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Key Terms acid a substance that donates hydrogen ions and therefore lowers pH adhesion the attraction between water molecules and molecules of a different substance amino acid a monomer of a protein anion a negative ion formed by gaining electrons atomic number the number of protons in an atom base a substance that absorbs hydrogen ions and therefore raises pH buffer a solution that resists a change in pH by absorbing or releasing hydrogen or hydroxide ions carbohydrate a biological macromolecule in which the ratio of carbon to hydrogen to oxygen is 1:2:1; carbohydrates serve as energy sources and structural support in cells cation a positive ion formed by losing electrons cellulose a polysaccharide that makes up the cell walls of plants and provides structural support to the cell chemical bond an interaction between two or more of the same or different elements that results in the formation of molecules chitin a type of carbohydrate that forms the outer
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the cell chemical bond an interaction between two or more of the same or different elements that results in the formation of molecules chitin a type of carbohydrate that forms the outer skeleton of arthropods, such as insects and crustaceans, and the cell walls of fungi cohesion the intermolecular forces between water molecules caused by the polar nature of water; creates surface tension covalent bond a type of strong bond between two or more of the same or different elements; forms when electrons are shared between elements denaturation the loss of shape in a protein as a result of changes in temperature, pH, or exposure to chemicals deoxyribonucleic acid (DNA) a double-stranded polymer of nucleotides that carries the hereditary information of the cell disaccharide two sugar monomers that are linked together by a glycodsidic bond electron a negatively charged particle that resides outside of the nucleus in the electron orbital; lacks functional mass and has a charge of –1
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together by a glycodsidic bond electron a negatively charged particle that resides outside of the nucleus in the electron orbital; lacks functional mass and has a charge of –1 electron transfer the movement of electrons from one element to another element one of 118 unique substances that cannot be broken down into smaller substances and retain the characteristic of that substance; each element has a specified number of protons and unique properties enzyme a catalyst in a biochemical reaction that is usually a complex or conjugated protein evaporation the release of water molecules from liquid water to form water vapor fat a lipid molecule composed of three fatty acids and a glycerol (triglyceride) that typically exists in a solid form at room temperature glycogen a storage carbohydrate in animals hormone a chemical signaling molecule, usually a protein or steroid, secreted by an endocrine gland or group of endocrine cells; acts to control or regulate specific physiological processes
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hormone a chemical signaling molecule, usually a protein or steroid, secreted by an endocrine gland or group of endocrine cells; acts to control or regulate specific physiological processes hydrogen bond a weak bond between partially positively charged hydrogen atoms and partially negatively charged elements or molecules hydrophilic describes a substance that dissolves in water; water-loving hydrophobic describes a substance that does not dissolve in water; water-fearing ion an atom or compound that does not contain equal numbers of protons and electrons, and therefore has a net charge ionic bond a chemical bond that forms between ions of opposite charges isotope one or more forms of an element that have different numbers of neutrons lipids a class of macromolecules that are nonpolar and insoluble in water litmus paper filter paper that has been treated with a natural water-soluble dye so it can be used as a pH indicator macromolecule a large molecule, often formed by
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and insoluble in water litmus paper filter paper that has been treated with a natural water-soluble dye so it can be used as a pH indicator macromolecule a large molecule, often formed by polymerization of smaller monomers mass number the number of protons plus neutrons in an atom matter anything that has mass and occupies space monosaccharide a single unit or monomer of carbohydrates neutron a particle with no charge that resides in the nucleus of an atom; has a mass of 1 nonpolar covalent bond a type of covalent bond that forms between atoms when electrons are shared equally between atoms, resulting in no regions with partial charges as in polar covalent bonds nucleic acid a biological macromolecule that carries the genetic information of a cell and carries instructions for the functioning of the cell nucleotide a monomer of nucleic acids; contains a pentose sugar, a phosphate group, and a nitrogenous base nucleus (chemistry) the dense center of an atom
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instructions for the functioning of the cell nucleotide a monomer of nucleic acids; contains a pentose sugar, a phosphate group, and a nitrogenous base nucleus (chemistry) the dense center of an atom made up of protons and (except in the case of a hydrogen atom) neutrons octet rule states that the outermost shell of an element with a low atomic number can hold eight 2 • Key Terms 51
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electrons oil an unsaturated fat that is a liquid at room temperature periodic table of elements an organizational chart of elements, indicating the atomic number and mass number of each element; also provides key information about the properties of elements pH scale a scale ranging from 0 to 14 that measures the approximate concentration of hydrogen ions of a substance phospholipid a major constituent of the membranes of cells; composed of two fatty acids and a phosphate group attached to the glycerol backbone polar covalent bond a type of covalent bond in which electrons are pulled toward one atom and away from another, resulting in slightly positive and slightly negative charged regions of the molecule polypeptide a long chain of amino acids linked by peptide bonds polysaccharide a long chain of monosaccharides; may be branched or unbranched protein a biological macromolecule composed of one or more chains of amino acids proton a positively charged particle that resides in
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may be branched or unbranched protein a biological macromolecule composed of one or more chains of amino acids proton a positively charged particle that resides in the nucleus of an atom; has a mass of 1 and a charge of +1 radioactive isotope an isotope that spontaneously emits particles or energy to form a more stable element ribonucleic acid (RNA) a single-stranded polymer of nucleotides that is involved in protein synthesis saturated fatty acid a long-chain hydrocarbon with single covalent bonds in the carbon chain; the number of hydrogen atoms attached to the carbon skeleton is maximized solvent a substance capable of dissolving another substance starch a storage carbohydrate in plants steroid a type of lipid composed of four fused hydrocarbon rings surface tension the cohesive force at the surface of a body of liquid that prevents the molecules from separating temperature a measure of molecular motion trans-fat a form of unsaturated fat with the hydrogen
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body of liquid that prevents the molecules from separating temperature a measure of molecular motion trans-fat a form of unsaturated fat with the hydrogen atoms neighboring the double bond across from each other rather than on the same side of the double bond triglyceride a fat molecule; consists of three fatty acids linked to a glycerol molecule unsaturated fatty acid a long-chain hydrocarbon that has one or more than one double bonds in the hydrocarbon chain van der Waals interaction a weak attraction or interaction between molecules caused by slightly positively charged or slightly negatively charged atoms Chapter Summary 2.1 The Building Blocks of Molecules Matter is anything that occupies space and has mass. It is made up of atoms of different elements. All of the 92 elements that occur naturally have unique qualities that allow them to combine in various ways to create compounds or molecules. Atoms, which consist of protons, neutrons, and electrons, are the smallest units
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that allow them to combine in various ways to create compounds or molecules. Atoms, which consist of protons, neutrons, and electrons, are the smallest units of an element that retain all of the properties of that element. Electrons can be donated or shared between atoms to create bonds, including ionic, covalent, and hydrogen bonds, as well as van der Waals interactions. 2.2 Water Water has many properties that are critical to maintaining life. It is polar, allowing for the formation of hydrogen bonds, which allow ions and other polar molecules to dissolve in water. Therefore, water is an excellent solvent. The hydrogen bonds between water molecules give water the ability to hold heat better than many other substances. As the temperature rises, the hydrogen bonds between water continually break and reform, allowing for the overall temperature to remain stable, although increased energy is added to the system. Water’s cohesive forces allow for the
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and reform, allowing for the overall temperature to remain stable, although increased energy is added to the system. Water’s cohesive forces allow for the property of surface tension. All of these unique properties of water are important in the chemistry of living organisms. The pH of a solution is a measure of the concentration of hydrogen ions in the solution. A solution with a high number of hydrogen ions is acidic and has a low pH value. A solution with a high number of hydroxide ions is basic and has a high pH value. The pH scale ranges from 0 to 14, with a pH of 7 being neutral. Buffers are solutions that moderate pH changes when an acid or base is added to the buffer system. Buffers are important in biological systems because of their ability to maintain constant pH conditions. 2.3 Biological Molecules Living things are carbon-based because carbon plays such a prominent role in the chemistry of living things. The four covalent bonding positions of the carbon atom
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2.3 Biological Molecules Living things are carbon-based because carbon plays such a prominent role in the chemistry of living things. The four covalent bonding positions of the carbon atom can give rise to a wide diversity of compounds with many functions, accounting for the importance of 52 2 • Chapter Summary Access for free at openstax.org
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carbon in living things. Carbohydrates are a group of macromolecules that are a vital energy source for the cell, provide structural support to many organisms, and can be found on the surface of the cell as receptors or for cell recognition. Carbohydrates are classified as monosaccharides, disaccharides, and polysaccharides, depending on the number of monomers in the molecule. Lipids are a class of macromolecules that are nonpolar and hydrophobic in nature. Major types include fats and oils, waxes, phospholipids, and steroids. Fats and oils are a stored form of energy and can include triglycerides. Fats and oils are usually made up of fatty acids and glycerol. Proteins are a class of macromolecules that can perform a diverse range of functions for the cell. They help in metabolism by providing structural support and by acting as enzymes, carriers or as hormones. The building blocks of proteins are amino acids. Proteins are organized at four levels: primary, secondary,
