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<span>Toggle Composition subsection</span> </button> <ul id="toc-Composition-sublist" class="vector-toc-list"> <li id="toc-Lipids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lipids"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Lipids</span> </div> </a> <ul id="toc-Lipids-sublist" class="vector-toc-list"> <li id="toc-DPPC" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#DPPC"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>DPPC</span> </div> </a> <ul id="toc-DPPC-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_phospholipids" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Other_phospholipids"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.2</span> <span>Other phospholipids</span> </div> </a> <ul id="toc-Other_phospholipids-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Proteins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Proteins</span> </div> </a> <ul id="toc-Proteins-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Artificial_surfactants" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Artificial_surfactants"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Artificial surfactants</span> </div> </a> <ul id="toc-Artificial_surfactants-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Surface_tension_magnitude_inside_the_lung" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Surface_tension_magnitude_inside_the_lung"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Surface tension magnitude inside the lung</span> </div> </a> <ul id="toc-Surface_tension_magnitude_inside_the_lung-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Production_and_degradation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Production_and_degradation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Production and degradation</span> </div> </a> <ul id="toc-Production_and_degradation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Diseases" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Diseases"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Diseases</span> </div> </a> <ul id="toc-Diseases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav 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class="firstHeading mw-first-heading"><span class="mw-page-title-main">Pulmonary surfactant</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 18 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-18" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">18 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%81%D8%A7%D8%B9%D9%84_%D8%A8%D8%A7%D9%84%D8%B3%D8%B7%D8%AD_%D8%A7%D9%84%D8%B1%D8%A6%D9%88%D9%8A" title="فاعل بالسطح الرئوي – Arabic" lang="ar" hreflang="ar" data-title="فاعل بالسطح الرئوي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Surfactant_pulmonar" title="Surfactant pulmonar – Catalan" lang="ca" hreflang="ca" data-title="Surfactant pulmonar" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Surfactant" title="Surfactant – German" lang="de" hreflang="de" data-title="Surfactant" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Surfactante_pulmonar" title="Surfactante pulmonar – Spanish" lang="es" hreflang="es" data-title="Surfactante pulmonar" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D9%88%D8%B1%D9%81%DA%A9%D8%AA%D8%A7%D9%86%D8%AA_%D8%B1%DB%8C%D9%88%DB%8C" title="سورفکتانت ریوی – Persian" lang="fa" hreflang="fa" data-title="سورفکتانت ریوی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Surfactant_pulmonaire" title="Surfactant pulmonaire – French" lang="fr" hreflang="fr" data-title="Surfactant pulmonaire" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Surfactante_pulmonar" title="Surfactante pulmonar – Galician" lang="gl" hreflang="gl" data-title="Surfactante pulmonar" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Surfaktan_paru" title="Surfaktan paru – Indonesian" lang="id" hreflang="id" data-title="Surfaktan paru" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Surfattante_polmonare" title="Surfattante polmonare – Italian" lang="it" hreflang="it" data-title="Surfattante polmonare" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%A1%D1%83%D1%80%D1%84%D0%B0%D0%BA%D1%82%D0%B0%D0%BD%D1%82" title="Сурфактант – Kazakh" lang="kk" hreflang="kk" data-title="Сурфактант" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B6%E0%B5%8D%E0%B4%B5%E0%B4%BE%E0%B4%B8%E0%B4%95%E0%B5%8B%E0%B4%B6%E0%B4%99%E0%B5%8D%E0%B4%99%E0%B4%B3%E0%B4%BF%E0%B4%B2%E0%B5%86_%E0%B4%B8%E0%B5%BC%E0%B4%AB%E0%B4%95%E0%B5%8D%E0%B4%B1%E0%B5%8D%E0%B4%B1%E0%B4%A8%E0%B5%8D%E0%B4%B1%E0%B5%8D" title="ശ്വാസകോശങ്ങളിലെ സർഫക്റ്റന്റ് – Malayalam" lang="ml" hreflang="ml" data-title="ശ്വാസകോശങ്ങളിലെ സർഫക്റ്റന്റ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E8%82%BA%E3%82%B5%E3%83%BC%E3%83%95%E3%82%A1%E3%82%AF%E3%82%BF%E3%83%B3%E3%83%88" title="肺サーファクタント – Japanese" lang="ja" hreflang="ja" data-title="肺サーファクタント" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Surfactante_pulmonar" title="Surfactante pulmonar – Portuguese" lang="pt" hreflang="pt" data-title="Surfactante pulmonar" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9B%D1%91%D0%B3%D0%BE%D1%87%D0%BD%D1%8B%D0%B9_%D1%81%D1%83%D1%80%D1%84%D0%B0%D0%BA%D1%82%D0%B0%D0%BD%D1%82" title="Лёгочный сурфактант – Russian" lang="ru" hreflang="ru" data-title="Лёгочный сурфактант" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Plu%C4%87ni_surfaktant" title="Plućni surfaktant – Serbian" lang="sr" hreflang="sr" data-title="Plućni surfaktant" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9B%D0%B5%D0%B3%D0%B5%D0%BD%D0%B5%D0%B2%D1%96_%D1%81%D1%83%D1%80%D1%84%D0%B0%D0%BA%D1%82%D0%B0%D0%BD%D1%82%D0%B8" title="Легеневі сурфактанти – Ukrainian" lang="uk" hreflang="uk" data-title="Легеневі сурфактанти" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Ch%E1%BA%A5t_ho%E1%BA%A1t_di%E1%BB%87n_ph%E1%BB%95i" title="Chất hoạt diện phổi – Vietnamese" lang="vi" hreflang="vi" data-title="Chất hoạt diện phổi" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%82%BA%E8%A1%A8%E9%9D%A2%E6%B4%BB%E6%80%A7%E7%89%A9%E8%B4%A8" title="肺表面活性物质 – Chinese" lang="zh" hreflang="zh" data-title="肺表面活性物质" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q1324199#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs 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href="/wiki/File:An_alveolus,_is_an_anatomical_structure_that_has_the_form_of_a_hollow_cavity._Mainly_found_in_the_lung,_the_pulmonary_alveoli_are_spherical_outcroppings_of_the_respiratory_bronchioles_and_are_the.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/An_alveolus%2C_is_an_anatomical_structure_that_has_the_form_of_a_hollow_cavity._Mainly_found_in_the_lung%2C_the_pulmonary_alveoli_are_spherical_outcroppings_of_the_respiratory_bronchioles_and_are_the.png/300px-thumbnail.png" decoding="async" width="300" height="224" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/An_alveolus%2C_is_an_anatomical_structure_that_has_the_form_of_a_hollow_cavity._Mainly_found_in_the_lung%2C_the_pulmonary_alveoli_are_spherical_outcroppings_of_the_respiratory_bronchioles_and_are_the.png/450px-thumbnail.