{"id":1929,"date":"2016-05-06T22:47:21","date_gmt":"2016-05-06T22:47:21","guid":{"rendered":"https:\/\/courses.lumenlearning.com\/biologyxwaymakerxmaster\/?post_type=chapter&#038;p=1929"},"modified":"2017-04-18T17:34:35","modified_gmt":"2017-04-18T17:34:35","slug":"the-cytoskeleton","status":"publish","type":"chapter","link":"https:\/\/courses.lumenlearning.com\/suny-wmopen-biology1\/chapter\/the-cytoskeleton\/","title":{"raw":"The Cytoskeleton","rendered":"The Cytoskeleton"},"content":{"raw":"<h2>Demonstrate familiarity with various components of the cytoskeleton, including monomeric units<\/h2>\r\nIf you were to remove all the organelles from a cell, would the plasma membrane and the cytoplasm be the only components left? No. Within the cytoplasm, there would still be ions and organic molecules, plus a network of protein fibers that help maintain the shape of the cell, secure some organelles in specific positions, allow cytoplasm and vesicles to move within the cell, and enable cells within multicellular organisms to move. Collectively, this network of protein fibers is known as the\u00a0<strong>cytoskeleton<\/strong>. There are three types of fibers within the cytoskeleton: microfilaments, intermediate filaments, and microtubules (Figure 1). In this outcome, we will examine each.\r\n\r\n[caption id=\"attachment_2020\" align=\"aligncenter\" width=\"1115\"]<img class=\"size-full wp-image-2020\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/10201632\/Figure_04_05_011.jpg\" alt=\"Microfilaments line the inside of the plasma membrane, whereas microfilaments radiate out from the center of the cell. Intermediate filaments form a network throughout the cell that holds organelles in place.\" width=\"1115\" height=\"373\" \/> Figure 1. Microfilaments thicken the cortex around the inner edge of a cell; like rubber bands, they resist tension. Microtubules are found in the interior of the cell where they maintain cell shape by resisting compressive forces. Intermediate filaments are found throughout the cell and hold organelles in place.[\/caption]\r\n\r\n<div class=\"textbox learning-objectives\">\r\n<h3>Learning Objectives<\/h3>\r\n<ul>\r\n \t<li>Describe the structure and function of microfilaments<\/li>\r\n \t<li>Describe the structure and function of intermediate filaments<\/li>\r\n \t<li>Describe the structure and function of microtubules<\/li>\r\n<\/ul>\r\n<\/div>\r\n<h2>Microfilaments<\/h2>\r\n[caption id=\"attachment_1707\" align=\"alignright\" width=\"300\"]<img class=\"wp-image-1707\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03184656\/Figure_04_05_02.jpg\" alt=\"This illustration shows two actin filaments wound together. Each actin filament is composed of many actin subunits connected together to form a chain.\" width=\"300\" height=\"415\" \/> Figure 2. Microfilaments are made of two intertwined strands of actin.[\/caption]\r\n\r\nOf the three types of protein fibers in the cytoskeleton, <strong>microfilaments<\/strong> are the narrowest. They function in cellular movement, have a diameter of about 7 nm, and are made of two intertwined strands of a globular protein called actin (Figure 2). For this reason, microfilaments are also known as actin filaments.\r\n\r\nActin is powered by ATP to assemble its filamentous form, which serves as a track for the movement of a motor protein called myosin. This enables actin to engage in cellular events requiring motion, such as cell division in animal cells and cytoplasmic streaming, which is the circular movement of the cell cytoplasm in plant cells. Actin and myosin are plentiful in muscle cells. When your actin and myosin filaments slide past each other, your muscles contract.\r\n\r\nMicrofilaments also provide some rigidity and shape to the cell. They can depolymerize (disassemble) and reform quickly, thus enabling a cell to change its shape and move. White blood cells (your body\u2019s infection-fighting cells) make good use of this ability. They can move to the site of an infection and phagocytize the pathogen.\r\n\r\nLook below see an example of a white blood cell in action.\u00a0Watch this\u00a0short time-lapse video of the cell capturing two bacteria. It engulfs one and then moves on to the other. Note that this video has no audio.\r\n\r\nhttps:\/\/youtu.be\/AUih856vaQY\r\n<h2>Intermediate Filaments<\/h2>\r\nIntermediate filaments are made of several strands of fibrous proteins that are wound together (Figure 3). These elements of the cytoskeleton get their name from the fact that their diameter, 8 to 10 nm, is between those of microfilaments and microtubules.