalation, the relaxing and flattening of the [[Abdominal external oblique muscle|oblique abdominis muscle]] pulls the transversus back down, allowing air back into the lungs.<ref name="Lyson 2014"/> Although many turtles spend large amounts of their lives underwater, all turtles breathe air and must surface at regular intervals to refill their lungs. Depending on the species, immersion periods vary between a minute and an hour.<ref>{{Cite journal |last1=Morera-Brenes |first1=Bernal |last2=Monge-Nájera |first2=Julián |year=2011 |title=Immersion Periods in Four Neotropical Turtles |url=https://revistas.uned.ac.cr/index.php/cuadernos/article/view/212 |journal=UNED Research Journal |volume=3 |issue=1 |pages=97 |doi=10.22458/urj.v3i1.212 |doi-access=free |access-date=June 11, 2020 |archive-date=August 9, 2020 |archive-url=https://web.archive.org/web/20200809164614/https://revistas.uned.ac.cr/index.php/cuadernos/article/view/212 |url-status=live }}</ref> Some species can [[Enteral respiration|respire through the cloaca]], which contains large sacs that are lined with many finger-like projections that take up dissolved [[oxygen]] from the water.<ref>{{Cite journal |last1=Priest |first1=Toni E. |date=December 2002 |last2=Franklin |first2=Craig E. |title=Effect of Water Temperature and Oxygen Levels on the Diving Behavior of Two Freshwater Turtles: ''Rheodytes leukops'' and ''Emydura macquarii'' |journal=Journal of Herpetology |volume=36 |issue=4 |pages=555–561 |jstor=1565924 |issn=0022-1511 |doi=10.1670/0022-1511(2002)036[0555:EOWTAO]2.0.CO;2 |s2cid=85279910 }}</ref>
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