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Respiratory Function and Mechanics in Pinnipeds and Cetaceans Andreas Fahlman1,2,*, Michael J

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Respiratory Function and Mechanics in Pinnipeds and Cetaceans Andreas Fahlman1,2,*, Michael J © 2017. Published by The Company of Biologists Ltd | Journal of Experimental Biology (2017) 220, 1761-1773 doi:10.1242/jeb.126870 REVIEW Respiratory function and mechanics in pinnipeds and cetaceans Andreas Fahlman1,2,*, Michael J. Moore3 and Daniel Garcia-Parraga1,4 ABSTRACT There are several reviews that describe anatomical features of In this Review, we focus on the functional properties of the respiratory diving marine mammals (e.g. Piscitelli et al., 2013), but these system of pinnipeds and cetaceans, and briefly summarize the reviews focus on the structural properties of excised tissues, which underlying anatomy; in doing so, we provide an overview of what is may not always reflect the functional properties of live animals currently known about their respiratory physiology and mechanics. (Kooyman, 1973; Ponganis, 2015). For example, compliance While exposure to high pressure is a common challenge among estimates of excised tissues do not account for the influence of breath-hold divers, there is a large variation in respiratory anatomy, surrounding structures that encase the respiratory system (Cozzi function and capacity between species – how are these traits adapted et al., 2005; Fahlman et al., 2011, 2014; Moore et al., 2014). to allow the animals to withstand the physiological challenges faced Likewise, pulmonary volume changes during compression of entire during dives? The ultra-deep diving feats of some marine mammals dead specimens (Moore et al., 2011) cannot account for the defy our current understanding of respiratory physiology and lung potential effects of blood engorgement of the tracheal mucosa in mechanics. These animals cope daily with lung compression, cetaceans (Leith, 1976; Cozzi et al., 2005; Davenport et al., 2013). alveolar collapse, transient hyperoxia and extreme hypoxia. By Thus, the functional properties of the whole living animal cannot be improving our understanding of respiratory physiology under these determined from deceased specimens. Here, we discuss recent conditions, we will be better able to define the physiological advances in our understanding of pulmonary mechanics and lung constraints imposed on these animals, and how these limitations function that we believe provide a theoretical framework that can may affect the survival of marine mammals in a changing merge past and future studies to enhance our knowledge of the traits environment. Many of the respiratory traits to survive exposure to that allow deep diving. an extreme environment may inspire novel treatments for a variety of ’ respiratory problems in humans. Scholander s legacy, the model of lung/alveolar collapse Scholander (1940) argued that the compliances of the respiratory KEY WORDS: Compliance, Marine mammal, Lung function, system, with a flexible thorax, would allow the elastic and highly Respiratory flow, Tidal volume, Residual volume, Total lung capacity, compliant alveoli to compress and push the air into the more rigid Respiratory frequency, Alveolar collapse (far less compliant) conducting airways. As the alveoli compress and collapse, the gas diffusion rate would decrease and cause a Introduction pulmonary shunt that increases with depth until the alveoli fully In 1940, Per Scholander published his 131-page-long monograph collapse and gas exchange ceases (Fig. 1A,D). Pulmonary shunt on cardiorespiratory function in marine mammals and birds. In his represents the amount of blood bypassing the lung and not treatise, he summarized the respiratory and cardiovascular traits participating in gas exchange, and it varies between 0% and required by marine mammals to manage life in an extreme 100%, where 0% represents a fully inflated lung with perfect gas environment and cope daily with challenges such as alveolar exchange, and 100% represents termination of gas exchange. collapse (atelectasis; see Glossary), alveolar recruitment (see Scholander assumed that the trachea behaved like an idealized non- Glossary), transient hyperoxia, extreme hypoxia and compressible pipe connected to a very compliant lung/alveolar decompression sickness (DCS; see Glossary). In this Review, we space (balloon-pipe model; Fig. 1A; Bostrom et al., 2008), allowing focus on the link between form and function in the respiratory the alveolar collapse depth to be estimated from Boyle’slaw systems of diving marine mammals, but emphasize studies that have (Scholander, 1940; Bostrom et al., 2008). This has been an attempted to understand lung function and mechanics in pinnipeds important assumption used to understand diving physiology and and cetaceans (the species where the majority of work has been how marine mammals avoid diving-related problems, such as DCS done). Scholander’s model of lung/alveolar collapse (see below) is and N2 narcosis, by reducing N2 uptake and blood and tissue N2 of particular interest to this Review; this model provides a tension. However, many aspects of marine mammal respiratory mechanism for how marine mammals avoid lung squeeze (see physiology are still not well understood; therefore, in this Review, Glossary), limit their uptake of N2, avoid inert gas narcosis (see we summarize past and recent studies with the aim of providing Glossary) and DCS, and are able to generate high respiratory flows some generalizations about the different traits that have evolved to that are sustained over the entire vital capacity (VC; see Glossary). allow marine mammals to manage a life in the ocean. Static respiratory variables 1Fundación Oceanográfic de la Comunidad Valenciana, Gran Vıá Marques del Turia 19, Valencia 46005, Spain. 