A Tour of the Cell You can probably identity the blue blobs in this beautiful micrograph as the nucll of the cells it depicts. But did you know that the brightly colored pink and green strands you also see form a cell's skeleton? These structures are part of a system of protein fibers called the cytoskeleton, Much tke the way your skeleton provides support and also How has our knowledge enables you to move, the cytoskeleton provides structural of cells grown? ‘support to a cell and allows some cells to craw! an others to ‘swim. But even stationary cells have movement: Many oftheir internal parts bustle about, often traveling on cytoskeletal “roads.” Later in the chapter you will learn more about the cytoskeleton and how ur knowiedge of its structures and functions has. frown. AS you will see, our understanding of nature offen goes hand in hand with the invention and refinement of instruments that extend our senses. This certainly aplies to how cells were first discovered. In 4665, Robert Hooke used a crude microscope to examine @ pieve of bark from an oak twee, Hooke compared the structures he saw to “ite fooms"—celulze in Latin—and the term cell stuck. His contemporary, Antonl van Leeuwenhock, working with more refined lenses, examined numerous subjects, from biood and sperm to pond water. He produced drawings and enthusiastic descriptions of his discoveries, such as the tiy “animalcules, very prettily amoving” he found in the scrapings from his teeth, Since the days of Hooke and Leeuwenhoek, improved microscopes and techniques have vastly expanded our view of the cll For example, fluorescently colored stains reveal the cytoskeleton in the cells pictured to the right. n this chapter, you will See mary micrographs using such techniques, and they wil often be paired with drawings that help emphasize specific details. Neither drawings nor micrographs, however, allow you to see the dynamic nature of living cals. For that, you need to look through a microscope or view videos. As you study the images in this chapter, keep in mind thatthe parts ofa cell are moving and interacting. Indeed, the phenomenon ‘we call fe emerges from the interactions ofthe many components of a cel Introduction to the Cell The Nucleus and Ribosomes (42-44) (45-46) Microscopes reveal the structures of ‘cols gonetc instructions are ‘celle—the fundamental units of Ite. housed inthe nucleus ‘and cared out by ‘ibosornes. zThe Endomembrane System (47-42) ‘The endomembrane system participates Inthe manufacture, distribution, and breakdown of materials. Energy-Converting Organelles (4.13-4.15) Mitochondria in all eukaryotic cells ‘and chloroplasts In plant ces function in energy processing. (x The Cytoskeleton and Cell Surfaces (4.16-4.22) Tre cytoskeleton and extracel ‘components provide support, moti ‘and funetonal connections.races Come oy 4.1 Microscopes reveal the world of the cell Before microscopes were first used in the 1600s, noone: as small as about 2 nanometers (nm), 100-old improvement knew that living organisms were composed of the tiny units over the light microscope. This high resolution has enabled wwe call cells. The first microscopes were light microscopes, biologists to explore cell ultzastuctue, the complex internal Uke the ones you may use ina biology laboratory. In alight anatomy of cell. Figures 4.48 and 4.4¢ show images microscope (LM), visible light is passed through a spec- produced by two kinds of electron microscopes. Imen, such asa microorganism or a thin slice of animal or plant tissue, and then theough glass lenses. The lenses bend the light in such a way that the image ofthe specimes ‘magnified a itis projected into your eye or a camera ‘Magnification i the nerease in an object’s image size compared with its actual size. Figure 4.1.8 shows a micrograph ofa single-celled organism called Paramecium, The notation “LM 230%" printed along the right edge tes you that this photograph was taken through alight microscope and that ‘he image is 230 times the actual size ofthe organism, This Paramecium is about 0.33 millimeter (mm) in Tength. Table 4.2. shows the most common units of length that biologists use, An important factor in microscopy is resolution, a mea- sure ofthe clarity of an image. Resolution is the ability to |A-Fleure 4.14 Ugh merogaph ofthe uneeitaroognism Paramectan distinguish two neazby objects as separate, For example, what You see asa single star in the sky may be resolved as twin Stars with a telescope, Each optical instrument —be itan eye, a telescope, ora