\r\n\r\n[caption id=\"attachment_1710\" align=\"aligncenter\" width=\"544\"]<img class=\"size-full wp-image-1710\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03185218\/Figure_04_05_03.jpg\" alt=\"This illustration shows 10 intermediate filament fibers bundled together.\" width=\"544\" height=\"116\" \/> Figure 3. Intermediate filaments consist of several intertwined strands of fibrous proteins.[\/caption]\r\n\r\nIntermediate filaments have no role in cell movement. Their function is purely structural. They bear tension, thus maintaining the shape of the cell, and anchor the nucleus and other organelles in place. Figure 4\u00a0shows how intermediate filaments create a supportive scaffolding inside the cell.\r\n\r\nThe intermediate filaments are the most diverse group of cytoskeletal elements. Several types of fibrous proteins are found in the intermediate filaments. You are probably most familiar with keratin, the fibrous protein that strengthens your hair, nails, and the epidermis of the skin.\r\n\r\n[caption id=\"attachment_2020\" align=\"aligncenter\" width=\"1115\"]<img class=\"size-full wp-image-2020\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/10201632\/Figure_04_05_011.jpg\" alt=\"Microfilaments line the inside of the plasma membrane, whereas microfilaments radiate out from the center of the cell. Intermediate filaments form a network throughout the cell that holds organelles in place.\" width=\"1115\" height=\"373\" \/> Figure 4. Microfilaments thicken the cortex around the inner edge of a cell; like rubber bands, they resist tension. Microtubules are found in the interior of the cell where they maintain cell shape by resisting compressive forces. Intermediate filaments are found throughout the cell and hold organelles in place.[\/caption]\r\n<h2>Microtubules<\/h2>\r\nAs their name implies, microtubules are small hollow tubes. The walls of the microtubule are made of polymerized dimers of <em>\u03b1<\/em>-tubulin and <em>\u03b2<\/em>-tubulin, two globular proteins (Figure 5). With a diameter of about 25 nm, microtubules are the widest components of the cytoskeleton. They help the cell resist compression, provide a track along which vesicles move through the cell, and pull replicated chromosomes to opposite ends of a dividing cell. Like microfilaments, microtubules can dissolve and reform quickly.\r\n\r\n[caption id=\"attachment_1708\" align=\"aligncenter\" width=\"800\"]<img class=\"size-full wp-image-1708\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03185007\/Figure_04_05_04ab.jpg\" alt=\"The left part of this figure is a molecular model of 13 polymerized dimers of alpha- and beta-tubulin joined together to form a hollow tube. The right part of this image shows the tubulin structure as a ring of spheres connected together.\" width=\"800\" height=\"426\" \/> Figure 5. Microtubules are hollow. Their walls consist of 13 polymerized dimers of \u03b1-tubulin and \u03b2-tubulin (right image). The left image shows the molecular structure of the tube.[\/caption]\r\n\r\nMicrotubules are also the structural elements of flagella, cilia, and centrioles (the latter are the two perpendicular bodies of the centrosome). In fact, in animal cells, the centrosome is the microtubule-organizing center. In eukaryotic cells, flagella and cilia are quite different structurally from their counterparts in prokaryotes, as discussed below.\r\n<h3>Flagella and Cilia<\/h3>\r\n[caption id=\"attachment_1709\" align=\"alignright\" width=\"350\"]<img class=\" wp-image-1709\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03185106\/Figure_04_05_05-300x236.jpg\" alt=\"This transmission electron micrograph shows a cross section of nine microtubule doublets that form a hollow tube. Another microtubule doublet sits in the center of the tube.\" width=\"350\" height=\"275\" \/> Figure 6. This transmission electron micrograph of two flagella shows the 9 + 2 array of microtubules (credit: modification of work by Dartmouth Electron Microscope Facility, Dartmouth College; scale-bar data from Matt Russell)[\/caption]\r\n\r\nTo refresh your memory, flagella (singular = <em>flagellum<\/em>) are long, hair-like structures that extend from the plasma membrane and are used to move an entire cell (for example, sperm, <em>Euglena<\/em>). When present, the cell has just one flagellum or a few flagella. When cilia (singular = <em>cilium<\/em>) are present, however, many of them extend along the entire surface of the plasma membrane. They are short, hair-like structures that are used to move entire cells (such as paramecia) or substances along the outer surface of the cell (for example, the cilia of cells lining the Fallopian tubes that move the ovum toward the uterus, or cilia lining the cells of the respiratory tract that trap particulate matter and move it toward your nostrils.)