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/77/An_alveolus%2C_is_an_anatomical_structure_that_has_the_form_of_a_hollow_cavity._Mainly_found_in_the_lung%2C_the_pulmonary_alveoli_are_spherical_outcroppings_of_the_respiratory_bronchioles_and_are_the.png/600px-thumbnail.png 2x" data-file-width="2000" data-file-height="1493" /></a><figcaption><a href="/wiki/Pulmonary_alveolus" title="Pulmonary alveolus">Alveoli</a> are the spherical outcroppings of the <a href="/wiki/Respiratory_bronchioles" class="mw-redirect" title="Respiratory bronchioles">respiratory bronchioles</a>.</figcaption></figure> <p><b>Pulmonary surfactant</b> is a surface-active complex of <a href="/wiki/Phospholipid" title="Phospholipid">phospholipids</a> and <a href="/wiki/Protein" title="Protein">proteins</a> formed by <a href="/wiki/Type_II_cells" class="mw-redirect" title="Type II cells">type II alveolar cells</a>.<sup id="cite_ref-Bernhard_1-0" class="reference"><a href="#cite_note-Bernhard-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The proteins and lipids that make up the <a href="/wiki/Surfactant" title="Surfactant">surfactant</a> have both <a href="/wiki/Hydrophilic" class="mw-redirect" title="Hydrophilic">hydrophilic</a> and <a href="/wiki/Hydrophobic" class="mw-redirect" title="Hydrophobic">hydrophobic</a> regions. By <a href="/wiki/Adsorption" title="Adsorption">adsorbing</a> to the air-water <a href="/wiki/Interface_(chemistry)" class="mw-redirect" title="Interface (chemistry)">interface</a> of <a href="/wiki/Pulmonary_alveolus" title="Pulmonary alveolus">alveoli</a>, with hydrophilic head groups in the water and the hydrophobic tails facing towards the air, the main lipid component of surfactant, <a href="/wiki/Dipalmitoylphosphatidylcholine" title="Dipalmitoylphosphatidylcholine">dipalmitoylphosphatidylcholine</a> (DPPC), reduces <a href="/wiki/Surface_tension" title="Surface tension">surface tension</a>. </p><p>As a medication, <a href="/wiki/Pulmonary_surfactant_(medication)" title="Pulmonary surfactant (medication)">pulmonary surfactant</a> is on the <a href="/wiki/WHO_Model_List_of_Essential_Medicines" title="WHO Model List of Essential Medicines">WHO Model List of Essential Medicines</a>, the most important medications needed in a basic <a href="/wiki/Health_system" title="Health system">health system</a>.<sup id="cite_ref-WHO2015E_2-0" class="reference"><a href="#cite_note-WHO2015E-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=1" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>To increase <a href="/wiki/Pulmonary_compliance" class="mw-redirect" title="Pulmonary compliance">pulmonary compliance</a>.</li> <li>To prevent <a href="/wiki/Atelectasis" title="Atelectasis">atelectasis</a> (collapse of the alveoli or atriums) at the end of expiration.</li> <li>To facilitate recruitment of collapsed airways.</li></ul> <p>Alveoli can be compared to gas in water, as the alveoli are wet and surround a central air space. The <a href="/wiki/Surface_tension" title="Surface tension">surface tension</a> acts at the air-water interface and tends to make the bubble smaller (by decreasing the surface area of the interface). The gas pressure (<i>P</i>) needed to keep an equilibrium between the collapsing force of <a href="/wiki/Surface_tension" title="Surface tension">surface tension</a> (<i>γ</i>) and the expanding force of gas in an alveolus of radius <i>r</i> is expressed by the <a href="/wiki/Young%E2%80%93Laplace_equation" title="Young–Laplace equation">Young–Laplace equation</a>: </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle P={\frac {2\gamma }{r}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>P</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mn>2</mn> <mi>γ</mi> </mrow> <mi>r</mi> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle P={\frac {2\gamma }{r}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/7b10f1ed1d9a8c80c52e0e6e8887cf6532286a92" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.838ex; width:8.105ex; height:5.343ex;" alt="{\displaystyle P={\frac {2\gamma }{r}}}"></span></dd></dl> <div class="mw-heading mw-heading3"><h3 id="Compliance">Compliance</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=2" title="Edit section: Compliance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Compliance_(physiology)" title="Compliance (physiology)">Compliance</a> is the ability of <a href="/wiki/Lung" title="Lung">lungs</a> and <a href="/wiki/Thorax" title="Thorax">thorax</a> to expand. Lung compliance is defined as the volume change per unit of pressure change across the lung. Measurements of lung volume obtained during the controlled inflation/deflation of a normal lung show that the volumes obtained during deflation exceed those during inflation, at a given pressure. This difference in inflation and deflation volumes at a given pressure is called <a href="/wiki/Hysteresis#Respiratory_physiology" title="Hysteresis">hysteresis</a> and is due to the air-water surface tension that occurs at the beginning of inflation. However, surfactant decreases the alveolar <a href="/wiki/Surface_tension" title="Surface tension">surface tension</a>, as seen in cases of premature infants with <a href="/wiki/Infant_respiratory_distress_syndrome" title="Infant respiratory distress syndrome">infant respiratory distress syndrome</a>. The normal surface tension for water is 70 dyn/cm (70 mN/m) and in the lungs, it is 25 dyn/cm (25 mN/m); however, at the end of the expiration, compressed surfactant phospholipid molecules decrease the surface tension to very low, near-zero levels. Pulmonary surfactant thus greatly reduces <a href="/wiki/Surface_tension" title="Surface tension">surface tension</a>, increasing compliance allowing the lung to inflate much more easily, thereby reducing the work of breathing. It reduces the pressure difference needed to allow the lung to inflate. The lung's compliance, and ventilation decrease when <a href="/wiki/Lung_parenchyma" class="mw-redirect" title="Lung parenchyma">lung tissue</a> becomes diseased and <a href="/wiki/Fibrosis" title="Fibrosis">fibrotic</a>.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Alveolar_size_regulation">Alveolar size regulation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=3" title="Edit section: Alveolar size regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As the alveoli increase in size, the surfactant becomes more spread out over the surface of the liquid. This increases surface tension effectively slowing the rate of expansion of the alveoli. This also helps all alveoli in the lungs expand at the same rate, as one that expands more quickly will experience a large rise in surface tension slowing its rate of expansion. It also means the rate of shrinking is more regular as if one reduces in size more quickly the surface tension will reduce more, so other alveoli can contract more easily than it can. Surfactant reduces surface tension more readily when the alveoli are smaller because the surfactant is more concentrated. </p> <div class="mw-heading mw-heading3"><h3 id="Prevention_of_fluid_accumulation_and_maintenance_of_dryness_of_airways">Prevention of fluid accumulation and maintenance of dryness of airways</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=4" title="Edit section: Prevention of fluid accumulation and maintenance of dryness of airways"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Surface tension draws fluid from capillaries to the alveolar spaces. Surfactant reduces fluid accumulation and keeps the airways dry by reducing surface tension.