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by acting as enzymes, carriers or as hormones. The building blocks of proteins are amino acids. Proteins are organized at four levels: primary, secondary, tertiary, and quaternary. Protein shape and function are intricately linked; any change in shape caused by changes in temperature, pH, or chemical exposure may lead to protein denaturation and a loss of function. Nucleic acids are molecules made up of repeating units of nucleotides that direct cellular activities such as cell division and protein synthesis. Each nucleotide is made up of a pentose sugar, a nitrogenous base, and a phosphate group. There are two types of nucleic acids: DNA and RNA. Visual Connection Questions 1. Figure 2.3How many neutrons do (K) potassium-39 and potassium-40 have, respectively? Review Questions 2. Magnesium has an atomic number of 12. Which of the following statements is true of a neutral magnesium atom? a. It has 12 protons, 12 electrons, and 12 neutrons. b. It has 12 protons, 12 electrons, and six
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the following statements is true of a neutral magnesium atom? a. It has 12 protons, 12 electrons, and 12 neutrons. b. It has 12 protons, 12 electrons, and six neutrons. c. It has six protons, six electrons, and no neutrons. d. It has six protons, six electrons, and six neutrons. 3. Which type of bond represents a weak chemical bond? a. hydrogen bond b. ionic bond c. covalent bond d. polar covalent bond 4. An isotope of sodium (Na) has a mass number of 22. How many neutrons does it have? a. 11 b. 12 c. 22 d. 44 5. Which of the following statements is not true? a. Water is polar. b. Water stabilizes temperature. c. Water is essential for life. d. Water is the most abundant atom in Earth’s atmosphere. 6. Using a pH meter, you find the pH of an unknown solution to be 8.0. How would you describe this solution? a. weakly acidic b. strongly acidic c. weakly basic d. strongly basic 7. The pH of lemon juice is about 2.0, whereas tomato juice's pH is about 4.0. Approximately how much of
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solution? a. weakly acidic b. strongly acidic c. weakly basic d. strongly basic 7. The pH of lemon juice is about 2.0, whereas tomato juice's pH is about 4.0. Approximately how much of an increase in hydrogen ion concentration is there between tomato juice and lemon juice? a. 2 times b. 10 times c. 100 times d. 1000 times 8. An example of a monosaccharide is ________. a. fructose b. glucose c. galactose d. all of the above 2 • Visual Connection Questions 53
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9. Cellulose and starch are examples of ________. a. monosaccharides b. disaccharides c. lipids d. polysaccharides 10. Phospholipids are important components of __________. a. the plasma membrane of cells b. the ring structure of steroids c. the waxy covering on leaves d. the double bond in hydrocarbon chains 11. The monomers that make up proteins are called _________. a. nucleotides b. disaccharides c. amino acids d. chaperones Critical Thinking Questions 12. Why are hydrogen bonds and van der Waals interactions necessary for cells? 13. Why can some insects walk on water? 14. Explain why water is an excellent solvent. 15. Explain at least three functions that lipids serve in plants and/or animals. 16. Explain what happens if even one amino acid is substituted for another in a polypeptide chain. Provide a specific example. 54 2 • Critical Thinking Questions Access for free at openstax.org
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INTRODUCTION CHAPTER 3 Cell Structure and Function 3.1How Cells Are Studied 3.2Comparing Prokaryotic and Eukaryotic Cells 3.3Eukaryotic Cells 3.4The Cell Membrane 3.5Passive Transport 3.6Active Transport Close your eyes and picture a brick wall. What is the basic building block of that wall? It is a single brick, of course. Like a brick wall, your body is composed of basic building blocks, and the building blocks of your body are cells. Your body has many kinds of cells, each specialized for a specific purpose. Just as a home is made from a variety of building materials, the human body is constructed from many cell types. For example, epithelial cells protect the surface of the body and cover the organs and body cavities within. Bone cells help to support and protect the body. Cells of the immune system fight invading bacteria. Additionally, red blood cells carry oxygen throughout the body. Each of these cell types
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within. Bone cells help to support and protect the body. Cells of the immune system fight invading bacteria. Additionally, red blood cells carry oxygen throughout the body. Each of these cell types plays a vital role during the growth, development, and day-to-day maintenance of the body. In spite of their enormous variety, however, all cells share certain fundamental characteristics. 3.1 How Cells Are Studied LEARNING OBJECTIVES By the end of this section, you will be able to: • Describe the roles of cells in organisms • Compare and contrast light microscopy and electron microscopy • Summarize the cell theory A cell is the smallest unit of a living thing. A living thing, like you, is called an organism. Thus, cells are the basic building blocks of all organisms. FIGURE 3.1 (a) Nasal sinus cells (viewed with a light microscope), (b) onion cells (viewed with a light microscope),
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are the basic building blocks of all organisms. FIGURE 3.1 (a) Nasal sinus cells (viewed with a light microscope), (b) onion cells (viewed with a light microscope), and (c)Vibrio tasmaniensisbacterial cells (viewed using a scanning electron microscope) are from very different organisms, yet all share certain characteristics of basic cell structure. (credit a: modification of work by Ed Uthman, MD; credit b: modification of work by Umberto Salvagnin; credit c: modification of work by Anthony D'Onofrio; scale- bar data from Matt Russell) CHAPTER OUTLINE
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In multicellular organisms, several cells of one particular kind interconnect with each other and perform shared functions to form tissues (for example, muscle tissue, connective tissue, and nervous tissue), several tissues combine to form an organ (for example, stomach, heart, or brain), and several organs make up an organ system (such as the digestive system, circulatory system, or nervous system). Several systems functioning together form an organism (such as an elephant, for example). There are many types of cells, and all are grouped into one of two broad categories: prokaryotic and eukaryotic. Animal cells, plant cells, fungal cells, and protist cells are classified as eukaryotic, whereas bacteria and archaea cells are classified as prokaryotic. Before discussing the criteria for determining whether a cell is prokaryotic or eukaryotic, let us first examine how biologists study cells. Microscopy
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determining whether a cell is prokaryotic or eukaryotic, let us first examine how biologists study cells. Microscopy Cells vary in size. With few exceptions, individual cells are too small to be seen with the naked eye, so scientists use microscopes to study them. Amicroscopeis an instrument that magnifies an object. Most images of cells are taken with a microscope and are called micrographs. Light Microscopes To give you a sense of the size of a cell, a typical human red blood cell is about eight millionths of a meter or eight micrometers (abbreviated as µm) in diameter; the head of a pin is about two thousandths of a meter (millimeters, or mm) in diameter. That means that approximately 250 red blood cells could fit on the head of a pin. The optics of the lenses of a light microscope changes the orientation of the image. A specimen that is right-side up and facing right on the microscope slide will appear upside-down and facing
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The optics of the lenses of a light microscope changes the orientation of the image. A specimen that is right-side up and facing right on the microscope slide will appear upside-down and facing left when viewed through a microscope, and vice versa. Similarly, if the slide is moved left while looking through the microscope, it will appear to move right, and if moved down, it will seem to move up. This occurs because microscopes use two sets of lenses to magnify the image. Due to the manner in which light travels through the lenses, this system of lenses produces an inverted image (binoculars and a dissecting microscope work in a similar manner, but include an additional magnification system that makes the final image appear to be upright). Most student microscopes are classified as light microscopes (Figure 3.2a). Visible light both passes through and is bent by the lens system to enable the user to see the specimen. Light
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Most student microscopes are classified as light microscopes (Figure 3.2a). Visible light both passes through and is bent by the lens system to enable the user to see the specimen. Light microscopes are advantageous for viewing living organisms, but since individual cells are generally transparent, their components are not distinguishable unless they are colored with special stains. Staining, however, usually kills the cells. Light microscopes commonly used in the undergraduate college laboratory magnify up to approximately 400 times. Two parameters that are important in microscopy are magnification and resolving power. Magnification is the degree of enlargement of an object. Resolving power is the ability of a microscope to allow the eye to distinguish two adjacent structures as separate; the higher the resolution, the closer those two objects can be, and the better the clarity and detail of
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ability of a microscope to allow the eye to distinguish two adjacent structures as separate; the higher the resolution, the closer those two objects can be, and the better the clarity and detail of the image. When oil immersion lenses are used, magnification is usually increased to 1,000 times for the study of smaller cells, like most prokaryotic cells. Because light entering a specimen from below is focused onto the eye of an observer, the specimen can be viewed using light microscopy. For this reason, for light to pass through a specimen, the sample must be thin or translucent. LINK T O LE ARNING For another perspective on cell size, try theHowBig(http://openstax.org/l/cell_sizes2)interactive. A second type of microscope used in laboratories is the dissecting microscope (Figure 3.2b). These microscopes have a lower magnification (20 to 80 times the object size) than light 56 3 • Cell Structure and Function Access for free at openstax.org
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microscopes and can provide a three-dimensional view of the specimen. Thick objects can be examined with many components in focus at the same time. These microscopes are designed to give a magnified and clear view of tissue structure as well as the anatomy of the whole organism. Like light microscopes, most modern dissecting microscopes are also binocular, meaning that they have two separate lens systems, one for each eye. The lens systems are separated by a certain distance, and therefore provide a sense of depth in the view of their subject to make manipulations by hand easier. Dissecting microscopes also have optics that correct the image so that it appears as if being seen by the naked eye and not as an inverted image. The light illuminating a sample under a dissecting microscope typically comes from above the sample, but may also be directed from below.