\r\n\r\nDespite their differences in length and number, flagella and cilia share a common structural arrangement of microtubules called a \"9 + 2 array.\" This is an appropriate name because a single flagellum or cilium is made of a ring of nine microtubule doublets, surrounding a single microtubule doublet in the center (Figure 6).\r\n<h2><strong>Check Your Understanding<\/strong><\/h2>\r\nAnswer the question(s) below to see how well you understand the topics covered in the previous section. This short quiz does\u00a0<strong>not<\/strong>\u00a0count toward your grade in the class, and you can retake it an unlimited number of times.\r\n\r\nUse this quiz to check your understanding and decide whether to (1) study the previous section further or (2) move on to the next section.\r\n\r\nhttps:\/\/assessments.lumenlearning.com\/assessments\/3328","rendered":"<h2>Demonstrate familiarity with various components of the cytoskeleton, including monomeric units<\/h2>\n<p>If you were to remove all the organelles from a cell, would the plasma membrane and the cytoplasm be the only components left? No. Within the cytoplasm, there would still be ions and organic molecules, plus a network of protein fibers that help maintain the shape of the cell, secure some organelles in specific positions, allow cytoplasm and vesicles to move within the cell, and enable cells within multicellular organisms to move. Collectively, this network of protein fibers is known as the\u00a0<strong>cytoskeleton<\/strong>. There are three types of fibers within the cytoskeleton: microfilaments, intermediate filaments, and microtubules (Figure 1). In this outcome, we will examine each.<\/p>\n<div id=\"attachment_2020\" style=\"width: 1125px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-2020\" class=\"size-full wp-image-2020\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/10201632\/Figure_04_05_011.jpg\" alt=\"Microfilaments line the inside of the plasma membrane, whereas microfilaments radiate out from the center of the cell. Intermediate filaments form a network throughout the cell that holds organelles in place.\" width=\"1115\" height=\"373\" \/><\/p>\n<p id=\"caption-attachment-2020\" class=\"wp-caption-text\">Figure 1. Microfilaments thicken the cortex around the inner edge of a cell; like rubber bands, they resist tension. Microtubules are found in the interior of the cell where they maintain cell shape by resisting compressive forces. Intermediate filaments are found throughout the cell and hold organelles in place.<\/p>\n<\/div>\n<div class=\"textbox learning-objectives\">\n<h3>Learning Objectives<\/h3>\n<ul>\n<li>Describe the structure and function of microfilaments<\/li>\n<li>Describe the structure and function of intermediate filaments<\/li>\n<li>Describe the structure and function of microtubules<\/li>\n<\/ul>\n<\/div>\n<h2>Microfilaments<\/h2>\n<div id=\"attachment_1707\" style=\"width: 310px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-1707\" class=\"wp-image-1707\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03184656\/Figure_04_05_02.jpg\" alt=\"This illustration shows two actin filaments wound together. Each actin filament is composed of many actin subunits connected together to form a chain.\" width=\"300\" height=\"415\" \/><\/p>\n<p id=\"caption-attachment-1707\" class=\"wp-caption-text\">Figure 2. Microfilaments are made of two intertwined strands of actin.<\/p>\n<\/div>\n<p>Of the three types of protein fibers in the cytoskeleton, <strong>microfilaments<\/strong> are the narrowest. They function in cellular movement, have a diameter of about 7 nm, and are made of two intertwined strands of a globular protein called actin (Figure 2). For this reason, microfilaments are also known as actin filaments.<\/p>\n<p>Actin is powered by ATP to assemble its filamentous form, which serves as a track for the movement of a motor protein called myosin. This enables actin to engage in cellular events requiring motion, such as cell division in animal cells and cytoplasmic streaming, which is the circular movement of the cell cytoplasm in plant cells. Actin and myosin are plentiful in muscle cells. When your actin and myosin filaments slide past each other, your muscles contract.<\/p>\n<p>Microfilaments also provide some rigidity and shape to the cell. They can depolymerize (disassemble) and reform quickly, thus enabling a cell to change its shape and move. White blood cells (your body\u2019s infection-fighting cells) make good use of this ability. They can move to the site of an infection and phagocytize the pathogen.<\/p>\n<p>Look below see an example of a white blood cell in action.