microscope—has a limit to its resolution The human eye can distinguish points as close together as 0.1 mm, about the sizeof avery fine grain of sand. A typical Light microscope cannot resoive detail finer than about (0.2 micrometer (umm), about the size of the smallest bacterium, No matter how many times the image of such a small cel is ‘magnified, the light microscope cannot resolve the details ofits structure. Indeed, light microscopes can effectively magnify objects only about 1,000 times, From the time that Hooke discovered cells in 1665 until 1 1900s, biologists had only light micro- scopes for viewing cells, With these microscopes and various | staining techniques to increase contrast betwen parts of cells, these early biologists discovered microorganisms, an ‘mal and plant cells, and even some structures within cells. BY ‘he mid: 1800s, this accumulation of evidence le to the eel theory, which states that all living things are composed of cellsand that all cells come from other cells (Our knowledge of cell structure took a giant leap forward as biologists began using the electron microscope in the 1950s, Instead of using light, an electron microscope (EM) focuses ‘beam of electrons through a specimen or onto ts surface Electron microscopes can distinguish biological structures the middle of CUE Eire ‘A Figure 4.4 Transmision electron micrograph of Toxoplasma 00 cm= 4,000 m= 394 inches ‘Tis parasite of eats can be vanamatted to humans, causing the 1 contmeter em) ~ 10% m9 (0.01 oF 2/100 m)~ 0.4 nen stoease toxoplasmosis) 4 murat rm) = 40° (0.001 9 1/2,000 my) Lmerameter n= 10% m (0.000001 m= 10% ram 4 nanometer fm) = 10°? m= 4 56 curren | A Tour of the CallBiologists use the scanning electron microscope (SEM) to study the detailed architecture of cell surfaces. The SEM uses anelectron beam to scan the surface of ace or other sample, ‘which is usually coated with thin fl of gold. The beam excites electrons on the surface, and these electrons are then detected by a device that translates their pattern into an image projected onto a video seen, The scanning electron micrograph in Figure 4B highlights the numerous iia on Paramecium, pro- jections it uses for movement. Notice the indentation, called ‘he oral groove, through which food enters the cell. As you can sce, the SEM prodices images that look three-dimensional The transmission electron microscope (TEM) is used to study the details of internal cll structure. The TEM aimnsan electron beam through a very thin section ofa specimen, just asa light microscope aims beam of ight through a specimen, ‘The section is stained with atoms of heavy metals, which attach, tocertan cellular structures more than others. Electrons are Scattered by these more dense parts, and the image s created by the pattem of transmitted electrons. Instead of using glass lenses, both the SEM and TEM use electromagnets as lenses to bend the paths of the electrons, magnifying and focusing the mage onto a monitor. The transmission eleczon micrograph in Figure 4.1C shows internal details ofa single-celled organism called Taxoplasma SEMs and TEMs are initially black and white butare often artificially colorized, as they are here, to highlight corclarify structural features. Flectron microscopes have truly revolutionized the study of cellsand their structures. Nonetheless, they have not replaced the light microscope: Electron microscopes cannot be used to study living specimens because the methods used to prepare the specimen kill he cells, Fora biologist studying aliving pro- cess, such as the movement of Paramecium, alight microscope equipped with a video camera is more suitable than either an SEM ora TEM. There ae different types of light microscopy, and major tech- nical advances in the past several decades have greatly expand- ed our ability to visualize cells, Figure 4.40 shows Paramecium as seen using differential interference contrast microscopy. ‘This optical technique amplifies differences in density so that ‘the structures in living cells appear almost three-dimensional Other techniques use fluorescent stains that selectively bind tovarlous cellular molecules (see the chapter introduction) You will see many beautiful and illuminating examples of ‘microscopy inthis textbook. But even with the magnification ‘A Figure 4.20 biterentia interference contrast merograph of Porameciam shown beside each micrograph, its often hard to imagine just how small cells are. Figure 4.45 shows the size range of cells compared with objects both