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dissecting microscope typically comes from above the sample, but may also be directed from below. FIGURE 3.2 (a) Most light microscopes used in a college biology lab can magnify cells up to approximately 400 times. (b) Dissecting microscopes have a lower magnification than light microscopes and are used to examine larger objects, such as tissues. Electron Microscopes In contrast to light microscopes, electron microscopes use a beam of electrons instead of a beam of light. Not only does this allow for higher magnification and, thus, more detail (Figure 3.3), it also provides higher resolving power. Preparation of a specimen for viewing under an electron microscope will kill it; therefore, live cells cannot be viewed using this type of microscopy. In addition, the electron beam moves best in a vacuum, making it impossible to view living materials. In a scanning electron microscope, a beam of electrons moves back and forth across a cell’s surface, rendering the
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living materials. In a scanning electron microscope, a beam of electrons moves back and forth across a cell’s surface, rendering the details of cell surface characteristics by reflection. Cells and other structures are usually coated with a metal like gold. In a transmission electron microscope, the electron beam is transmitted through the cell and provides details of a cell’s internal structures. As you might imagine, electron microscopes are significantly more bulky and expensive than are light microscopes. 3.1 • How Cells Are Studied 57
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FIGURE 3.3 (a)Salmonellabacteria are viewed with a light microscope. (b) This scanning electron micrograph showsSalmonellabacteria (in red) invading human cells. (credit a: modification of work by CDC, Armed Forces Institute of Pathology, Charles N. Farmer; credit b: modification of work by Rocky Mountain Laboratories, NIAID, NIH; scale-bar data from Matt Russell) CAREER C ONNE CTION Cytotechnologist Have you ever heard of a medical test called a Pap smear (Figure 3.4)? In this test, a doctor takes a small sample of cells from the uterine cervix of a patient and sends it to a medical lab where a cytotechnologist stains the cells and examines them for any changes that could indicate cervical cancer or a microbial infection. Cytotechnologists (cyto- = cell) are professionals who study cells through microscopic examinations and other laboratory tests. They are trained to determine which cellular changes are within normal limits or are abnormal.
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laboratory tests. They are trained to determine which cellular changes are within normal limits or are abnormal. Their focus is not limited to cervical cells; they study cellular specimens that come from all organs. When they notice abnormalities, they consult a pathologist, who is a medical doctor who can make a clinical diagnosis. Cytotechnologists play vital roles in saving people’s lives. When abnormalities are discovered early, a patient’s treatment can begin sooner, which usually increases the chances of successful treatment. 58 3 • Cell Structure and Function Access for free at openstax.org
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FIGURE 3.4 These uterine cervix cells, viewed through a light microscope, were obtained from a Pap smear. Normal cells are on the left. The cells on the right are infected with human papillomavirus. (credit: modification of work by Ed Uthman; scale-bar data from Matt Russell) Cell Theory The microscopes we use today are far more complex than those used in the 1600s by Antony van Leeuwenhoek, a Dutch shopkeeper who had great skill in crafting lenses. Despite the limitations of his now-ancient lenses, van Leeuwenhoek observed the movements of protists (a type of single-celled organism) and sperm, which he collectively termed “animalcules.” In a 1665 publication calledMicrographia, experimental scientist Robert Hooke coined the term “cell” (from the Latincella, meaning “small room”) for the box-like structures he observed when viewing cork tissue through a lens. In the 1670s, van Leeuwenhoek discovered bacteria and protozoa. Later advances in lenses and microscope
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In the 1670s, van Leeuwenhoek discovered bacteria and protozoa. Later advances in lenses and microscope construction enabled other scientists to see different components inside cells. By the late 1830s, botanist Matthias Schleiden and zoologist Theodor Schwann were studying tissues and proposed theunified cell theory, which states that all living things are composed of one or more cells, that the cell is the basic unit of life, and that all new cells arise from existing cells. These principles still stand today. 3.2 Comparing Prokaryotic and Eukaryotic Cells LEARNING OBJECTIVES By the end of this section, you will be able to: • Name examples of prokaryotic and eukaryotic organisms • Compare and contrast prokaryotic cells and eukaryotic cells • Describe the relative sizes of different kinds of cells Cells fall into one of two broad categories: prokaryotic and eukaryotic. The predominantly single-celled organisms of
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• Describe the relative sizes of different kinds of cells Cells fall into one of two broad categories: prokaryotic and eukaryotic. The predominantly single-celled organisms of the domains Bacteria and Archaea are classified as prokaryotes (pro- = before; -karyon- = nucleus). Animal cells, plant cells, fungi, and protists are eukaryotes (eu- = true). Components of Prokaryotic Cells All cells share four common components: 1) a plasma membrane, an outer covering that separates the cell’s interior from its surrounding environment; 2) cytoplasm, consisting of a jelly-like region within the cell in which other cellular components are found; 3) DNA, the genetic material of the cell; and 4) ribosomes, particles that synthesize proteins. However, prokaryotes differ from eukaryotic cells in several ways. Aprokaryotic cellis a simple, single-celled (unicellular) organism that lacks a nucleus, or any other membrane-
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proteins. However, prokaryotes differ from eukaryotic cells in several ways. Aprokaryotic cellis a simple, single-celled (unicellular) organism that lacks a nucleus, or any other membrane- bound organelle. We will shortly come to see that this is significantly different in eukaryotes. Prokaryotic DNA is found in the central part of the cell: a darkened region called the nucleoid (Figure 3.5). 3.2 • Comparing Prokaryotic and Eukaryotic Cells 59
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FIGURE 3.5 This figure shows the generalized structure of a prokaryotic cell. Unlike Archaea and eukaryotes, bacteria have a cell wall made of peptidoglycan, comprised of sugars and amino acids, and many have a polysaccharide capsule (Figure 3.5). The cell wall acts as an extra layer of protection, helps the cell maintain its shape, and prevents dehydration. The capsule enables the cell to attach to surfaces in its environment. Some prokaryotes have flagella, pili, or fimbriae. Flagella are used for locomotion, while most pili are used to exchange genetic material during a type of reproduction called conjugation. Eukaryotic Cells In nature, the relationship between form and function is apparent at all levels, including the level of the cell, and this will become clear as we explore eukaryotic cells. The principle “form follows function” is found in many contexts. For example, birds and fish have streamlined bodies that allow them to move quickly through the medium in which
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For example, birds and fish have streamlined bodies that allow them to move quickly through the medium in which they live, be it air or water. It means that, in general, one can deduce the function of a structure by looking at its form, because the two are matched. Aeukaryotic cellis a cell that has a membrane-bound nucleus and other membrane-bound compartments or sacs, calledorganelles, which have specialized functions. The word eukaryotic means “true kernel” or “true nucleus,” alluding to the presence of the membrane-bound nucleus in these cells. The word “organelle” means “little organ,” and, as already mentioned, organelles have specialized cellular functions, just as the organs of your body have specialized functions. Cell Size At 0.1–5.0 µm in diameter, prokaryotic cells are significantly smaller than eukaryotic cells, which have diameters ranging from 10–100 µm (Figure 3.6). The small size of prokaryotes allows ions and organic molecules that enter
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ranging from 10–100 µm (Figure 3.6). The small size of prokaryotes allows ions and organic molecules that enter them to quickly spread to other parts of the cell. Similarly, any wastes produced within a prokaryotic cell can quickly move out. However, larger eukaryotic cells have evolved different structural adaptations to enhance cellular transport. Indeed, the large size of these cells would not be possible without these adaptations. In general, cell size is limited because volume increases much more quickly than does cell surface area. As a cell becomes larger, it becomes more and more difficult for the cell to acquire sufficient materials to support the processes inside the cell, because the relative size of the surface area across which materials must be transported declines. 60 3 • Cell Structure and Function Access for free at openstax.org
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FIGURE 3.6 This figure shows the relative sizes of different kinds of cells and cellular components. An adult human is shown for comparison. 3.3 Eukaryotic Cells LEARNING OBJECTIVES By the end of this section, you will be able to: • Describe the structure of eukaryotic plant and animal cells • State the role of the plasma membrane • Summarize the functions of the major cell organelles • Describe the cytoskeleton and extracellular matrix At this point, it should be clear that eukaryotic cells have a more complex structure than do prokaryotic cells. Organelles allow for various functions to occur in the cell at the same time. Before discussing the functions of organelles within a eukaryotic cell, let us first examine two important components of the cell: the plasma membrane and the cytoplasm. 3.3 • Eukaryotic Cells 61
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VISU AL CONNE CTION FIGURE 3.7 This figure shows (a) a typical animal cell and (b) a typical plant cell. What structures does a plant cell have that an animal cell does not have? What structures does an animal cell have that a plant cell does not have? 62 3 • Cell Structure and Function Access for free at openstax.org
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The Plasma Membrane Like prokaryotes, eukaryotic cells have aplasma membrane(Figure 3.8) made up of a phospholipid bilayer with embedded proteins that separates the internal contents of the cell from its surrounding environment. A phospholipid is a lipid molecule composed of two fatty acid chains, a glycerol backbone, and a phosphate group. The plasma membrane regulates the passage of some substances, such as organic molecules, ions, and water, preventing the passage of some to maintain internal conditions, while actively bringing in or removing others. Other compounds move passively across the membrane. FIGURE 3.8 The plasma membrane is a phospholipid bilayer with embedded proteins. There are other components, such as cholesterol and carbohydrates, which can be found in the membrane in addition to phospholipids and protein. The plasma membranes of cells that specialize in absorption are folded into fingerlike projections called microvilli