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Innate_immunity">Innate immunity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=5" title="Edit section: Innate immunity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Surfactant immune function is primarily attributed to two proteins: <a href="/wiki/Surfactant_protein_A" title="Surfactant protein A">SP-A</a> and <a href="/wiki/Surfactant_protein_D" title="Surfactant protein D">SP-D</a>. These proteins can bind to sugars on the surface of pathogens and thereby <a href="/wiki/Opsonize" class="mw-redirect" title="Opsonize">opsonize</a> them for uptake by phagocytes. It also regulates inflammatory responses and interacts with the adaptive immune response. Surfactant degradation or inactivation may contribute to enhanced susceptibility to lung inflammation and infection.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Composition">Composition</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=6" title="Edit section: Composition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>~40% <a href="/wiki/Dipalmitoylphosphatidylcholine" title="Dipalmitoylphosphatidylcholine">dipalmitoylphosphatidylcholine</a> (DPPC);<sup id="cite_ref-NkadiMerritt2009_6-0" class="reference"><a href="#cite_note-NkadiMerritt2009-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li> <li>~40% other <a href="/wiki/Phospholipid" title="Phospholipid">phospholipids</a> (PC);<sup id="cite_ref-NkadiMerritt2009_6-1" class="reference"><a href="#cite_note-NkadiMerritt2009-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li> <li>~10% surfactant <a href="/wiki/Proteins" class="mw-redirect" title="Proteins">proteins</a> (<a href="/wiki/Surfactant_protein_A" title="Surfactant protein A">SP-A</a>, <a href="/wiki/Pulmonary_surfactant-associated_protein_B" class="mw-redirect" title="Pulmonary surfactant-associated protein B">SP-B</a>, <a href="/wiki/Surfactant_protein_C" title="Surfactant protein C">SP-C</a> and <a href="/wiki/Surfactant_protein_D" title="Surfactant protein D">SP-D</a>);<sup id="cite_ref-NkadiMerritt2009_6-2" class="reference"><a href="#cite_note-NkadiMerritt2009-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li> <li>~10% neutral <a href="/wiki/Lipids" class="mw-redirect" title="Lipids">lipids</a> (<a href="/wiki/Cholesterol" title="Cholesterol">Cholesterol</a>);<sup id="cite_ref-NkadiMerritt2009_6-3" class="reference"><a href="#cite_note-NkadiMerritt2009-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li> <li>Traces of other <a href="/wiki/Chemical_substance" title="Chemical substance">substances</a>.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Lipids">Lipids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=7" title="Edit section: Lipids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="DPPC">DPPC</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=8" title="Edit section: DPPC"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Dipalmitoylphosphatidylcholine" title="Dipalmitoylphosphatidylcholine">Dipalmitoylphosphatidylcholine</a> (DPPC) is a <a href="/wiki/Phospholipid" title="Phospholipid">phospholipid</a> with two 16-carbon saturated chains and a <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> group with quaternary amine group attached. The DPPC is the strongest surfactant molecule in the pulmonary surfactant mixture. It also has a higher compaction capacity than the other phospholipids, because the apolar tail is less bent. Nevertheless, without the other substances of the pulmonary surfactant mixture, the DPPC's <a href="/wiki/Adsorption" title="Adsorption">adsorption</a> kinetics is very slow. This happens primarily because the phase <a href="/wiki/Transition_temperature" title="Transition temperature">transition temperature</a> between <a href="/wiki/Gel" title="Gel">gel</a> to <a href="/wiki/Liquid_crystal" title="Liquid crystal">liquid crystal</a> of pure DPPC is 41.5 °C, which is higher than the <a href="/wiki/Human_body_temperature" title="Human body temperature">human body's temperature</a> of 37 °C.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Other_phospholipids">Other phospholipids</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=9" title="Edit section: Other phospholipids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Phosphatidylcholine molecules form ~85% of the lipid in surfactant and have saturated acyl chains. Phosphatidylglycerol (PG) forms about 11% of the lipids in the surfactant, it has unsaturated fatty acid chains that fluidize the lipid monolayer at the interface. Neutral lipids and cholesterol are also present. The components for these lipids diffuse from the blood into type II alveolar cells where they are assembled and packaged for secretion into secretory organelles called <a href="/wiki/Lamellar_bodies" title="Lamellar bodies">lamellar bodies</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2015)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Proteins">Proteins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=10" title="Edit section: Proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Proteins" class="mw-redirect" title="Proteins">Proteins</a> make up the remaining 10% of the surfactant. Half of this 10% is <a href="/wiki/Plasma_proteins" class="mw-redirect" title="Plasma proteins">plasma proteins</a> but the rest is formed by the <a href="/wiki/Apolipoprotein" title="Apolipoprotein">apolipoproteins</a>, surfactant proteins SP-A, SP-B, SP-C, and SP-D. The apolipoproteins are produced by the secretory pathway in type II cells. They undergo much post-translational modification, ending up in the lamellar bodies. These are concentric rings of lipid and protein, about 1 μm in diameter. </p> <ul><li><a href="/wiki/Surfactant_protein_A" title="Surfactant protein A">SP-A</a> and <a href="/wiki/Surfactant_protein_D" title="Surfactant protein D">SP-D</a> are <a href="/wiki/Collectin" title="Collectin">collectins</a>. They confer innate immunity as they have carbohydrate recognition domains that allow them to coat bacteria and viruses promoting phagocytosis by macrophages.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> SP-A is also thought to be involved in a negative feedback mechanism to control the production of surfactant.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2015)">citation needed</span></a></i>]</sup></li> <li><a href="/wiki/Surfactant_protein_B" title="Surfactant protein B">SP-B</a> and <a href="/wiki/Surfactant_protein_C" title="Surfactant protein C">SP-C</a> are hydrophobic membrane proteins that increase the rate that surfactant spreads over the surface. SP-B and SP-C are required for the proper biophysical function of the lung. Humans and animals born with a congenital absence of the <a href="/wiki/Saposin_protein_domain" title="Saposin protein domain">saposin family</a> SP-B experience intractable respiratory failure whereas those born lacking SP-C tend to develop progressive interstitial pneumonitis.<sup id="cite_ref-pmid11181967_9-0" class="reference"><a href="#cite_note-pmid11181967-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup></li></ul> <p>The SP proteins reduce the critical temperature of DPPC's <a href="/wiki/Phase_transition" title="Phase transition">phase transition</a> to a value lower than 37 °C,<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> which improves its adsorption and interface spreading velocity.