larger and smaller and the ‘optical instrument that allows us to view them, Notice that the scale along the left side of the figures logarithmic to accommodate the range of sizes shown, Starting at the top with 10 meters (am, each reference measurement matksa ten fold decrease in length. Most cells ae between 1 and 300 pm, 1 diameter (yellow region of the figure) and are therefore visible only with a microscope, Certain bacteria areas small, 0.2 pm and can barely be seen with alight microscope, whereas chicken eggs are large enough to be seen with the ‘unaided eye. A single nerve cell running from the base of your spinal cord to your big toe may be I min length, although it Js 0 thin you would still need a microscope to see it. In the next module, we explore why cells are so small, EP Wien ype of miroscope would you use to study (a) the anges in shape of a Iving human white blood cel (b) the ‘nest details of surface texture of a human hair; () the detlled structure of an organelle ina lver elt? soyesusunn (9) scons anda jess (a sebor a tom [suman height im (1,000 rm) E—Lenath of some nerve and muscle cells o1m (100 mm) F chicken egg 201m orm Fre9 000 S28) (1.0001) Es mecium 100 ym F=-Human 099 Most plant and srimal ets TOES Nucous Most bacteria E> 1 ym EEMtocrension um (0.900 0m) sooo Sas Envcsome ronins Lids small molecules SB i 10 nm 01 nm E=Atoms ‘A Figure 4.4 The size range o cols and related objec. Invoduetion tote colt 874.2 The small size of cells relates to the need to exchange materials across the plasma membrane As you saw in Figure 4.18, most cell are microscopic, Are there advantages to being so small? The logistics of carrying outa cell's functions appear to set both lower and upper limits on cell size, Atminimum,a cell must be large enough to house enough DNA, protein molecules, and structures to survive and reproduce, But why aren't most cells as large as chicken eggs? ‘The maximum size ofa cell isinfluenced by geometry—the need to have a surface area large enough to service the volume ofacell Active cell havea huge amount of traffic across thelr outer surface. Achicken egg cell isn't very active, btoncea chick embryo start to develop, the egg is divided into many ‘microscopic cells, each bounded by a membrane that allows the essential flow of oxygen, nutrients, and wastes across its surface Surface-tovVolume Ratio. Large cells have moze surface ayea than small cells, but they havea much smaller surface area relative to thee volume than small cells. Figure 4.28 illustrates this by comparing I large cube to 27 small ones Using arbitrary units of measurement, the total volume isthe same in both cases: 27 units (height x wide length). The total surface areas, however, are quite different. ‘cube has sxsdes; tus, its surface area is six times the area ofeach side theight % width). The surface area of the large cubes 54 units’, while the total surface area of all 27 cubes is to2units? 27 6 x 1 x 1), three times greater than the surface area ofthe large cube. Thus, the combined smaller cubes havea much greater surface-to-volume ratio than the large cube. How about those neurons that extend from the base of your spe to your tes? Very thin, elongated shapes also provide a large surface area relative oa cell's volume ‘The Plasma Membrane Sowhatisacell's surface ike? And how does it control the traffic of molecules across it? The plasma membrane, also referred toas the cell membrane, forms a flexible boundary between the living cell and is sur roundings. Forastructure that separates life from nonlife, this membranes amazingly thin Itwouldtakeastack of more than 8,000 plasma membranes to equal the thickness ofthis page And, as you have come to expect with all things biological, the structure ofthe plasma membrane correlates with ts function, Tertvetane [arnt rus Teotce sus teens Surace volume ratio 2 6 ‘A Figure 4.28 effet of col size on surface area and volume 5B curren | A Tourof the Call ydrophitc heads Hydrophobic Phospholipid Channel Hydrophilic Hydrophobic protein regions of regions of aproten a protein ‘A Figure 4.28 The stture of plasma membrane. Phospholipid molecules are well suited to thelr role asa ‘major constituent of biological membranes. Each phospho- lipid is composed of two distinct regions—a head with a neg: atively charged phosphate group and two nonpolat fatty acid tails (see Module 3.10) Phospholipids group together to form, a two-layer sheet called a phospholipid bilayer. As you can see inFigure 4.28, the phospholipids hydrophilic (water-loving) heads face outward, exposed to the aqueous solutions on both sides of a membrane, Theis hydrophobic (water-earing) tails point inward, mingling togetiner and shielded from water. Embedded in this lipid bilayer are diverse proteins, floating like icebergsin a phospholipid se. The regions ofthe proteins within the center of the membrane ae hydrophobic; the exte- rior sections exposed to water are hydrophillc. Iustrating our theme of EMEMALORTNTIND, the properties of the phospholipid bilayer and the proteins sus- pended int relate to the plasma membrane’s job asa traffic cop, regulating the flow of material into and out ofthe cel Nonpolar molecules, such as Op and CO, can easily move across the membrane’s hydrophobic interior. Some of the ‘membrane’s proteins form channels tunnels that shield ions and polar molecules as they pass through the hydrophobic cen- terofthe membrane, Sil other proves serve as pumps, using energy to actively transport moleculesinto or out of the cell ‘We will etumn to the structure and function of biological ‘membranes later (see Chapter In the next module, we consider other features common to all ellsand take a closer lookat the prokaryotic cells found in two ofthe three major ‘groups of organisms. To convince yourself that a small call has a greater surface area relative to volume than a large eel, compare the ‘surface-to-volume ratios of the large cube and one of the small cubes in Figure 4.24, (cin 5 Fc Ft oumen gene 9 = s0ie xr pe com cnepns) 9 9 oan ous Leg 949 Be ©4.3 Prokaryotic cells are structurally simpler than eukaryotic cells Cells are of two distinet types: prokaryotic and eukaryotic Prokaryotic cells were the firs to evolve and were Earth's sole inhabitants for more than 1.5 billion years. Evidence indicates that eukaryotic cells evolved from some of these ancestral cells about 1.8 billion years ago. Biologists recognize three domains or major groups of organisms. The microor sganisms placed in domains Bacteria and Archaea consist of prokaryotic cells. These organisms are known as prokaryotes, Allother forms of if are placed in domain Fukarya, They are ‘composed of eukaryotic cells and are referred to as eukaryotes. Fukaryotic cells are distinguished by having a membrane- enclosed nucleus, which houses most oftheir DNA, and many membrane-enclosed organelles that perform specific functions. Prokaryotic eels are smaller and simpler in structure oth types of cells, however, share certain basic In addition to being bounded by a plasma membrane, the interior of all cells is filled with a thick, eliylike fluid called cytosol, in which cellular components are suspended. ll cells have one or more chromosomes, which carry genes ‘made of DNA, They also contain ribosomes, tiny structures ‘hat make proteins according to instructions from the genes, ‘The inside of bath types of cells is called the cytoplasm. However, n eukaryotic cells, this term refers only to the region between the nucleus and the plasma membrane, Figure 4.3 explores the structure ofa generalized prokary otic cell, Notice thatthe DNA is coiled into a region called the mucleoid ("nuclews-ike”), but no membrane surrounds | the DNA. The ribosomes of prokaryotes are smaller and dif fer somewhat from those of eukaryotes. These molecular atures Fimbriae: atachmert structures on the surace of some prokaryotes é Ribosomes: stetures that synthesize proteins Nucleoia att wal Capsule Flagel “A Figure 4.3 & dagram (et) ond fection mictograph ng) o a typleal prokaryote el region where the cell's DNA is located (not encloses by a membrane) Plasma membrane: membrane enclosing the cytoplasm "gid structure outside the plasma membrane jallyiko outer coating of many prokaryotes locometion ‘organelles of some bactera differences are the basis for the ation of some antibiotics, which specifically target prokaryotic ribosomes. Thus, p tein synthesis can be blocked fo the bacterium that’s invaded, you, but not for you, the eukaryote who is taking the drug. Outside the plasma membrane of most prokaryotes isa fairly rigid, chemically complex cell wall. The wall protects ‘he cell and helps maintain its shape. Some antibiotics such aspenicllin, prevent the formation of these protective walls. Again, because your cells don’t have such walls, these antbi ties can kill invading bacteria without harming your cells. Certain prokaryotes have a sticky outer coat called a capsule around the cel wall, helping to glue the cells to surfaces or toother cells na colony: In addition to capsules, some