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The plasma membranes of cells that specialize in absorption are folded into fingerlike projections called microvilli (singular = microvillus). This folding increases the surface area of the plasma membrane. Such cells are typically found lining the small intestine, the organ that absorbs nutrients from digested food. This is an excellent example of form matching the function of a structure. People with celiac disease have an immune response to gluten, which is a protein found in wheat, barley, and rye. The immune response damages microvilli, and thus, afflicted individuals cannot absorb nutrients. This leads to malnutrition, cramping, and diarrhea. Patients suffering from celiac disease must follow a gluten-free diet. The Cytoplasm Thecytoplasmcomprises the contents of a cell between the plasma membrane and the nuclear envelope (a structure to be discussed shortly). It is made up of organelles suspended in the gel-likecytosol, the cytoskeleton,
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structure to be discussed shortly). It is made up of organelles suspended in the gel-likecytosol, the cytoskeleton, and various chemicals (Figure 3.7). Even though the cytoplasm consists of 70 to 80 percent water, it has a semi- solid consistency, which comes from the proteins within it. However, proteins are not the only organic molecules found in the cytoplasm. Glucose and other simple sugars, polysaccharides, amino acids, nucleic acids, fatty acids, and derivatives of glycerol are found there too. Ions of sodium, potassium, calcium, and many other elements are also dissolved in the cytoplasm. Many metabolic reactions, including protein synthesis, take place in the cytoplasm. The Cytoskeleton 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
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components left? No. Within the cytoplasm, there would still be ions and organic molecules, plus a network of protein fibers that helps to maintain the shape of the cell, secures certain organelles in specific positions, allows cytoplasm and vesicles to move within the cell, and enables unicellular organisms to move independently. Collectively, this network of protein fibers is known as thecytoskeleton. There are three types of fibers within the cytoskeleton: microfilaments, also known as actin filaments, intermediate filaments, and microtubules (Figure 3.9). 3.3 • Eukaryotic Cells 63
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FIGURE 3.9 Microfilaments, intermediate filaments, and microtubules compose a cell’s cytoskeleton. Microfilaments are the thinnest of the cytoskeletal fibers and function in moving cellular components, for example, during cell division. They also maintain the structure of microvilli, the extensive folding of the plasma membrane found in cells dedicated to absorption. These components are also common in muscle cells and are responsible for muscle cell contraction. Intermediate filaments are of intermediate diameter and have structural functions, such as maintaining the shape of the cell and anchoring organelles. Keratin, the compound that strengthens hair and nails, forms one type of intermediate filament. Microtubules are the thickest of the cytoskeletal fibers. These are hollow tubes that can dissolve and reform quickly. Microtubules guide organelle movement and are the structures that pull
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tubes that can dissolve and reform quickly. Microtubules guide organelle movement and are the structures that pull chromosomes to their poles during cell division. They are also the structural components of flagella and cilia. In cilia and flagella, the microtubules are organized as a circle of nine double microtubules on the outside and two microtubules in the center. The centrosome is a region near the nucleus of animal cells that functions as a microtubule-organizing center. It contains a pair of centrioles, two structures that lie perpendicular to each other. Each centriole is a cylinder of nine triplets of microtubules. The centrosome replicates itself before a cell divides, and the centrioles play a role in pulling the duplicated chromosomes to opposite ends of the dividing cell. However, the exact function of the centrioles in cell division is not clear, since cells that have the centrioles removed can still divide, and plant cells, which lack centrioles, are
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not clear, since cells that have the centrioles removed can still divide, and plant cells, which lack centrioles, are capable of cell division. Flagella and Cilia Flagella(singular = flagellum) are long, hair-like structures that extend from the plasma membrane and are used to move an entire cell, (for example, sperm,Euglena). When present, the cell has just one flagellum or a few flagella. Whencilia(singular = cilium) are present, however, they are many in number and extend along the entire surface of the plasma membrane. They are short, hair-like structures that are used to move entire cells (such as paramecium) or move 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 move particulate matter toward the throat that mucus has trapped). 64 3 • Cell Structure and Function Access for free at openstax.org
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The Endomembrane System Theendomembrane system(endo= within) is a group of membranes and organelles (Figure 3.13) in eukaryotic cells that work together to modify, package, and transport lipids and proteins. It includes the nuclear envelope, lysosomes, and vesicles, the endoplasmic reticulum and Golgi apparatus, which we will cover shortly. Although not technicallywithinthe cell, the plasma membrane is included in the endomembrane system because, as you will see, it interacts with the other endomembranous organelles. The Nucleus Typically, the nucleus is the most prominent organelle in a cell (Figure 3.7). Thenucleus(plural = nuclei) houses the cell’s DNA in the form of chromatin and directs the synthesis of ribosomes and proteins. Let us look at it in more detail (Figure 3.10). FIGURE 3.10 The outermost boundary of the nucleus is the nuclear envelope. Notice that the nuclear envelope consists of two
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detail (Figure 3.10). FIGURE 3.10 The outermost boundary of the nucleus is the nuclear envelope. Notice that the nuclear envelope consists of two phospholipid bilayers (membranes)—an outer membrane and an inner membrane—in contrast to the plasma membrane (Figure 3.8), which consists of only one phospholipid bilayer. (credit: modification of work by NIGMS, NIH) Thenuclear envelopeis a double-membrane structure that constitutes the outermost portion of the nucleus (Figure 3.10). Both the inner and outer membranes of the nuclear envelope are phospholipid bilayers. The nuclear envelope is punctuated with pores that control the passage of ions, molecules, and RNA between the nucleoplasm and the cytoplasm. To understandchromatin, it is helpful to first consider chromosomes. Achromosomeis a structure within the nucleus that is made up of DNA, the hereditary material, and proteins. This combination of DNA and proteins is
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nucleus that is made up of DNA, the hereditary material, and proteins. This combination of DNA and proteins is called chromatin. In eukaryotes, chromosomes are linear structures. Every species has a specific number of chromosomes in the nucleus of its body cells. For example, in humans, the chromosome number is 46, whereas in fruit flies, the chromosome number is eight. Chromosomes are only visible and distinguishable from one another when the cell is getting ready to divide. When the cell is in the growth and maintenance phases of its life cycle, the chromosomes resemble an unwound, jumbled bunch of threads. We already know that the nucleus directs the synthesis of ribosomes, but how does it do this? Some chromosomes have sections of DNA that encode ribosomal RNA. A darkly staining area within the nucleus, called thenucleolus (plural = nucleoli), aggregates the ribosomal RNA with associated proteins to assemble the ribosomal subunits that
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(plural = nucleoli), aggregates the ribosomal RNA with associated proteins to assemble the ribosomal subunits that are then transported through the nuclear pores into the cytoplasm. The Endoplasmic Reticulum Theendoplasmic reticulum (ER)(Figure 3.13) is a series of interconnected membranous tubules that collectively modify proteins and synthesize lipids. However, these two functions are performed in separate areas of the endoplasmic reticulum: the rough endoplasmic reticulum and the smooth endoplasmic reticulum, respectively. The hollow portion of the ER tubules is called the lumen or cisternal space. The membrane of the ER, which is a phospholipid bilayer embedded with proteins, is continuous with the nuclear envelope. 3.3 • Eukaryotic Cells 65
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Therough endoplasmic reticulum (RER)is so named because the ribosomes attached to its cytoplasmic surface give it a studded appearance when viewed through an electron microscope. The ribosomes synthesize proteins while attached to the ER, resulting in transfer of their newly synthesized proteins into the lumen of the RER where they undergo modifications such as folding or addition of sugars. The RER also makes phospholipids for cell membranes. If the phospholipids or modified proteins are not destined to stay in the RER, they will be packaged within vesicles and transported from the RER by budding from the membrane (Figure 3.13). Since the RER is engaged in modifying proteins that will be secreted from the cell, it is abundant in cells that secrete proteins, such as the liver. Thesmooth endoplasmic reticulum (SER)is continuous with the RER but has few or no ribosomes on its cytoplasmic surface (seeFigure 3.7). The SER’s functions include synthesis of carbohydrates, lipids (including
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cytoplasmic surface (seeFigure 3.7). The SER’s functions include synthesis of carbohydrates, lipids (including phospholipids), and steroid hormones; detoxification of medications and poisons; alcohol metabolism; and storage of calcium ions. The Golgi Apparatus We have already mentioned that vesicles can bud from the ER, but where do the vesicles go? Before reaching their final destination, the lipids or proteins within the transport vesicles need to be sorted, packaged, and tagged so that they wind up in the right place. The sorting, tagging, packaging, and distribution of lipids and proteins take place in theGolgi apparatus(also called the Golgi body), a series of flattened membranous sacs (Figure 3.11). FIGURE 3.11 The Golgi apparatus in this transmission electron micrograph of a white blood cell is visible as a stack of semicircular flattened rings in the lower portion of this image. Several vesicles can be seen near the Golgi apparatus. (credit: modification of work by
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flattened rings in the lower portion of this image. Several vesicles can be seen near the Golgi apparatus. (credit: modification of work by Louisa Howard; scale-bar data from Matt Russell) The Golgi apparatus has a receiving face near the endoplasmic reticulum and a releasing face on the side away from the ER, toward the cell membrane. The transport vesicles that form from the ER travel to the receiving face, fuse with it, and empty their contents into the lumen of the Golgi apparatus. As the proteins and lipids travel through the Golgi, they undergo further modifications. The most frequent modification is the addition of short chains of sugar molecules. The newly modified proteins and lipids are then tagged with small molecular groups to enable them to be routed to their proper destinations. Finally, the modified and tagged proteins are packaged into vesicles that bud from the opposite face of the Golgi.