<sup id="cite_ref-Schuerch_1992_11-0" class="reference"><a href="#cite_note-Schuerch_1992-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Possmayer_12-0" class="reference"><a href="#cite_note-Possmayer-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The compression of the interface causes a phase change of the surfactant molecules to liquid-gel or even gel-solid. The fast adsorption velocity is necessary to maintain the integrity of the gas exchange region of the lungs. </p><p>Each SP protein has distinct functions, which act <a href="/wiki/Synergy" title="Synergy">synergistically</a> to keep an interface rich in DPPC during lung's expansion and contraction. Changes in the surfactant mixture composition alter the pressure and temperature conditions for phase changes and the phospholipids' crystal shape as well.<sup id="cite_ref-Veldhuizena_13-0" class="reference"><a href="#cite_note-Veldhuizena-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Only the liquid phase can freely spread on the surface to form a monolayer. Nevertheless, it has been observed that if a lung region is abruptly expanded the floating crystals crack like "<a href="/wiki/Icebergs" class="mw-redirect" title="Icebergs">icebergs</a>". Then the SP proteins selectively attract more DPPC to the interface than other phospholipids or cholesterol, whose surfactant properties are worse than DPPC's. The SP also fastens the DPPC on the interface to prevent the DPPC from being squeezed out when the surface area decreases <sup id="cite_ref-Possmayer_12-1" class="reference"><a href="#cite_note-Possmayer-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> This also reduces the interface compressibility.<sup id="cite_ref-Schuerch_2001_14-0" class="reference"><a href="#cite_note-Schuerch_2001-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Artificial_surfactants">Artificial surfactants</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=11" title="Edit section: Artificial surfactants"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Pulmonary_surfactant_(medication)" title="Pulmonary surfactant (medication)">Pulmonary surfactant (medication)</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Pulmonary_surfactant.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ea/Pulmonary_surfactant.JPG/200px-Pulmonary_surfactant.JPG" decoding="async" width="200" height="267" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ea/Pulmonary_surfactant.JPG/300px-Pulmonary_surfactant.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ea/Pulmonary_surfactant.JPG/400px-Pulmonary_surfactant.JPG 2x" data-file-width="2604" data-file-height="3472" /></a><figcaption><a href="/wiki/Survanta" class="mw-redirect" title="Survanta">Survanta</a>, surrounded by devices for its application.</figcaption></figure> <p>There are a number of types of pulmonary surfactants available. Ex-situ measurements of <a href="/wiki/Surface_tension" title="Surface tension">surface tension</a> and <a href="/wiki/Interfacial_rheology" title="Interfacial rheology">interfacial rheology</a> can help to understand the functionality of pulmonary surfactants.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Synthetic pulmonary surfactants </p> <ol><li><a href="/wiki/Colfosceril_palmitate" title="Colfosceril palmitate">Colfosceril palmitate</a> (Exosurf) - a mixture of DPPC with hexadecanol and tyloxapol added as spreading agents</li> <li><a href="/wiki/Pumactant" class="mw-redirect" title="Pumactant">Pumactant</a> (Artificial Lung Expanding Compound or ALEC) - a mixture of DPPC and PG</li> <li><a href="/wiki/KL-4" class="mw-redirect" title="KL-4">KL-4</a> - composed of DPPC, palmitoyl-oleoyl phosphatidylglycerol, and palmitic acid, combined with a 21 amino acid synthetic peptide that mimics the structural characteristics of SP-B.</li> <li><a href="/wiki/Venticute" class="mw-redirect" title="Venticute">Venticute</a> - DPPC, PG, palmitic acid and recombinant SP-C</li> <li><a href="/wiki/Lucinactant" title="Lucinactant">Lucinactant</a> - <a href="/wiki/Dipalmitoylphosphatidylcholine" title="Dipalmitoylphosphatidylcholine">DPPC</a>, <a href="/w/index.php?title=1-palmitoyl-2-oleoyl-sn-glycero-3-phosphoglycerol&action=edit&redlink=1" class="new" title="1-palmitoyl-2-oleoyl-sn-glycero-3-phosphoglycerol (page does not exist)">POPG</a>, and <a href="/wiki/Palmitic_acid" title="Palmitic acid">palmitic acid</a>.</li></ol> <p>Animal derived surfactants </p> <ol><li><a href="/wiki/Beractant" title="Beractant">Beractant</a> <ol><li>(Alveofact) - extracted from cow lung lavage fluid</li> <li>(Survanta) - extracted from minced cow lung with additional DPPC, palmitic acid, and tripalmitin</li> <li>(Beraksurf) -extracted from minced calf lung with additional DPPC, palmitic acid, and tripalmitin</li></ol></li> <li><a href="/wiki/Calfactant" title="Calfactant">Calfactant</a> (Infasurf) - extracted from calf lung lavage fluid</li> <li><a href="/wiki/Poractant_alfa" title="Poractant alfa">Poractant alfa</a> (Curosurf) - extracted from material derived from minced pig lung</li> <li><a href="/w/index.php?title=Ovinactant&action=edit&redlink=1" class="new" title="Ovinactant (page does not exist)">Ovinactant</a> (Varasurf) - extracted from material derived from minced sheep lung</li></ol> <div class="mw-heading mw-heading2"><h2 id="Surface_tension_magnitude_inside_the_lung">Surface tension magnitude inside the lung</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=12" title="Edit section: Surface tension magnitude inside the lung"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Even though the surface tension can be greatly reduced by pulmonary surfactant, this effect will depend on the surfactant's concentration on the interface. The interface concentration has a saturation limit, which depends on temperature and mixture composition. Because during ventilation there is a variation of the lung surface area, the surfactant's interface concentration is not usually at the level of saturation. The surface increases during inspiration, which consequently opens space for new surfactant molecules to be recruited to the interface. Meanwhile, during expiration the surface area decreases at a rate which is always in excess of the rate at which the surfactant molecules are driven from the interface into the water film. Thus, the surfactant density at the air water interface remains high and is relatively preserved throughout expiration, decreasing the surface tension even further. This also explains why the compliance is greater during expiration than during inspiration. <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2015)">citation needed</span></a></i>]</sup> </p><p>SP molecules contribute to increasing the surfactant interface adsorption kinetics, when the concentration is below the <a href="/wiki/Solubility" title="Solubility">saturation</a> level. They also make <a href="/wiki/Van_der_Waals_Forces" class="mw-redirect" title="Van der Waals Forces">weak bonds</a> with the surfactant molecules at the interface and hold them longer there when the interface is compressed. Therefore, during ventilation, surface tension is usually lower than at equilibrium. Therefore, the surface tension varies according to the volume of air in the lungs, which protects them from atelectasis at low volumes and tissue damage at high volume levels.