2Department of Life Sciences, Texas A&M Static indices of respiratory capacity are those that do not change University-Corpus Christi, 6300 Ocean Drive, Corpus Christi, TX 78412, USA. between breaths. Data on these variables exist for a limited 3Biology Department, Woods Hole Oceanographic Institution, Woods Hole, MA 02543, USA. 4Oceanográfic-Avanqua, Ciudad de las Artes y las Ciencias, Valencia number of marine mammal species and include information on the 46013, Spain. total lung capacity (TLC) and minimum air volume (MAV; see Glossary) of the relaxed lung, as discussed below (Kooyman, *Author for correspondence ([email protected]) 1973; Kooyman and Sinnett, 1979; Piscitelli et al., 2010; Fahlman A.F., 0000-0002-8675-6479 et al., 2011). Journal of Experimental Biology 1761 REVIEW Journal of Experimental Biology (2017) 220, 1761-1773 doi:10.1242/jeb.126870 4 A V A Glossary V DS Alveolar recruitment 3 V A V The point when collapsed alveoli open up and gas exchange resumes. DS Atelectasis V V 2 AI 252 m A=0 ml, DS=420 ml Alveolar collapse, resulting in cessation of gas exchange. Collateral ventilation (I) Volume 1 Ventilatory flow through the lung parenchyma through alternative flow pathways, such as pores of Kohn. 0 Dead space 02040 60 80 100 120 The volume of air in the respiratory system not participating in gas Depth (m) exchange, e.g. air in the trachea. Decompression sickness Also called the ‘bends’ or caisson disease; a collection of symptoms Trachea observed following a reduction in ambient pressure, which causes bubbles to form in the blood and tissues. In humans, symptoms include Balloon–pipe model Alveoli dizziness, numbness, fatigue and, in more severe cases, paralysis, problems breathing and death. Inert gas narcosis Caused by the anesthetic effect of lipid-soluble gases at high pressure. Rigid but compressible In air-breathing divers the symptoms may ultimately lead to loss of trachea consciousness as pressure increases. Increasing pressure Lung squeeze Pulmonary edema caused by intrathoracic pressures that are lower than BC environmental pressures during breath-hold diving. Maximal/forced breath Often used in human lung-function testing to assess the maximal capacity of lung function such as VC, PEF, PIF and airway obstruction. The individual is asked to expire maximally, followed by an inspiration. Minimum air volume The volume of air left in the relaxed lung. Respiratory frequency 1 cm The number of breaths per unit time. Rete A network of arteries or veins D ) 50 Tidal volume A ϫ The volume of air exhaled or inhaled during a normal breath. P 40 Vital capacity The maximal volume of air that can be exchanged in one breath. In 30 marine mammals, VC is close to TLC. 20 Alveolar 10 collapse TLC or lung size Compliant trachea Overall, maximal lung volumes, or TLC (generally defined as the Rigid trachea Relative diffusion rate ( Relative diffusion 0 volume of air in the lung when the transpulmonary pressure is 123456789101112 30 cmH O, where 1 cmH O≈98 Pa), of diving mammals are in the 2 2 Pressure (ATA) general range of those of terrestrial mammals (Kooyman, 1973; Fahlman et al., 2011; Piscitelli et al., 2013; Ponganis, 2015). Fig. 1. The effect of pressure on lung volume and diffusion rate. (A) Graph Exceptions are the smaller lungs of deep-diving cetaceans and the showing how the compression of the respiratory system is affected when the enlarged lungs of shallow-diving species, e.g. sea otters compliance of the upper and lower airways are accounted for. The figure (Scholander, 1940; Kooyman, 1973; Leith, 1989; Piscitelli et al., assumes a Weddell seal with a diving lung volume of 11 liters, an alveolar volume (VA, solid lines) of 10 liters and a dead space volume (VDS,broken 2013; Ponganis, 2015). While the classification of deep and shallow lines) of 1 liter. The black lines represent the volume of the respiratory system in divers is not well defined, and changes as we find out more about the relation to depth for Scholander’s original balloon-pipe model, with a stiff dead life history of different species, one study defined a shallow diver to space that does not compress, and the red lines represent the volume of the be a species where most dives are shallower than 100 m, e.g. lung based on the lung compression model presented in Bostrom et al. (2008). bottlenose dolphin (Tursiops truncatus) and harbor porpoise The schematic below panel A provides a qualitative explanation of the two (Phocoena phocoena) (Piscitelli et al., 2010).
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