\u00a0Watch this\u00a0short time-lapse video of the cell capturing two bacteria. It engulfs one and then moves on to the other. Note that this video has no audio.<\/p>\n<p><iframe loading=\"lazy\" id=\"oembed-1\" title=\"white blood cell chases bacteria\" width=\"500\" height=\"375\" src=\"https:\/\/www.youtube.com\/embed\/AUih856vaQY?feature=oembed&#38;rel=0\" frameborder=\"0\" allowfullscreen=\"allowfullscreen\"><\/iframe><\/p>\n<h2>Intermediate Filaments<\/h2>\n<p>Intermediate filaments are made of several strands of fibrous proteins that are wound together (Figure 3). These elements of the cytoskeleton get their name from the fact that their diameter, 8 to 10 nm, is between those of microfilaments and microtubules.<\/p>\n<div id=\"attachment_1710\" style=\"width: 554px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-1710\" class=\"size-full wp-image-1710\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03185218\/Figure_04_05_03.jpg\" alt=\"This illustration shows 10 intermediate filament fibers bundled together.\" width=\"544\" height=\"116\" \/><\/p>\n<p id=\"caption-attachment-1710\" class=\"wp-caption-text\">Figure 3. Intermediate filaments consist of several intertwined strands of fibrous proteins.<\/p>\n<\/div>\n<p>Intermediate filaments have no role in cell movement. Their function is purely structural. They bear tension, thus maintaining the shape of the cell, and anchor the nucleus and other organelles in place. Figure 4\u00a0shows how intermediate filaments create a supportive scaffolding inside the cell.<\/p>\n<p>The intermediate filaments are the most diverse group of cytoskeletal elements. Several types of fibrous proteins are found in the intermediate filaments. You are probably most familiar with keratin, the fibrous protein that strengthens your hair, nails, and the epidermis of the skin.<\/p>\n<div id=\"attachment_2020\" style=\"width: 1125px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-2020\" class=\"size-full wp-image-2020\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/10201632\/Figure_04_05_011.jpg\" alt=\"Microfilaments line the inside of the plasma membrane, whereas microfilaments radiate out from the center of the cell. Intermediate filaments form a network throughout the cell that holds organelles in place.\" width=\"1115\" height=\"373\" \/><\/p>\n<p id=\"caption-attachment-2020\" class=\"wp-caption-text\">Figure 4. Microfilaments thicken the cortex around the inner edge of a cell; like rubber bands, they resist tension. Microtubules are found in the interior of the cell where they maintain cell shape by resisting compressive forces. Intermediate filaments are found throughout the cell and hold organelles in place.<\/p>\n<\/div>\n<h2>Microtubules<\/h2>\n<p>As their name implies, microtubules are small hollow tubes. The walls of the microtubule are made of polymerized dimers of <em>\u03b1<\/em>-tubulin and <em>\u03b2<\/em>-tubulin, two globular proteins (Figure 5). With a diameter of about 25 nm, microtubules are the widest components of the cytoskeleton. They help the cell resist compression, provide a track along which vesicles move through the cell, and pull replicated chromosomes to opposite ends of a dividing cell. Like microfilaments, microtubules can dissolve and reform quickly.<\/p>\n<div id=\"attachment_1708\" style=\"width: 810px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-1708\" class=\"size-full wp-image-1708\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03185007\/Figure_04_05_04ab.jpg\" alt=\"The left part of this figure is a molecular model of 13 polymerized dimers of alpha- and beta-tubulin joined together to form a hollow tube. The right part of this image shows the tubulin structure as a ring of spheres connected together.\" width=\"800\" height=\"426\" \/><\/p>\n<p id=\"caption-attachment-1708\" class=\"wp-caption-text\">Figure 5. Microtubules are hollow. Their walls consist of 13 polymerized dimers of \u03b1-tubulin and \u03b2-tubulin (right image). The left image shows the molecular structure of the tube.<\/p>\n<\/div>\n<p>Microtubules are also the structural elements of flagella, cilia, and centrioles (the latter are the two perpendicular bodies of the centrosome). In fact, in animal cells, the centrosome is the microtubule-organizing center. In eukaryotic cells, flagella and cilia are quite different structurally from their counterparts in prokaryotes, as discussed below.<\/p>\n<h3>Flagella and Cilia<\/h3>\n<div id=\"attachment_1709\" style=\"width: 360px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-1709\" class=\"wp-image-1709\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/03185106\/Figure_04_05_05-300x236.jpg\" alt=\"This transmission electron micrograph shows a cross section of nine microtubule doublets that form a hollow tube. Another microtubule doublet sits in the center of the tube.