<sup id="cite_ref-Schuerch_1992_11-1" class="reference"><a href="#cite_note-Schuerch_1992-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Veldhuizena_13-1" class="reference"><a href="#cite_note-Veldhuizena-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Schuerch_2001_14-1" class="reference"><a href="#cite_note-Schuerch_2001-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <table class="wikitable" style="text-align:center;"> <caption>Surface tension values </caption> <tbody><tr> <th>Condition </th> <th>Tension (mN/m) </th></tr> <tr> <td style="text-align:left;">Water at 25 °C </td> <td>70 </td></tr> <tr> <td style="text-align:left;">Pulmonary surfactant in equilibrium at 36 °C </td> <td>25 </td></tr> <tr> <td style="text-align:left;">Healthy lung at 100% of <a href="/wiki/Total_lung_capacity" class="mw-redirect" title="Total lung capacity">TLC</a> </td> <td>30 </td></tr> <tr> <td style="text-align:left;">Healthy lung between 40 and 60% of <a href="/wiki/Total_lung_capacity" class="mw-redirect" title="Total lung capacity">TLC</a> </td> <td>1~6 </td></tr> <tr> <td style="text-align:left;">Healthy lung below 40% of <a href="/wiki/Total_lung_capacity" class="mw-redirect" title="Total lung capacity">TLC</a> </td> <td><1 </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Production_and_degradation">Production and degradation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=13" title="Edit section: Production and degradation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Surfactant production in humans begins in <a href="/wiki/Type_II_cells" class="mw-redirect" title="Type II cells">type II cells</a> during the <a href="/wiki/Pulmonary_alveolus" title="Pulmonary alveolus">alveolar sac</a> stage of lung development. <a href="/wiki/Lamellar_bodies" title="Lamellar bodies">Lamellar bodies</a> appear in the cytoplasm at about 20 weeks gestation.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> These lamellar bodies are secreted by <a href="/wiki/Exocytosis" title="Exocytosis">exocytosis</a> into the <a href="/wiki/Epithelial_lining_fluid" class="mw-redirect" title="Epithelial lining fluid">alveolar lining fluid</a>, where the surfactant forms a meshwork of <b>tubular myelin</b><sup id="cite_ref-Young_17-0" class="reference"><a href="#cite_note-Young-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Franciosi_18-0" class="reference"><a href="#cite_note-Franciosi-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Full_term" class="mw-redirect" title="Full term">Full term</a> infants are estimated to have an alveolar storage pool of approximately 100 mg/kg of surfactant, while preterm infants have an estimated 4–5 mg/kg at birth.<sup id="cite_ref-Niemarkt_Kramer_p._19-0" class="reference"><a href="#cite_note-Niemarkt_Kramer_p.-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Club_cell" title="Club cell">Club cells</a> also produce a component of lung surfactant.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>Alveolar surfactant has a half-life of 5 to 10 hours once secreted. It can be both broken down by macrophages and/or reabsorbed into the lamellar structures of type II pneumocytes. Up to 90% of surfactant DPPC (dipalmitoylphosphatidylcholine) is recycled from the alveolar space back into the type II pneumocyte. This process is believed to occur through <a href="/wiki/Surfactant_protein_A" title="Surfactant protein A">SP-A</a> stimulating receptor-mediated, <a href="/wiki/Clathrin" title="Clathrin">clathrin</a> dependent <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> The other 10% is taken up by alveolar macrophages and digested. </p> <div class="mw-heading mw-heading2"><h2 id="Diseases">Diseases</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=14" title="Edit section: Diseases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Infant_respiratory_distress_syndrome" title="Infant respiratory distress syndrome">Infant respiratory distress syndrome</a> (IRDS) is caused by lack of surfactant, commonly seen in premature babies born before 28–32 weeks of gestation. <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2015)">citation needed</span></a></i>]</sup></li> <li><a href="/wiki/Congenital_surfactant_deficiency" class="mw-redirect" title="Congenital surfactant deficiency">Congenital surfactant deficiency</a></li> <li><a href="/wiki/Pulmonary_alveolar_proteinosis" title="Pulmonary alveolar proteinosis">Pulmonary alveolar proteinosis</a></li> <li><a href="/wiki/Surfactant_metabolism_dysfunction" title="Surfactant metabolism dysfunction">Surfactant metabolism dysfunction</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=15" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In late 1920s von Neergaard<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> identified the function of the pulmonary surfactant in increasing the compliance of the lungs by reducing surface tension. However the significance of his discovery was not understood by the scientific and medical community at that time. He also realized the importance of having low surface tension in lungs of newborn infants. Later, in the middle of the 1950s, Pattle and <a href="/wiki/John_Allen_Clements" title="John Allen Clements">Clements</a> rediscovered the importance of surfactant and low surface tension in the lungs. At the end of that decade it was discovered that the lack of surfactant caused <a href="/wiki/Infant_respiratory_distress_syndrome" title="Infant respiratory distress syndrome">infant respiratory distress syndrome</a> (IRDS).<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Veldhuizena_13-2" class="reference"><a href="#cite_note-Veldhuizena-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=16" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Bernhard-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bernhard_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBernhard2016" class="citation journal cs1">Bernhard W (November 2016). "Lung surfactant: Function and composition in the context of development and respiratory physiology". <i>Annals of Anatomy</i>. <b>208</b>: 146–150. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.aanat.2016.08.003">10.1016/j.aanat.2016.08.003</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27693601">27693601</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+of+Anatomy&rft.atitle=Lung+surfactant%3A+Function+and+composition+in+the+context+of+development+and+respiratory+physiology.&rft.volume=208&rft.pages=146-150&rft.date=2016-11&rft_id=info%3Adoi%2F10.1016%2Fj.aanat.2016.08.003&rft_id=info%3Apmid%2F27693601&rft.aulast=Bernhard&rft.aufirst=W&rfr_id=info%3Asid%2Fen.wikipedia.org%3APulmonary+surfactant" class="Z3988"></span></span> </li> <li id="cite_note-WHO2015E-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-WHO2015E_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.who.int/medicines/publications/essentialmedicines/EML2015_8-May-15.pdf">"19th WHO Model List of Essential Medicines (April 2015)"</a> <span class="cs1-format">(PDF)</span>. WHO. April 2015<span class="reference-accessdate">. Retrieved <span class="nowrap">May 10,</span> 2015</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=19th+WHO+Model+List+of+Essential+Medicines+%28April+2015%29&rft.pub=WHO&rft.date=2015-04&rft_id=https%3A%2F%2Fwww.who.int%2Fmedicines%2Fpublications%2Fessentialmedicines%2FEML2015_8-May-15.