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be routed to their proper destinations. Finally, the modified and tagged proteins are packaged into vesicles that bud from the opposite face of the Golgi. While some of these vesicles, transport vesicles, deposit their contents into other parts of the cell where they will be used, others, secretory vesicles, fuse with the plasma membrane and release their contents outside the cell. The amount of Golgi in different cell types again illustrates that form follows function within cells. Cells that engage in a great deal of secretory activity (such as cells of the salivary glands that secrete digestive enzymes or cells of the immune system that secrete antibodies) have an abundant number of Golgi. In plant cells, the Golgi has an additional role of synthesizing polysaccharides, some of which are incorporated into the cell wall and some of which are used in other parts of the cell. Lysosomes
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In plant cells, the Golgi has an additional role of synthesizing polysaccharides, some of which are incorporated into the cell wall and some of which are used in other parts of the cell. Lysosomes In animal cells, thelysosomesare the cell’s “garbage disposal.” Digestive enzymes within the lysosomes aid the breakdown of proteins, polysaccharides, lipids, nucleic acids, and even worn-out organelles. In single-celled 66 3 • Cell Structure and Function Access for free at openstax.org
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eukaryotes, lysosomes are important for digestion of the food they ingest and the recycling of organelles. These enzymes are active at a much lower pH (more acidic) than those located in the cytoplasm. Many reactions that take place in the cytoplasm could not occur at a low pH, thus the advantage of compartmentalizing the eukaryotic cell into organelles is apparent. Lysosomes also use their hydrolytic enzymes to destroy disease-causing organisms that might enter the cell. A good example of this occurs in a group of white blood cells called macrophages, which are part of your body’s immune system. In a process known as phagocytosis, a section of the plasma membrane of the macrophage invaginates (folds in) and engulfs a pathogen. The invaginated section, with the pathogen inside, then pinches itself off from the plasma membrane and becomes a vesicle. The vesicle fuses with a lysosome. The lysosome’s hydrolytic enzymes then destroy the pathogen (Figure 3.12).
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plasma membrane and becomes a vesicle. The vesicle fuses with a lysosome. The lysosome’s hydrolytic enzymes then destroy the pathogen (Figure 3.12). FIGURE 3.12 A macrophage has phagocytized a potentially pathogenic bacterium into a vesicle, which then fuses with a lysosome within the cell so that the pathogen can be destroyed. Other organelles are present in the cell, but for simplicity, are not shown. Vesicles and Vacuoles Vesiclesandvacuolesare membrane-bound sacs that function in storage and transport. Vacuoles are somewhat larger than vesicles, and the membrane of a vacuole does not fuse with the membranes of other cellular components. Vesicles can fuse with other membranes within the cell system. Additionally, enzymes within plant vacuoles can break down macromolecules. 3.3 • Eukaryotic Cells 67
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VISU AL CONNE CTION FIGURE 3.13 The endomembrane system works to modify, package, and transport lipids and proteins. (credit: modification of work by Magnus Manske) Why does thecisface of the Golgi not face the plasma membrane? Ribosomes Ribosomesare the cellular structures responsible for protein synthesis. When viewed through an electron microscope, free ribosomes appear as either clusters or single tiny dots floating freely in the cytoplasm. Ribosomes may be attached to either the cytoplasmic side of the plasma membrane or the cytoplasmic side of the endoplasmic reticulum (Figure 3.7). Electron microscopy has shown that ribosomes consist of large and small subunits. Ribosomes are enzyme complexes that are responsible for protein synthesis. Because protein synthesis is essential for all cells, ribosomes are found in practically every cell, although they are smaller in prokaryotic cells. They are particularly abundant in immature red blood cells for the synthesis of
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smaller in prokaryotic cells. They are particularly abundant in immature red blood cells for the synthesis of hemoglobin, which functions in the transport of oxygen throughout the body. Mitochondria Mitochondria(singular = mitochondrion) are often called the “powerhouses” or “energy factories” of a cell because they are responsible for making adenosine triphosphate (ATP), the cell’s main energy-carrying molecule. The formation of ATP from the breakdown of glucose is known as cellular respiration. Mitochondria are oval-shaped, double-membrane organelles (Figure 3.14) that have their own ribosomes and DNA. Each membrane is a phospholipid bilayer embedded with proteins. The inner layer has folds called cristae, which increase the surface area of the inner membrane. The area surrounded by the folds is called the mitochondrial matrix. The cristae and the matrix have different roles in cellular respiration.
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area of the inner membrane. The area surrounded by the folds is called the mitochondrial matrix. The cristae and the matrix have different roles in cellular respiration. In keeping with our theme of form following function, it is important to point out that muscle cells have a very high concentration of mitochondria because muscle cells need a lot of energy to contract. 68 3 • Cell Structure and Function Access for free at openstax.org
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FIGURE 3.14 This transmission electron micrograph shows a mitochondrion as viewed with an electron microscope. Notice the inner and outer membranes, the cristae, and the mitochondrial matrix. (credit: modification of work by Matthew Britton; scale-bar data from Matt Russell) Peroxisomes Peroxisomesare small, round organelles enclosed by single membranes. They carry out oxidation reactions that break down fatty acids and amino acids. They also detoxify many poisons that may enter the body. Alcohol is detoxified by peroxisomes in liver cells. A byproduct of these oxidation reactions is hydrogen peroxide, H2O2, which is contained within the peroxisomes to prevent the chemical from causing damage to cellular components outside of the organelle. Hydrogen peroxide is safely broken down by peroxisomal enzymes into water and oxygen. Animal Cells versus Plant Cells Despite their fundamental similarities, there are some striking differences between animal and plant cells (seeTable
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Animal Cells versus Plant Cells Despite their fundamental similarities, there are some striking differences between animal and plant cells (seeTable 3.1). Animal cells have centrioles, centrosomes (discussed under the cytoskeleton), and lysosomes, whereas plant cells do not. Plant cells have a cell wall, chloroplasts, plasmodesmata, and plastids used for storage, and a large central vacuole, whereas animal cells do not. The Cell Wall InFigure 3.7b, the diagram of a plant cell, you see a structure external to the plasma membrane called the cell wall. Thecell wallis a rigid covering that protects the cell, provides structural support, and gives shape to the cell. Fungal and protist cells also have cell walls. While the chief component of prokaryotic cell walls is peptidoglycan, the major organic molecule in the plant cell wall is cellulose, a polysaccharide made up of long, straight chains of glucose units. When nutritional information
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wall is cellulose, a polysaccharide made up of long, straight chains of glucose units. When nutritional information refers to dietary fiber, it is referring to the cellulose content of food. Chloroplasts Like mitochondria, chloroplasts also have their own DNA and ribosomes.Chloroplastsfunction in photosynthesis and can be found in eukaryotic cells such as plants and algae. In photosynthesis, carbon dioxide, water, and light energy are used to make glucose and oxygen. This is the major difference between plants and animals: Plants (autotrophs) are able to make their own food, like glucose, whereas animals (heterotrophs) must rely on other organisms for their organic compounds or food source. Like mitochondria, chloroplasts have outer and inner membranes, but within the space enclosed by a chloroplast’s inner membrane is a set of interconnected and stacked, fluid-filled membrane sacs called thylakoids (Figure 3.15).