pro- karyotes have surface projections. Short projections help attach prokaryotes to each other or their substrate. Longer projections called flagella (singular, flagellum) propel cel ‘hhrough ts liquid environment, Tetakes an electron microscope to see the internal details. ‘of any cell, and thisis especially true of prokaryotic cells, Notice that the TEM of the bacterium in Figure 4.3 hasa mag: nificatlon of 20,940x, Most prokaryotic cells are about one- tenth the sizeof atypical eukaryotic cell (Prokaryotes will be described in more detail in Chapter 16.) Eukaryotic cells are ‘de main focus ofthis chapter, so we turn to these next. st three features that are common to prokaryote and eukaryotic coli. List troe features that dir. seuesoas say teunowos ane pu 'soyoueB0 oosepue-ovesauu “ya Benes sotosouons suerguou buses one se 8c wee © Heicobactor por ‘abacteram tat Causes stomach ulcers Introduction tote colt 59Eukaryo' All eukaryoUe cells—whether from prolist (a diverse group of ‘mostly unicellular organisms), fungi, animals, or plants—are fundamentally similar to one another and profoundly diff ‘om prokaryotic cells Le’ look plant cel as representatives of the eukaryotes, Figure 4.4 isa diagram of a generalized animal cell, and Figure 4.45 shows a generalize plant cell. We colorcode the various structures in the diagrams for easier Identification, and you wil se miniature versions of these cells to orient you during our in-depth tour in the rest ofthe chapter. But no cells would look exactly ike these. For one thing, cells have mult ple copies ofall of these structures (except forthe nucl Your cells have hundreds of mitochondria and millions of sbosomes. A plant cell may have 30 chloroplasts packed inside. Cells also have different shapes and relative propor tions of cell parts, depending on their specialized functions. The most obvious hallmark ofa eukaryotic cellists muclews, Butitalso.contains various other organelles ("little organs ent ‘an animal cell anda rset for ysosome: Ct cells are partitioned into functional compartments Which perform specific tasks, just asthe cll seis wrapped in a membrane made of phospholipids and proteins that perform various functions, each organelles bounded by a membrane with a lipid and protein composition that suits its function The organelles and other structures of eukaryotic cells can be organized into four basic functional groups: (1) The nucleus and ribosomes carry out the genetic control of the cell, (2) Oxganelles involved in the manufacture, distribu: don, and breakdown of molecules include the endoplasmic reticulum, Golgi apparatus, lysosomes, vacuoles, and peroxl somes. (3) Mitochondria in all cells and chloroplasts in plant cells function in energy processing. (4) Structural support, movement, and communication between cells are the func: tions of the cytoskeleton, plasma membrane, and plant cell wall. The cellular components identified in these two figures will be examined in detail in the modules that follow. In essence, the internal membranes of a eukaryotic cell partition it into functional compartments in which many Mn ae CYTOSKELETON Pecie Centrosome with pairof centrioles =ofits chemical activities colle metabolism —take place. Infact, various enzymes essen: | for metabolic processes are built into the membranes of organelles. The fluid-filled spaces within such compart ‘ments are locations where specific chemical conditions are ‘maintained, These conditions vary among organelles and favor the metabolic processes occurring in each. For exam ple, while a part of the endoplasmic reticulum is engaged Jn making hormones, neighboring peroxisomes may be detoxifying harmful compounds and making hydrogen per- oxide (H;02) asa poisonous by-product of their activities. But because the 1,0; is confined within the peroxisomes, where itis converted to HO by resident enzymes, the rest of the cell is protected. Except for lysosomes and centrosomes, the organelles and other structures of animal cells are found in plant cells. Also, although some animal cells have Nagella or cila (not shown ‘in Figure 4.44), among plants, only the sperm cells of few species have flagella, A plant cell Figure 4.48) also has some structure: ‘animal cell lacks. For example, a plant cell has a rigid, rather ively called cellular tan Oe Entel smooth eri poate CYTOSKELETON ees es thick cell wall. Chemically different from prokaryotic cell wall, plant cell walls contain the polysaccharide cellulose, Plasmodesmata (singular, plasmodesma) are