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APulmonary+surfactant" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20150518094937/http://www.medicalassistantonlineprograms.org/alveoli/">"Alveoli and the Breathing Process"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.medicalassistantonlineprograms.org/alveoli/">the original</a> on 2015-05-18<span class="reference-accessdate">. Retrieved <span class="nowrap">2013-10-30</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Alveoli+and+the+Breathing+Process&rft_id=http%3A%2F%2Fwww.medicalassistantonlineprograms.org%2Falveoli%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3APulmonary+surfactant" class="Z3988"></span><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Identifying_reliable_sources_(medicine)" title="Wikipedia:Identifying reliable sources (medicine)"><span title="Material near this tag may rely on an unreliable or less reliable medical source. (May 2015)">unreliable medical source?</span></a></i>]</sup></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWest,_John_B.1994" class="citation book cs1">West, John B. (1994). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/respiratoryphysi00west"><i>Respiratory physiology-- the essentials</i></a></span>. 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"Endocytic pathway for surfactant protein a in human macrophages: Binding, clathrin-mediated uptake, and trafficking through the endolysosomal pathway". <i>AJP: Lung Cellular and Molecular Physiology</i>. <b>290</b> (2): L334–42. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1152%2Fajplung.00267.2005">10.1152/ajplung.00267.2005</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16169899">16169899</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:12759872">12759872</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=AJP%3A+Lung+Cellular+and+Molecular+Physiology&rft.atitle=Endocytic+pathway+for+surfactant+protein+a+in+human+macrophages%3A+Binding%2C+clathrin-mediated+uptake%2C+and+trafficking+through+the+endolysosomal+pathway&rft.volume=290&rft.issue=2&rft.pages=L334-42&rft.date=2005&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A12759872%23id-name%3DS2CID&rft_id=info%3Apmid%2F16169899&rft_id=info%3Adoi%2F10.1152%2Fajplung.00267.2005&rft.aulast=Crowther&rft.aufirst=J.+E.&rft.au=Schlesinger%2C+L.+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APulmonary+surfactant" class="Z3988"></span></span> </li> <li id="cite_note-22"><span class="mw-cite-backlink"><b><a href="#cite_ref-22">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNeergaard1929" class="citation journal cs1 cs1-prop-foreign-lang-source">Neergaard K (1929). "Neue Auffassungen über einen Grundbegriff der Atemmechanik" [New views on a fundamental concept of respiratory mechanics]. <i>Zeitschrift für die Gesamte Experimentelle Medizin</i> (in German). <b>66</b> (1): 373–94. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fbf02621963">10.1007/bf02621963</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:87610289">87610289</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+die+Gesamte+Experimentelle+Medizin&rft.atitle=Neue+Auffassungen+%C3%BCber+einen+Grundbegriff+der+Atemmechanik&rft.volume=66&rft.issue=1&rft.pages=373-94&rft.date=1929&rft_id=info%3Adoi%2F10.1007%2Fbf02621963&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A87610289%23id-name%3DS2CID&rft.aulast=Neergaard&rft.aufirst=K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APulmonary+surfactant" class="Z3988"></span></span> </li> <li id="cite_note-23"><span class="mw-cite-backlink"><b><a href="#cite_ref-23">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAvery1959" class="citation journal cs1">Avery ME (1959-05-01). "Surface Properties in Relation to Atelectasis and Hyaline Membrane Disease". <i>Archives of Pediatrics & Adolescent Medicine</i>. <b>97</b> (5_PART_I): 517–23. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1001%2Farchpedi.1959.02070010519001">10.1001/archpedi.1959.02070010519001</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1072-4710">1072-4710</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/13649082">13649082</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:23200589">23200589</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archives+of+Pediatrics+%26+Adolescent+Medicine&rft.atitle=Surface+Properties+in+Relation+to+Atelectasis+and+Hyaline+Membrane+Disease&rft.volume=97&rft.issue=5_PART_I&rft.pages=517-23&rft.date=1959-05-01&rft.issn=1072-4710&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A23200589%23id-name%3DS2CID&rft_id=info%3Apmid%2F13649082&rft_id=info%3Adoi%2F10.1001%2Farchpedi.1959.02070010519001&rft.aulast=Avery&rft.aufirst=Mary+Ellen&rfr_id=info%3Asid%2Fen.wikipedia.org%3APulmonary+surfactant" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pulmonary_surfactant&action=edit&section=17" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> 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physiology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Respiration</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Breath" class="mw-redirect" title="Breath">breath</a> <ul><li><a href="/wiki/Inhalation" title="Inhalation">inhalation</a></li> <li><a href="/wiki/Exhalation" title="Exhalation">exhalation</a></li> <li><a href="/wiki/Obligate_nasal_breathing" title="Obligate nasal breathing">obligate nasal breathing</a></li></ul></li> <li><a href="/wiki/Respiratory_rate" title="Respiratory rate">respiratory rate</a></li> <li><a href="/wiki/Respirometer" title="Respirometer">respirometer</a></li> <li><a class="mw-selflink selflink">pulmonary surfactant</a></li> <li><a href="/wiki/Pulmonary_compliance" class="mw-redirect" title="Pulmonary compliance">compliance</a></li> <li><a href="/wiki/Elastic_recoil" title="Elastic recoil">elastic recoil</a></li> <li><a href="/wiki/Hysteresivity" title="Hysteresivity">hysteresivity</a></li> <li><a href="/wiki/Airway_resistance" title="Airway resistance">airway resistance</a></li> <li>bronchial <ul><li><a href="/wiki/Bronchial_hyperresponsiveness" title="Bronchial hyperresponsiveness">hyperresponsiveness</a></li> <li><a href="/wiki/Bronchoconstriction" title="Bronchoconstriction">constriction</a></li> <li><a href="/wiki/Bronchodilatation" title="Bronchodilatation">dilatation</a></li></ul></li> <li><a href="/wiki/Mechanical_ventilation" title="Mechanical ventilation">mechanical ventilation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Control_of_ventilation" title="Control of ventilation">Control</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Pons" title="Pons">pons</a></i> <ul><li><a href="/wiki/Pneumotaxic_center" class="mw-redirect" title="Pneumotaxic center">pneumotaxic center</a></li> <li><a href="/wiki/Apneustic_center" class="mw-redirect" title="Apneustic center">apneustic center</a></li></ul></li> <li><i><a href="/wiki/Medulla_oblongata" title="Medulla oblongata">medulla</a></i> <ul><li><a href="/wiki/Dorsal_respiratory_group" class="mw-redirect" title="Dorsal