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inner membrane is a set of interconnected and stacked, fluid-filled membrane sacs called thylakoids (Figure 3.15). Each stack of thylakoids is called a granum (plural = grana). The fluid enclosed by the inner membrane and surrounding the grana is called the stroma. 3.3 • Eukaryotic Cells 69
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FIGURE 3.15 This simplified diagram of a chloroplast shows the outer membrane, inner membrane, thylakoids, grana, and stroma. The chloroplasts contain a green pigment called chlorophyll, which captures the energy of sunlight for photosynthesis. Like plant cells, photosynthetic protists also have chloroplasts. Some bacteria also perform photosynthesis, but they do not have chloroplasts. Their photosynthetic pigments are located in the thylakoid membrane within the cell itself. EVOLUTION C ONNE CTION Endosymbiosis We have mentioned that both mitochondria and chloroplasts contain DNA and ribosomes. Have you wondered why? Strong evidence points to endosymbiosis as the explanation. Symbiosis is a relationship in which organisms from two separate species live in close association and typically exhibit specific adaptations to each other. Endosymbiosis (endo-= within) is a relationship in which one organism
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exhibit specific adaptations to each other. Endosymbiosis (endo-= within) is a relationship in which one organism lives inside the other. Endosymbiotic relationships abound in nature. Microbes that produce vitamin K live inside the human gut. This relationship is beneficial for us because we are unable to synthesize vitamin K. It is also beneficial for the microbes because they are protected from other organisms and are provided a stable habitat and abundant food by living within the large intestine. Scientists have long noticed that bacteria, mitochondria, and chloroplasts are similar in size and have other similar features. In the 1950s and 1960s, scientists discovered that mitochondria and chloroplasts have their own DNA and ribosomes, just as bacteria do. In 1967, Lynn Margulis used microbial evidence in her proposal of endosymbiotic theory, which indicated that these organelles originated from separate organisms. Although Margulis’s work was
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theory, which indicated that these organelles originated from separate organisms. Although Margulis’s work was met with resistance, this basic component of this once-revolutionary hypothesis is now widely accepted. Scientists believe that host cells and bacteria formed a mutually beneficial endosymbiotic relationship when the host cells ingested aerobic bacteria and cyanobacteria but did not destroy them. Through evolution, these ingested bacteria became more specialized in their functions, with the aerobic bacteria becoming mitochondria and the photosynthetic bacteria becoming chloroplasts. The Central Vacuole Previously, we mentioned vacuoles as essential components of plant cells. If you look atFigure 3.7, you will see that plant cells each have a large, central vacuole that occupies most of the cell. Thecentral vacuoleplays a key role in regulating the cell’s concentration of water in changing environmental conditions. In plant cells, the liquid inside the
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regulating the cell’s concentration of water in changing environmental conditions. In plant cells, the liquid inside the central vacuole provides turgor pressure, which is the outward pressure caused by the fluid inside the cell. Have you ever noticed that if you forget to water a plant for a few days, it wilts? That is because as the water concentration in the soil becomes lower than the water concentration in the plant, water moves out of the central vacuoles and cytoplasm and into the soil. As the central vacuole shrinks, it leaves the cell wall unsupported. This loss of support to the cell walls of a plant results in the wilted appearance. Additionally, this fluid has a very bitter taste, which discourages consumption by insects and animals. The central vacuole also functions to store proteins in developing 70 3 • Cell Structure and Function Access for free at openstax.org
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seed cells. Extracellular Matrix of Animal Cells Most animal cells release materials into the extracellular space. The primary components of these materials are glycoproteins and the protein collagen. Collectively, these materials are called theextracellular matrix(Figure 3.16). Not only does the extracellular matrix hold the cells together to form a tissue, but it also allows the cells within the tissue to communicate with each other. FIGURE 3.16 The extracellular matrix consists of a network of substances secreted by cells. Blood clotting provides an example of the role of the extracellular matrix in cell communication. When the cells lining a blood vessel are damaged, they display a protein receptor called tissue factor. When tissue factor binds with another factor in the extracellular matrix, it causes platelets to adhere to the wall of the damaged blood vessel, stimulates adjacent smooth muscle cells in the blood vessel to contract (thus constricting the blood vessel), and
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stimulates adjacent smooth muscle cells in the blood vessel to contract (thus constricting the blood vessel), and initiates a series of steps that stimulate the platelets to produce clotting factors. Intercellular Junctions Cells can also communicate with each other by direct contact, referred to as intercellular junctions. There are some differences in the ways that plant and animal cells do this.Plasmodesmata(singular = plasmodesma) are junctions between plant cells, whereas animal cell contacts include tight and gap junctions, and desmosomes. In general, long stretches of the plasma membranes of neighboring plant cells cannot touch one another because they are separated by the cell walls surrounding each cell. Plasmodesmata are numerous channels that pass between the cell walls of adjacent plant cells, connecting their cytoplasm and enabling signal molecules and nutrients to be transported from cell to cell (Figure 3.17a). 3.3 • Eukaryotic Cells 71
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FIGURE 3.17 There are four kinds of connections between cells. (a) A plasmodesma is a channel between the cell walls of two adjacent plant cells. (b) Tight junctions join adjacent animal cells. (c) Desmosomes join two animal cells together. (d) Gap junctions act as channels between animal cells. (credit b, c, d: modification of work by Mariana Ruiz Villareal) Atight junctionis a watertight seal between two adjacent animal cells (Figure 3.17b). Proteins hold the cells tightly against each other. This tight adhesion prevents materials from leaking between the cells. Tight junctions are typically found in the epithelial tissue that lines internal organs and cavities, and composes most of the skin. For example, the tight junctions of the epithelial cells lining the urinary bladder prevent urine from leaking into the extracellular space. Also found only in animal cells aredesmosomes, which act like spot welds between adjacent epithelial cells (Figure
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extracellular space. Also found only in animal cells aredesmosomes, which act like spot welds between adjacent epithelial cells (Figure 3.17c). They keep cells together in a sheet-like formation in organs and tissues that stretch, like the skin, heart, and muscles. Gap junctionsin animal cells are like plasmodesmata in plant cells in that they are channels between adjacent cells that allow for the transport of ions, nutrients, and other substances that enable cells to communicate (Figure 3.17d). Structurally, however, gap junctions and plasmodesmata differ. 72 3 • Cell Structure and Function Access for free at openstax.org
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Components of Prokaryotic and Eukaryotic Cells and Their Functions Cell Component Function Present in Prokaryotes? Present in Animal Cells? Present in Plant Cells? Plasma membrane Separates cell from external environment; controls passage of organic molecules, ions, water, oxygen, and wastes into and out of the cell Yes Yes Yes Cytoplasm Provides structure to cell; site of many metabolic reactions; medium in which organelles are found Yes Yes Yes Nucleoid Location of DNA Yes No No Nucleus Cell organelle that houses DNA and directs synthesis of ribosomes and proteins No Yes Yes Ribosomes Protein synthesis Yes Yes Yes Mitochondria ATP production/cellular respiration No Yes Yes Peroxisomes Oxidizes and breaks down fatty acids and amino acids, and detoxifies poisons No Yes Yes Vesicles and vacuoles Storage and transport; digestive function in plant cells No Yes Yes Centrosome Unspecified role in cell division in animal cells; organizing center of microtubules in animal cells No Yes No
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vacuoles Storage and transport; digestive function in plant cells No Yes Yes Centrosome Unspecified role in cell division in animal cells; organizing center of microtubules in animal cells No Yes No Lysosomes Digestion of macromolecules; recycling of worn-out organelles No Yes No Cell wall Protection, structural support and maintenance of cell shape Yes, primarily peptidoglycan in bacteria but not Archaea No Yes, primarily cellulose Chloroplasts Photosynthesis No No Yes Endoplasmic reticulum Modifies proteins and synthesizes lipids No Yes Yes Golgi apparatus Modifies, sorts, tags, packages, and distributes lipids and proteins No Yes Yes TABLE 3.1 3.3 • Eukaryotic Cells 73
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Cell Component Function Present in Prokaryotes? Present in Animal Cells? Present in Plant Cells? Cytoskeleton Maintains cell’s shape, secures organelles in specific positions, allows cytoplasm and vesicles to move within the cell, and enables unicellular organisms to move independently Yes Yes Yes Flagella Cellular locomotion Some Some No, except for some plant sperm. Cilia Cellular locomotion, movement of particles along extracellular surface of plasma membrane, and filtration No Some No TABLE 3.1 This table provides the components of prokaryotic and eukaryotic cells and their respective functions. 3.4 The Cell Membrane LEARNING OBJECTIVES By the end of this section, you will be able to: • Understand the fluid mosaic model of membranes • Describe the functions of phospholipids, proteins, and carbohydrates in membranes A cell’s plasma membrane defines the boundary of the cell and determines the nature of its contact with the
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• Describe the functions of phospholipids, proteins, and carbohydrates in membranes A cell’s plasma membrane defines the boundary of the cell and determines the nature of its contact with the environment. Cells exclude some substances, take in others, and excrete still others, all in controlled quantities. Plasma membranes enclose the borders of cells, but rather than being a static bag, they are dynamic and constantly in flux. The plasma membrane must be sufficiently flexible to allow certain cells, such as red blood cells and white blood cells, to change shape as they pass through narrow capillaries. These are the more obvious functions of a plasma membrane. In addition, the surface of the plasma membrane carries markers that allow cells to recognize one another, which is vital as tissues and organs form during early development, and which later plays a role in the “self” versus “non-self” distinction of the immune response.