cytoplasmic channels through cel walls that connect adjacent cells. An Important organelle found in plant cells isthe chloroplas where photosynthesis occurs. Unique to plant cells sa large central vacuole, a compartment that stores wateranda variety of chemicals Bukaryotic cells contain nonmembranous st well. The cytoskeleton, which you were introduced to in the chapter introduction, is composed of different types of protein fibers that extend throughout the cell. And ribo- somes are found in the cytosol as well as attached to certain ‘membranes. After you preview these cell diagrams, let’s move to the first stop on our detailed tour ofthe eukaryotic cell—the nucleus. tures as ldentity the structures In the plant cll that are not present In the animal cel. sh ae root een ee aSYou just savra preview of the many intricate structures that can be found ina eukaryotic cell. Acell must build and ‘maintain these structures and also process energy to support lts work of transport, movement, and communication. But who isin charge ofthis bustling factory? Who stores the master plans, gives the orders, changes course in response to environmental input, and, when called on, makes another factory just hike itself? The cell's nucle command center 1 functions as this The nucleus contains the cell's genetic instructions encoded in DNA. These master plans control the cel’s activ ities by directing protein synthesis, The DNA is associated ‘with many proteins and organized into structures called cho: ‘mosomes, The proteins help coll these long DNA molecules. Indeed, the DNA of the 46 chromosomes in one of yous cells laid end to end would stretch toa length of more than 2m, Dut must coll up to it intoa nucleus only S pm in diameter When a cell is not dividing, this complex of proteins and DNA, called chromatin, appears as a diffuse mass within the nucleus, as shown in the TEM (eight half) and diagram (lett half) of amucleus in Figure 4.5, Asa cell prepares to divide, the DNA is copied so that each daughter cell can later receive an identical set of genetic instructions, Jus prior to cell division, the thin chromatin fibers coil up further, Becoming thick enough to be visible with a light microscope as the familiar separate structures you would probably recognize as chromosomes. roe crs Endoplasm ‘A Figure 4.5 A cos seeton ofthe nclous with 9 superposed TEM 62 curren | A Tour of the call SUC UCSC U CMe) The nucleus contains the cell's geneti structions Enclosing the nuclewsis a double membrane called the nuclear envelope. Fach of the two membranes is separate phospholipid bilayer with associated proteins, Similar in func tion to the plasma membrane, the nuclear envelope controls the flow of materials into and out of the nucleus. As you can see In the diagram ofa nucleus in Figure 4.5, the nuclear envelope is perforated with protein-lined pores. These pore regulate the entry and exit of large molecules and also connect with the cell's network of membranes called the endoplasmic reticulum. The mucleolus, a prominent stracture inthe nucleus, is the site where a special type of RNA called ribosomal RNA (FRNA) is synthesized according to instructions in the DNA, Proteins brought in from the cytoplasm are assembled with {hls 2RNA to form the subunits of ribosomes. These subunits then exit to the eytoplasm, where they will jon to form, functional ribosomes. Another type of RNA, messenger RNA (mRNA), directs protein synthesis. Essentially, mRNA is transcription of protein-syathesizing instructions written in a gene's DNA, (see Figure 10.7). The mRNA moves into the eytoplasm, where ribosomes sanslate it into the amino acid sequences of proteins, Let's look at ribosomes next @omive ne processes tat scrim he nls4.6 Ribosomes make proteins for use in the cell and for export the nucleusis the cell’s command center, then ribosomes are the machines that carry out those commands. Ribosomes ate the cellular components that use instructions from the nucleus, written in mRNA, to build proteins. Cells that ‘make alot of proteins have. large number of ribosomes For example, a cell in your pancreas that produces digestive enzymes may contain a few million ribosomes. What other structure is prominent in cells that are active in protein synthesis? Remember thatthe nucleolus in the nucleus isthe site where the subunits of ribosomes are assembled. ‘As shown In Figute 4.6, ibosomes are found in two locations in the cell. Free ribosomes are suspended in the cytosol, while bound ribosomes are attached to the