respiratory group">dorsal respiratory group</a></li> <li><a href="/wiki/Ventral_respiratory_group" class="mw-redirect" title="Ventral respiratory group">ventral respiratory group</a></li></ul></li> <li><i><a href="/wiki/Chemoreceptor" title="Chemoreceptor">chemoreceptors</a></i> <ul><li><a href="/wiki/Central_chemoreceptors" title="Central chemoreceptors">central</a></li> <li><a href="/wiki/Peripheral_chemoreceptors" title="Peripheral chemoreceptors">peripheral</a></li></ul></li> <li><a href="/wiki/Pulmonary_stretch_receptors" title="Pulmonary stretch receptors">pulmonary stretch receptors</a> <ul><li><a href="/wiki/Hering%E2%80%93Breuer_reflex" title="Hering–Breuer reflex">Hering–Breuer reflex</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lung_volumes" title="Lung volumes">Lung volumes</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Vital_capacity" title="Vital capacity">VC</a></li> <li><a href="/wiki/Functional_residual_capacity" title="Functional residual capacity">FRC</a></li> <li><a href="/wiki/Tidal_volume" title="Tidal volume">Vt</a></li> <li><a href="/wiki/Dead_space_(physiology)" title="Dead space (physiology)">dead space</a></li> <li><a href="/wiki/Closing_capacity" title="Closing capacity">CC</a></li> <li><a href="/wiki/Peak_expiratory_flow" title="Peak expiratory flow">PEF</a></li></ul> <dl><dt>calculations</dt> <dd></dd> <dd><a href="/wiki/Respiratory_minute_volume" class="mw-redirect" title="Respiratory minute volume">respiratory minute volume</a></dd> <dd><a href="/wiki/FEV1/FVC_ratio" title="FEV1/FVC ratio">FEV1/FVC ratio</a></dd></dl> <dl><dt><a href="/wiki/Lung_function_test" class="mw-redirect" title="Lung function test">Lung function tests</a></dt> <dd></dd> <dd><a href="/wiki/Spirometry" title="Spirometry">spirometry</a></dd> <dd><a href="/wiki/Body_plethysmography" class="mw-redirect" title="Body plethysmography">body plethysmography</a></dd> <dd><a href="/wiki/Peak_flow_meter" class="mw-redirect" title="Peak flow meter">peak flow meter</a></dd> <dd><a href="/wiki/Nitrogen_washout" title="Nitrogen washout">nitrogen washout</a></dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Circulation</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pulmonary_circulation" title="Pulmonary circulation">pulmonary circulation</a></li> <li><a href="/wiki/Hypoxic_pulmonary_vasoconstriction" title="Hypoxic pulmonary vasoconstriction">hypoxic pulmonary vasoconstriction</a></li> <li><a href="/wiki/Pulmonary_shunt" title="Pulmonary shunt">pulmonary shunt</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Interactions</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ventilation_(physiology)" class="mw-redirect" title="Ventilation (physiology)">ventilation (V)</a></li> <li><a href="/wiki/Perfusion" title="Perfusion">Perfusion</a> (Q) <ul><li><a href="/wiki/Ventilation/perfusion_ratio" title="Ventilation/perfusion ratio">Ventilation/perfusion ratio</a></li> <li><a href="/wiki/Ventilation/perfusion_scan" title="Ventilation/perfusion scan">V/Q scan</a></li></ul></li> <li><a href="/wiki/Zones_of_the_lung" title="Zones of the lung">zones of the lung</a></li> <li><a href="/wiki/Gas_exchange" title="Gas exchange">gas exchange</a></li> <li><a href="/wiki/Pulmonary_gas_pressures" title="Pulmonary gas pressures">pulmonary gas pressures</a></li> <li><a href="/wiki/Alveolar_gas_equation" title="Alveolar gas equation">alveolar gas equation</a></li> <li><a href="/wiki/Alveolar%E2%80%93arterial_gradient" title="Alveolar–arterial gradient">alveolar–arterial gradient</a></li> <li><a href="/wiki/Hemoglobin" title="Hemoglobin">hemoglobin</a></li> <li><a href="/wiki/Oxygen%E2%80%93hemoglobin_dissociation_curve" title="Oxygen–hemoglobin dissociation curve">oxygen–hemoglobin dissociation curve</a> (<a href="/wiki/Oxygen_saturation" title="Oxygen saturation">Oxygen saturation</a></li> <li><a href="/wiki/2,3-Bisphosphoglyceric_acid" title="2,3-Bisphosphoglyceric acid">2,3-BPG</a></li> <li><a href="/wiki/Bohr_effect" title="Bohr effect">Bohr effect</a></li> <li><a href="/wiki/Haldane_effect" title="Haldane effect">Haldane effect</a>)</li> <li><a href="/wiki/Carbonic_anhydrase" title="Carbonic anhydrase">carbonic anhydrase</a> (<a href="/wiki/Chloride_shift" title="Chloride shift">chloride shift</a>)</li> <li><a href="/wiki/Oxyhemoglobin" class="mw-redirect" title="Oxyhemoglobin">oxyhemoglobin</a></li> <li><a href="/wiki/Respiratory_quotient" title="Respiratory quotient">respiratory quotient</a></li> <li><a href="/wiki/Arterial_blood_gas" class="mw-redirect" title="Arterial blood gas">arterial blood gas</a></li> <li><a href="/wiki/Diffusion_capacity" class="mw-redirect" title="Diffusion capacity">diffusion capacity</a> (<a href="/wiki/DLCO" class="mw-redirect" title="DLCO">DLCO</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Insufficiency</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Effects_of_high_altitude_on_humans" title="Effects of high altitude on humans">high altitude</a> <ul><li><a href="/wiki/Death_zone" title="Death zone">death zone</a></li></ul></li> <li><a href="/wiki/Oxygen_toxicity" title="Oxygen toxicity">oxygen toxicity</a></li> <li><a href="/wiki/Hypoxia_(medical)" class="mw-redirect" title="Hypoxia (medical)">hypoxia</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Protein:_cell_membrane_proteins_(other_than_Cell_surface_receptor,_enzymes,_and_cytoskeleton)" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Other_cell_membrane_proteins" title="Template:Other cell membrane proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Other_cell_membrane_proteins" title="Template talk:Other cell membrane proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Other_cell_membrane_proteins" title="Special:EditPage/Template:Other cell membrane proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Protein:_cell_membrane_proteins_(other_than_Cell_surface_receptor,_enzymes,_and_cytoskeleton)" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein" title="Protein">Protein</a>: cell <a href="/wiki/Membrane_protein" title="Membrane protein">membrane proteins</a> (other than <a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">Cell surface receptor</a>, <a href="/wiki/Enzyme" title="Enzyme">enzymes</a>, and <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeleton</a>)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Arrestin" title="Arrestin">Arrestin</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/SAG_(gene)" title="SAG (gene)">SAG</a></li> <li><a href="/wiki/Arrestin_beta_1" title="Arrestin beta 1">ARRB1</a></li> <li><a href="/wiki/Arrestin_beta_2" title="Arrestin beta 2">ARRB2</a></li> <li><a href="/wiki/ARR3" title="ARR3">ARR3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Membrane-spanning_4A" title="Membrane-spanning 4A">Membrane-spanning 4A</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CD20" title="CD20">MS4A1</a></li> <li><a href="/wiki/MS4A2" title="MS4A2">MS4A2</a></li> <li><a href="/wiki/MS4A3" title="MS4A3">MS4A3</a></li> <li><a href="/w/index.php?title=MS4A4A&action=edit&redlink=1" class="new" title="MS4A4A (page does not exist)">MS4A4A</a></li> <li><a href="/w/index.php?title=MS4A4E&action=edit&redlink=1" class="new" title="MS4A4E (page does not exist)">MS4A4E</a></li> <li><a href="/w/index.php?title=MS4A5&action=edit&redlink=1" class="new" title="MS4A5 (page does not exist)">MS4A5</a></li> <li><a href="/w/index.php?title=MS4A6A&action=edit&redlink=1" class="new" title="MS4A6A (page does not exist)">MS4A6A</a></li> <li><a href="/w/index.php?title=MS4A6E&action=edit&redlink=1" class="new" title="MS4A6E (page does not exist)">MS4A6E</a></li> <li><a href="/wiki/MS4A7" title="MS4A7">MS4A7</a></li> <li><a href="/w/index.php?title=MS4A8B&action=edit&redlink=1" class="new" title="MS4A8B (page does not exist)">MS4A8B</a></li> <li><a href="/w/index.php?title=MS4A9&action=edit&redlink=1" class="new" title="MS4A9 (page does not exist)">MS4A9</a></li> <li><a href="/w/index.php?title=MS4A10&action=edit&redlink=1" class="new" title="MS4A10 (page does not exist)">MS4A10</a></li> <li><a href="/w/index.php?title=MS4A12&action=edit&redlink=1" class="new" title="MS4A12 (page does not exist)">MS4A12</a></li> <li><a href="/w/index.php?title=MS4A13&action=edit&redlink=1" class="new" title="MS4A13 (page does not exist)">MS4A13</a></li> <li><a href="/w/index.php?title=MS4A14&action=edit&redlink=1" class="new" title="MS4A14 (page does not exist)">MS4A14</a></li> <li><a href="/w/index.php?title=MS4A15&action=edit&redlink=1" class="new" title="MS4A15 (page does not exist)">MS4A15</a></li> <li><a href="/w/index.php?title=MS4A18&action=edit&redlink=1" class="new" title="MS4A18 (page does not exist)">MS4A18</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Myelin" title="Myelin">Myelin</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Myelin_basic_protein" title="Myelin basic protein">Myelin basic protein</a> <ul><li><a href="/wiki/PMP2" title="PMP2">PMP2</a></li></ul></li> <li><a href="/wiki/Myelin_proteolipid_protein" title="Myelin proteolipid protein">Myelin proteolipid protein</a> <ul><li><a href="/wiki/Proteolipid_protein_1" title="Proteolipid protein 1">PLP1</a></li></ul></li> <li><a href="/wiki/Myelin_oligodendrocyte_glycoprotein" title="Myelin oligodendrocyte glycoprotein">Myelin oligodendrocyte glycoprotein</a></li> <li><a href="/wiki/Myelin-associated_glycoprotein" title="Myelin-associated glycoprotein">Myelin-associated glycoprotein</a></li> <li><a href="/wiki/Myelin_protein_zero" title="Myelin protein zero">Myelin protein zero</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Pulmonary surfactant</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pulmonary_surfactant-associated_protein_B" class="mw-redirect" title="Pulmonary surfactant-associated protein B">Pulmonary surfactant-associated protein B</a></li> <li><a href="/wiki/Pulmonary_surfactant-associated_protein_C" class="mw-redirect" title="Pulmonary surfactant-associated protein C">Pulmonary surfactant-associated protein C</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tetraspanin" title="Tetraspanin">Tetraspanin</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/w/index.php?title=TSPAN1&action=edit&redlink=1" class="new" title="TSPAN1 (page does not exist)">TSPAN1</a></li> <li><a href="/wiki/TSPAN2" title="TSPAN2">TSPAN2</a></li> <li><a href="/wiki/TSPAN3" title="TSPAN3">TSPAN3</a></li> <li><a href="/wiki/TSPAN4" title="TSPAN4">TSPAN4</a></li> <li><a href="/w/index.php?title=TSPAN5&action=edit&redlink=1" class="new" title="TSPAN5 (page does not exist)">TSPAN5</a></li> <li><a href="/wiki/TSPAN6" title="TSPAN6">TSPAN6</a></li> <li><a href="/wiki/TSPAN7" title="TSPAN7">TSPAN7</a></li> <li><a href="/wiki/TSPAN8" title="TSPAN8">TSPAN8</a></li> <li><a href="/wiki/TSPAN9" class="mw-redirect" title="TSPAN9">TSPAN9</a></li> <li><a href="/wiki/TSPAN10" class="mw-redirect" title="TSPAN10">TSPAN10</a></li> <li><a href="/w/index.php?title=TSPAN11&action=edit&redlink=1" class="new" title="TSPAN11 (page does not exist)">TSPAN11</a></li> <li><a href="/wiki/TSPAN12" title="TSPAN12">TSPAN12</a></li> <li><a href="/wiki/TSPAN13" title="TSPAN13">TSPAN13</a></li> <li><a href="/wiki/TSPAN14" title="TSPAN14">TSPAN14</a></li> <li><a href="/w/index.php?title=TSPAN15&action=edit&redlink=1" class="new" title="TSPAN15 (page does not exist)">TSPAN15</a></li> <li><a href="/w/index.php?title=TSPAN16&action=edit&redlink=1" class="new" title="TSPAN16 (page does not exist)">TSPAN16</a></li> <li><a href="/w/index.php?title=TSPAN17&action=edit&redlink=1" class="new" title="TSPAN17 (page does not exist)">TSPAN17</a></li> <li><a href="/wiki/TSPAN18" class="mw-redirect" title="TSPAN18">TSPAN18</a></li> <li><a href="/w/index.php?title=TSPAN19&action=edit&redlink=1" class="new" title="TSPAN19 (page does not exist)">TSPAN19</a></li> <li><a href="/w/index.php?title=TSPAN20&action=edit&redlink=1" class="new" title="TSPAN20 (page does not exist)">TSPAN20</a></li> <li><a href="/wiki/TSPAN21" class="mw-redirect" title="TSPAN21">TSPAN21</a></li> <li><a href="/w/index.php?title=TSPAN22&action=edit&redlink=1" class="new" title="TSPAN22 (page does not exist)">TSPAN22</a></li> <li><a href="/wiki/TSPAN23" class="mw-redirect" title="TSPAN23">TSPAN23</a></li> <li><a href="/wiki/CD151" title="CD151">TSPAN24</a></li> <li><a href="/wiki/CD53" title="CD53">TSPAN25</a></li> <li><a href="/wiki/CD37" title="CD37">TSPAN26</a></li> <li><a href="/wiki/CD82_(gene)" title="CD82 (gene)">TSPAN27</a></li> <li><a href="/wiki/CD81" title="CD81">TSPAN28</a></li> <li><a href="/wiki/CD9" title="CD9">TSPAN29</a></li> <li><a href="/wiki/CD63" title="CD63">TSPAN30</a></li> <li><a href="/wiki/TSPAN31" title="TSPAN31">TSPAN31</a></li> <li><a href="/wiki/TSPAN32" title="TSPAN32">TSPAN32</a></li> <li><a href="/w/index.php?title=TSPAN33&action=edit&redlink=1" class="new" title="TSPAN33 (page does not exist)">TSPAN33</a></li> <li><a href="/w/index.php?title=TSPAN34&action=edit&redlink=1" class="new" title="TSPAN34 (page does not exist)">TSPAN34</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/ungrouped</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Calnexin" title="Calnexin">Calnexin</a></li> <li><a href="/wiki/LDL-receptor-related_protein-associated_protein" title="LDL-receptor-related protein-associated protein">LDL-receptor-related protein-associated protein</a></li> <li><a href="/wiki/Merlin_(protein)" title="Merlin (protein)">Neurofibromin 2</a></li> <li><a href="/wiki/Presenilin" title="Presenilin">Presenilin</a> <ul><li><a href="/wiki/PSEN1" class="mw-redirect" title="PSEN1">PSEN1</a></li> <li><a href="/wiki/PSEN2" class="mw-redirect" title="PSEN2">PSEN2</a></li></ul></li> <li><a href="/wiki/HFE_(gene)" title="HFE (gene)">HFE</a></li> <li><a href="/wiki/Phospholipid_scramblase" title="Phospholipid scramblase">Phospholipid transfer proteins</a></li> <li><a href="/wiki/Dysferlin" title="Dysferlin">Dysferlin</a></li> <li><a href="/wiki/STRC" title="STRC">STRC</a></li> <li><a href="/wiki/OTOF" class="mw-redirect" title="OTOF">OTOF</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div><i>see also <a href="/wiki/Template:Other_cell_membrane_protein_disorders" title="Template:Other cell membrane protein disorders">other cell membrane protein disorders</a></i></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Pulmonary_surfactant&oldid=1224542251">https://en.wikipedia.org/w/index.php?title=Pulmonary_surfactant&oldid=1224542251</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Respiratory_physiology" title="Category:Respiratory physiology">Respiratory physiology</a></li><li><a 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