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one another, which is vital as tissues and organs form during early development, and which later plays a role in the “self” versus “non-self” distinction of the immune response. The plasma membrane also carries receptors, which are attachment sites for specific substances that interact with the cell. Each receptor is structured to bind with a specific substance. For example, surface receptors of the membrane create changes in the interior, such as changes in enzymes of metabolic pathways. These metabolic pathways might be vital for providing the cell with energy, making specific substances for the cell, or breaking down cellular waste or toxins for disposal. Receptors on the plasma membrane’s exterior surface interact with hormones or neurotransmitters, and allow their messages to be transmitted into the cell. Some recognition sites are used by viruses as attachment points. Although they are highly specific, pathogens like viruses may evolve to exploit
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viruses as attachment points. Although they are highly specific, pathogens like viruses may evolve to exploit receptors to gain entry to a cell by mimicking the specific substance that the receptor is meant to bind. This specificity helps to explain why human immunodeficiency virus (HIV) or any of the five types of hepatitis viruses invade only specific cells. Fluid Mosaic Model In 1972, S. J. Singer and Garth L. Nicolson proposed a new model of the plasma membrane that, compared to earlier understanding, better explained both microscopic observations and the function of the plasma membrane. This was called thefluid mosaic model. The model has evolved somewhat over time, but still best accounts for the 74 3 • Cell Structure and Function Access for free at openstax.org
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structure and functions of the plasma membrane as we now understand them. The fluid mosaic model describes the structure of the plasma membrane as a mosaic of components—including phospholipids, cholesterol, proteins, and carbohydrates—in which the components are able to flow and change position, while maintaining the basic integrity of the membrane. Both phospholipid molecules and embedded proteins are able to diffuse rapidly and laterally in the membrane. The fluidity of the plasma membrane is necessary for the activities of certain enzymes and transport molecules within the membrane. Plasma membranes range from 5–10 nm thick. As a comparison, human red blood cells, visible via light microscopy, are approximately 8 µm thick, or approximately 1,000 times thicker than a plasma membrane. (Figure 3.18) FIGURE 3.18 The fluid mosaic model of the plasma membrane structure describes the plasma membrane as a fluid combination of phospholipids, cholesterol, proteins, and carbohydrates.
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FIGURE 3.18 The fluid mosaic model of the plasma membrane structure describes the plasma membrane as a fluid combination of phospholipids, cholesterol, proteins, and carbohydrates. The plasma membrane is made up primarily of a bilayer of phospholipids with embedded proteins, carbohydrates, glycolipids, and glycoproteins, and, in animal cells, cholesterol. The amount of cholesterol in animal plasma membranes regulates the fluidity of the membrane and changes based on the temperature of the cell’s environment. In other words, cholesterol acts as antifreeze in the cell membrane and is more abundant in animals that live in cold climates. The main fabric of the membrane is composed of two layers of phospholipid molecules, and the polar ends of these molecules (which look like a collection of balls in an artist’s rendition of the model) (Figure 3.18) are in contact with aqueous fluid both inside and outside the cell. Thus, both surfaces of the plasma membrane are hydrophilic. In
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aqueous fluid both inside and outside the cell. Thus, both surfaces of the plasma membrane are hydrophilic. In contrast, the interior of the membrane, between its two surfaces, is a hydrophobic or nonpolar region because of the fatty acid tails. This region has no attraction for water or other polar molecules. Proteins make up the second major chemical component of plasma membranes. Integral proteins are embedded in the plasma membrane and may span all or part of the membrane. Integral proteins may serve as channels or pumps to move materials into or out of the cell. Peripheral proteins are found on the exterior or interior surfaces of membranes, attached either to integral proteins or to phospholipid molecules. Both integral and peripheral proteins may serve as enzymes, as structural attachments for the fibers of the cytoskeleton, or as part of the cell’s recognition sites. Carbohydrates are the third major component of plasma membranes. They are always found on the exterior surface
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recognition sites. Carbohydrates are the third major component of plasma membranes. They are always found on the exterior surface of cells and are bound either to proteins (forming glycoproteins) or to lipids (forming glycolipids). These carbohydrate chains may consist of 2–60 monosaccharide units and may be either straight or branched. Along with peripheral proteins, carbohydrates form specialized sites on the cell surface that allow cells to recognize each other. EVOLUTION C ONNE CTION How Viruses Infect Specific Organs Specific glycoprotein molecules exposed on the surface of the cell membranes of host cells are exploited by many viruses to infect specific organs. For example, HIV is able to penetrate the plasma membranes of specific kinds of 3.4 • The Cell Membrane 75
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white blood cells called T-helper cells and monocytes, as well as some cells of the central nervous system. The hepatitis virus attacks only liver cells. These viruses are able to invade these cells, because the cells have binding sites on their surfaces that the viruses have exploited with equally specific glycoproteins in their coats. (Figure 3.19). The cell is tricked by the mimicry of the virus coat molecules, and the virus is able to enter the cell. Other recognition sites on the virus’s surface interact with the human immune system, prompting the body to produce antibodies. Antibodies are made in response to the antigens (or proteins associated with invasive pathogens). These same sites serve as places for antibodies to attach, and either destroy or inhibit the activity of the virus. Unfortunately, these sites on HIV are encoded by genes that change quickly, making the production of an effective vaccine against the virus very difficult. The virus population
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change quickly, making the production of an effective vaccine against the virus very difficult. The virus population within an infected individual quickly evolves through mutation into different populations, or variants, distinguished by differences in these recognition sites. This rapid change of viral surface markers decreases the effectiveness of the person’s immune system in attacking the virus, because the antibodies will not recognize the new variations of the surface patterns. FIGURE 3.19 HIV docks at and binds to the CD4 receptor, a glycoprotein on the surface of T cells, before entering, or infecting, the cell. (credit: modification of work by US National Institutes of Health/National Institute of Allergy and Infectious Diseases) 3.5 Passive Transport LEARNING OBJECTIVES By the end of this section, you will be able to: • Explain why and how passive transport occurs • Understand the processes of osmosis and diffusion
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3.5 Passive Transport LEARNING OBJECTIVES By the end of this section, you will be able to: • Explain why and how passive transport occurs • Understand the processes of osmosis and diffusion • Define tonicity and describe its relevance to passive transport Plasma membranes must allow certain substances to enter and leave a cell, while preventing harmful material from entering and essential material from leaving. In other words, plasma membranes areselectively permeable (semipermeable)—they allow some substances through but not others. If they were to lose this selectivity, the cell would no longer be able to sustain itself, and it would be destroyed. Some cells require larger amounts of specific substances than do other cells; they must have a way of obtaining these materials from the extracellular fluids. This 76 3 • Cell Structure and Function Access for free at openstax.org
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may happen passively, as certain materials move back and forth, or the cell may have special mechanisms that ensure transport. Most cells expend most of their energy, in the form of adenosine triphosphate (ATP), to create and maintain an uneven distribution of ions on the opposite sides of their membranes. The structure of the plasma membrane contributes to these functions, but it also presents some problems. The most direct forms of membrane transport are passive.Passive transportis a naturally occurring phenomenon and does not require the cell to expend energy to accomplish the movement. In passive transport, substances move from an area of higher concentration to an area of lower concentration in a process called diffusion. A physical space in which there is a different concentration of a single substance is said to have aconcentration gradient. Selective Permeability Plasma membranes are asymmetric, meaning that despite the mirror image formed by the phospholipids, the
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Selective Permeability Plasma membranes are asymmetric, meaning that despite the mirror image formed by the phospholipids, the interior of the membrane is not identical to the exterior of the membrane. Integral proteins that act as channels or pumps work in one direction. Carbohydrates, attached to lipids or proteins, are also found on the exterior surface of the plasma membrane. These carbohydrate complexes help the cell bind substances that the cell needs in the extracellular fluid. This adds considerably to the selective nature of plasma membranes. Recall that plasma membranes have hydrophilic and hydrophobic regions. This characteristic helps the movement of certain materials through the membrane and hinders the movement of others. Lipid-soluble material can easily slip through the hydrophobic lipid core of the membrane. Substances such as the fat-soluble vitamins A, D, E, and K