outside of the endoplasmic reticulum or nuclear envelope. Fre and bound ribosomes are structurally identical, and they can function in either location, depending on the protein they are making. Most of the proteins made on free ribosomes function within the cytosol; examples are enzymes that catalyze the first steps of sugar breakdown for cellular respiration. In Module 48, you wll see how bound ribosomes make proteins ‘that will be exported from the cel. ‘Atte bottom right in Figure 4.6, you see how ribosomes Interact with messenger RNA (carrying the instructions. froma gene) to build a protein. The nucleotide sequence of an mRNA molecule is translated into the amino acid sequence ofa polypeptide. The pathway from DNA to RNA. to protein isa prime example of our theme ofthe flow of {Protein synthesis is explored in more detall Jn Chapter 103 Cea reticulum Ce era A Figure 4.6 The lcations and structure of ribosomes ‘What role do ribosomes play in carrying out the genet Instructions of aco? seavessou yo suononas oo prone Sled anaite soso MU ee) CL) 4.7 Many organelles are connected in the endomembrane system Ribosomes may be a cell's protein-making machines, but sunning a factory as complex as a cell requires infrastructure and many different departments that perform separate but ated functions. Internal membranes, a distinguishing feature of eukaryotic cells, are involved in most ofa cell’s functions. Many of the membranes of the eukaryotic cell are part ofan endomembrane system. Some of these mem- branes are physically connected and others are linked when tiny vesicles (sacs made of membrane) transfer membrane segments between them. “The endomembrane system includes the nuclear enve- lope, endoplasmic reticulum, Golgi apparatus, lysosomes, various types of vesicles and vacuoles, and the plasma membrane. (The plasma membrane is not exactly an endo (Gnner) membrane in physical location, but itis related to the other membranes by the transfer of vesicles.) Many of these organelles interact in the synthesis, distribution, storage, and export of molecules “The largest component of the endomembrane system is the endoplasmic reticulum (ER), an extensive network offlattened sacs and tubules. (The word endoplasmic means “within the cytoplasm,” and reticulum is Latin fr “little net”) ‘The ERisa prime example of the direct and indirect interre Jatedness of parts ofthe endomembrane system. As shown in Figure 4.5 on the facing page, membranes ofthe ERare con tinuous with the nuclear envelope. And when vesicles bud from the ER, they travel to many other components of the endomembrane system, ‘The membranes ofthe ER enclose a space separate fom the cytosol. Indeed, an important aspect ofthe components of the endomembranesystem is dividing the cell into functional com- partments, each of which may requir different conditions Which structure Includes all thers inthe lst ER, vesicle, mndomembrane system, nuclear envelope? ‘The Endomembrane System 6.4.8 The endoplasmic reticulum is a biosynthetic workshop One ofthe major manufacturing sites in a cells the endoplas- mic reticulum, The diagram in Figure 4.8A shows a cutaway view ofthe interconnecting membranes of the smooth and rough ER, which can be distinguished in the superimposed electron micrograph. Smooth endoplasmic reticulum is called smooth because its outer surface lacks attached ribo: somes. Rough endoplasmic reticulum has bound ribo- somes that stud the outer surface ofthe membrane; thus, appears rough in the electron micrograph, ‘Smooth ER The smooth ER of various cell types functions Ina variety of metabolic processes. Enzymes of the smooth ER are important in the synthess of lipids, including olls, phos- phtolipids, and steroids. Invertebrates, for example, cells ofthe ‘ovaries and testes synthesize the steroid sex hormones. These cellsazerich in smooth ER, a structural feature that fits their function by providing ample machinery for steroid synthesis. (Our live cells also have large amounts of smooth ER, ‘with enzymes that help process drugs, alcohol, and other potentially harmful substances. The sedative phenobarbital and other barbiturates are examples of drugs detoxified by these enzymes. As liver cells are exposed to such chemicals, ‘the amount of smooth ER and its detoxifying enzymes Increases, thereby increasing the rate of detoxification and thus the body's tolerance to the drugs. The result sa need for higher doses ofa drug toachieve a particular effec, such, as sedation. Also, because detoxifying enzymes