\" width=\"350\" height=\"275\" \/><\/p>\n<p id=\"caption-attachment-1709\" class=\"wp-caption-text\">Figure 6. This transmission electron micrograph of two flagella shows the 9 + 2 array of microtubules (credit: modification of work by Dartmouth Electron Microscope Facility, Dartmouth College; scale-bar data from Matt Russell)<\/p>\n<\/div>\n<p>To refresh your memory, flagella (singular = <em>flagellum<\/em>) are long, hair-like structures that extend from the plasma membrane and are used to move an entire cell (for example, sperm, <em>Euglena<\/em>). When present, the cell has just one flagellum or a few flagella. When cilia (singular = <em>cilium<\/em>) are present, however, many of them extend along the entire surface of the plasma membrane. They are short, hair-like structures that are used to move entire cells (such as paramecia) or substances along the outer surface of the cell (for example, the cilia of cells lining the Fallopian tubes that move the ovum toward the uterus, or cilia lining the cells of the respiratory tract that trap particulate matter and move it toward your nostrils.)<\/p>\n<p>Despite their differences in length and number, flagella and cilia share a common structural arrangement of microtubules called a &#8220;9 + 2 array.&#8221; This is an appropriate name because a single flagellum or cilium is made of a ring of nine microtubule doublets, surrounding a single microtubule doublet in the center (Figure 6).<\/p>\n<h2><strong>Check Your Understanding<\/strong><\/h2>\n<p>Answer the question(s) below to see how well you understand the topics covered in the previous section. This short quiz does\u00a0<strong>not<\/strong>\u00a0count toward your grade in the class, and you can retake it an unlimited number of times.<\/p>\n<p>Use this quiz to check your understanding and decide whether to (1) study the previous section further or (2) move on to the next section.<\/p>\n<p>\t<iframe id=\"lumen_assessment_3328\" class=\"resizable\" src=\"https:\/\/assessments.lumenlearning.com\/assessments\/load?assessment_id=3328&#38;embed=1&#38;external_user_id=&#38;external_context_id=&#38;iframe_resize_id=lumen_assessment_3328\" frameborder=\"0\" style=\"border:none;width:100%;height:100%;min-height:400px;\"><br \/>\n\t<\/iframe><\/p>\n\n\t\t\t <section class=\"citations-section\" role=\"contentinfo\">\n\t\t\t <h3>Candela Citations<\/h3>\n\t\t\t\t\t <div>\n\t\t\t\t\t\t <div id=\"citation-list-1929\">\n\t\t\t\t\t\t\t <div class=\"licensing\"><div class=\"license-attribution-dropdown-subheading\">CC licensed content, Original<\/div><ul class=\"citation-list\"><li><strong>Authored by<\/strong>: Shelli Carter and Lumen Learning. <strong>Provided by<\/strong>: Lumen Learning. <strong>License<\/strong>: <em><a target=\"_blank\" rel=\"license\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY: Attribution<\/a><\/em><\/li><\/ul><div class=\"license-attribution-dropdown-subheading\">CC licensed content, Shared previously<\/div><ul class=\"citation-list\"><li>Biology. <strong>Provided by<\/strong>: OpenStax CNX. <strong>Located at<\/strong>: <a target=\"_blank\" href=\"http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8\">http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8<\/a>. <strong>License<\/strong>: <em><a target=\"_blank\" rel=\"license\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY: Attribution<\/a><\/em>. <strong>License Terms<\/strong>: Download for free at http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8<\/li><\/ul><div class=\"license-attribution-dropdown-subheading\">All rights reserved content<\/div><ul class=\"citation-list\"><li>white blood cell chases bacteria. <strong>Authored by<\/strong>: Brad Cameron. <strong>Located at<\/strong>: <a target=\"_blank\" href=\"https:\/\/youtu.be\/AUih856vaQY\">https:\/\/youtu.be\/AUih856vaQY<\/a>. <strong>License<\/strong>: <em>All Rights Reserved<\/em>. <strong>License Terms<\/strong>: Standard YouTube License<\/li><\/ul><\/div>\n\t\t\t\t\t\t <\/div>\n\t\t\t\t\t <\/div>\n\t\t\t <\/section>","protected":false},"author":17,"menu_order":4,"template":"","meta":{"_candela_citation":"[{\"type\":\"original\",\"description\":\"\",\"author\":\"Shelli Carter and Lumen Learning\",\"organization\":\"Lumen Learning\",\"url\":\"\",\"project\":\"\",\"license\":\"cc-by\",\"license_terms\":\"\"},{\"type\":\"cc\",\"description\":\"Biology\",\"author\":\"\",\"organization\":\"OpenStax 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