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slip through the hydrophobic lipid core of the membrane. Substances such as the fat-soluble vitamins A, D, E, and K readily pass through the plasma membranes in the digestive tract and other tissues. Fat-soluble drugs also gain easy entry into cells and are readily transported into the body’s tissues and organs. Molecules of oxygen and carbon dioxide have no charge and pass through by simple diffusion. Polar substances present problems for the membrane. While some polar molecules connect easily with the outside of a cell, they cannot readily pass through the lipid core of the plasma membrane. Additionally, whereas small ions could easily slip through the spaces in the mosaic of the membrane, their charge prevents them from doing so. Ions such as sodium, potassium, calcium, and chloride must have a special means of penetrating plasma membranes. Simple sugars and amino acids also need help with transport across plasma membranes. Diffusion
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Simple sugars and amino acids also need help with transport across plasma membranes. Diffusion Diffusionis a passive process of transport. A single substance tends to move from an area of high concentration to an area of low concentration until the concentration is equal across the space. You are familiar with diffusion of substances through the air. For example, think about someone opening a bottle of perfume in a room filled with people. The perfume is at its highest concentration in the bottle and is at its lowest at the edges of the room. The perfume vapor will diffuse, or spread away, from the bottle, and gradually, more and more people will smell the perfume as it spreads. Materials move within the cell’s cytosol by diffusion, and certain materials move through the plasma membrane by diffusion (Figure 3.20). Diffusion expends no energy. Rather the different concentrations of
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plasma membrane by diffusion (Figure 3.20). Diffusion expends no energy. Rather the different concentrations of materials in different areas are a form of potential energy, and diffusion is the dissipation of that potential energy as materials move down their concentration gradients, from high to low. FIGURE 3.20 Diffusion through a permeable membrane follows the concentration gradient of a substance, moving the substance from an area of high concentration to one of low concentration. (credit: modification of work by Mariana Ruiz Villarreal) 3.5 • Passive Transport 77
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Each separate substance in a medium, such as the extracellular fluid, has its own concentration gradient, independent of the concentration gradients of other materials. Additionally, each substance will diffuse according to that gradient. Several factors affect the rate of diffusion. • Extent of the concentration gradient: The greater the difference in concentration, the more rapid the diffusion. The closer the distribution of the material gets to equilibrium, the slower the rate of diffusion becomes. • Mass of the molecules diffusing: More massive molecules move more slowly, because it is more difficult for them to move between the molecules of the substance they are moving through; therefore, they diffuse more slowly. • Temperature: Higher temperatures increase the energy and therefore the movement of the molecules, increasing the rate of diffusion. • Solvent density: As the density of the solvent increases, the rate of diffusion decreases. The molecules slow
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increasing the rate of diffusion. • Solvent density: As the density of the solvent increases, the rate of diffusion decreases. The molecules slow down because they have a more difficult time getting through the denser medium. LINK T O LE ARNING For an animation of the diffusion process in action, viewthis short video(http://openstax.org/l/passive_trnsprt)on cell membrane transport. Facilitated transport Infacilitated transport, also called facilitated diffusion, material moves across the plasma membrane with the assistance of transmembrane proteins down a concentration gradient (from high to low concentration) without the expenditure of cellular energy. However, the substances that undergo facilitated transport would otherwise not diffuse easily or quickly across the plasma membrane. The solution to moving polar substances and other substances across the plasma membrane rests in the proteins that span its surface. The material being transported
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substances across the plasma membrane rests in the proteins that span its surface. The material being transported is first attached to protein or glycoprotein receptors on the exterior surface of the plasma membrane. This allows the material that is needed by the cell to be removed from the extracellular fluid. The substances are then passed to specific integral proteins that facilitate their passage, because they form channels or pores that allow certain substances to pass through the membrane. The integral proteins involved in facilitated transport are collectively referred to as transport proteins, and they function as either channels for the material or carriers. Osmosis Osmosisis the movement of free water molecules through a semipermeable membrane according to the water's concentration gradient across the membrane, which is inversely proportional to the solutes' concentration. Whereas
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concentration gradient across the membrane, which is inversely proportional to the solutes' concentration. Whereas diffusion transports material across membranes and within cells, osmosis transportsonly wateracross a membrane and the membrane limits the diffusion of solutes in the water. Osmosis is a special case of diffusion. Water, like other substances, moves from an area of high concentration of free water molecules to one of low free water molecule concentration. Imagine a beaker with a semipermeable membrane, separating the two sides or halves (Figure 3.21). On both sides of the membrane, the water level is the same, but there are different concentrations on each side of a dissolved substance, orsolute, that cannot cross the membrane. If the volume of the water is the same, but the concentrations of solute are different, then there are also different concentrations of water, the solvent, on either side of the membrane. 78 3 • Cell Structure and Function
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same, but the concentrations of solute are different, then there are also different concentrations of water, the solvent, on either side of the membrane. 78 3 • Cell Structure and Function Access for free at openstax.org
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FIGURE 3.21 In osmosis, water always moves from an area of higher concentration (of water) to one of lower concentration (of water). In this system, the solute cannot pass through the selectively permeable membrane. A principle of diffusion is that the molecules move around and will spread evenly throughout the medium if they can. However, only the material capable of getting through the membrane will diffuse through it. In this example, the solute cannot diffuse through the membrane, but the water can. Water has a concentration gradient in this system. Therefore, water will diffuse down its concentration gradient, crossing the membrane to the side where it is less concentrated. This diffusion of water through the membrane—osmosis—will continue until the concentration gradient of water goes to zero. Osmosis proceeds constantly in living systems. LINK T O LE ARNING Watch thisvideo(http://openstax.org/l/dispersion)that illustrates diffusion in hot versus cold solutions. Tonicity
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LINK T O LE ARNING Watch thisvideo(http://openstax.org/l/dispersion)that illustrates diffusion in hot versus cold solutions. Tonicity Tonicitydescribes the amount of solute in a solution. The measure of the tonicity of a solution, or the total amount of solutes dissolved in a specific amount of solution, is called itsosmolarity. Three terms—hypotonic, isotonic, and hypertonic—are used to relate the osmolarity of a cell to the osmolarity of the extracellular fluid that contains the cells. In ahypotonicsolution, such as tap water, the extracellular fluid has a lower concentration of solutes than the fluid inside the cell, and water enters the cell. (In living systems, the point of reference is always the cytoplasm, so the prefixhypo- means that the extracellular fluid has a lower concentration of solutes, or a lower osmolarity, than the cell cytoplasm.) It also means that the extracellular fluid has a higher concentration of water than does the cell.
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the cell cytoplasm.) It also means that the extracellular fluid has a higher concentration of water than does the cell. In this situation, water will follow its concentration gradient and enter the cell. This may cause an animal cell to burst, or lyse. In ahypertonicsolution (the prefixhyper- refers to the extracellular fluid having a higher concentration of solutes than the cell’s cytoplasm), the fluid contains less water than the cell does, such as seawater. Because the cell has a lower concentration of solutes, the water will leave the cell. In effect, the solute is drawing the water out of the cell. This may cause an animal cell to shrivel, or crenate. In anisotonicsolution, the extracellular fluid has the same osmolarity as the cell. If the concentration of solutes of the cell matches that of the extracellular fluid, there will be no net movement of water into or out of the cell. Blood
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the cell matches that of the extracellular fluid, there will be no net movement of water into or out of the cell. Blood cells in hypertonic, isotonic, and hypotonic solutions take on characteristic appearances (Figure 3.22). 3.5 • Passive Transport 79
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VISU AL CONNE CTION FIGURE 3.22 Osmotic pressure changes the shape of red blood cells in hypertonic, isotonic, and hypotonic solutions. (credit: modification of work by Mariana Ruiz Villarreal) A doctor injects a patient with what the doctor thinks is isotonic saline solution. The patient dies, and autopsy reveals that many red blood cells have been destroyed. Do you think the solution the doctor injected was really isotonic? Some organisms, such as plants, fungi, bacteria, and some protists, have cell walls that surround the plasma membrane and prevent cell lysis. The plasma membrane can only expand to the limit of the cell wall, so the cell will not lyse. In fact, the cytoplasm in plants is always slightly hypertonic compared to the cellular environment, and water will always enter a cell if water is available. This influx of water produces turgor pressure, which stiffens the
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