often cannot distinguish among related chemicals, the growth of smooth ER in response to one drug can increase the need for higher doses of other drugs. Barbiturate abuse, for example, can decrease the effectiveness of certain antibiotics and other useful drugs. ‘Smooth ER has yet another function, the storage of alcium ions. In muscle cells, for example, a specialized smooth ER membrane pumps calcium ions into the interior of the ER. When a nerve signal stimulates a muscle cell, calet= lum ions rush from the smooth ER into the cytosol and tigger contraction of the cel. Rough ER Many types of cells secrete proteins produced by bosomes attached to rough ER. An example of secretory protein is insulin, a hormone produced and secreted by eet= tain cells of the pancreas and transported in the bloodstream. ‘Type I diabetes results when these cells are destroyed and a lack of insulin disrupts glucose metabolism in the body. Figure 4.88 follows the synthesis, modification, and packaging ofa secretory protein. As the polypeptide is syn- thesized by a bound ribosome following the instructions of| an mRNA, @ its threaded into the cavity ofthe rough ER. Asit enters, the new protein folds into its three-dimensional shape. @ Short chains of sugars are often linked to the poly- peptide, making the molecule a glycoprotein (glyco means sugar”). @ When the molecule is ready for export from the ER itis packaged ina transport vesicle, a vesicle that moves from one part of the cell to another. @ This vesicle buds off from the ER membrane, 64 curren | A Tour of the Call “A Figure 4.88 symhesis and packeging of 2 secretory protein bythe rough ef ar) ‘The vesicle now carries the protein to the Golgi apparatus for further processing. From there, a transport vesicle containing the finished molecule makes ts way to the plasma membrane nd releases its contents from the cell.dition to making secretory proteins, rough ER isa mem brane-making machine for the cell It grows in place by adding _membrane proteins and phospholipids to its own membrane, Aspolypeptides destined to be membrane proteins ow from bound ribosomes, they are inserted into the ER membrane, Phospiolipids are made by enzymes of the rough ER and also inserted into the membrane, Thus, the ER membrane grows, 1d portions oft are transferred to other components ofthe tendomembranesystem in the form of transport vesicles, Now lets follow a transport vesicle carrying products of ‘he rough ER to the Golgi apparatus Explain why we say thatthe endopl isa biosynthetic workshop. The Golgi apparatus modifies, sorts, and ships cell products After leaving the ER, many transport vesicles travel tothe Golgi apparatus. Using alight microscope and a staining -hnique he developed, Italian scientist Camillo Golgi dis- covered this membranous organelle in 1898, The electron ‘microscope confirmed his discover revealing a stack of flattened sacs, looking much like a pile of pita bread, a cell may contain many, even hundreds, of these stacks, The number of Golgi stacks correlates with how active he ce isin secreting proteins—a multistep process that, as you have ust seen, Is initiated in the rough ER. The Golgt apparatus serves as a molecular warehouse and, processing station for products manufactured by the ER. You ‘can follow these activities in Figure 4.9, Note that, unlike DR sacs, the flattened Golgi sacs are not connected. @ One side of a Golgi stack serves asa receiving dock for transport vesicles produced by the ER. @ A vesicle fuses with a Golg! sac, adding ts membrane and contents to the “receiving” side, © Products of the ER are modified as they progress through che tack. © The “shipping” side of the Golgi functions as more than SO years late, bud offand depot, dispatching its products in vesicles th travel to other sites, ow might ER products be processed during their transit through the Golgi? Various Golgi enzymes modify the carbo- hydrate portions of the glycoproteins madein the ER, removing, some sugars and substituting others. Molecular identification tags such as phosphate groups, may be added that help the Golgi sort molecules into different batches fr different destinations, Finished secretory products, packaged in transport ves: cles, move to the plasma membrane for export from the cell, Alternatively, finished products may become part of the plasma membrane itself or part of another organelle, such asa lysosome, which we discuss next What Is the relationship ofthe Goig apparatus to the ER in a protelnsecroting coll? Sa Ey ent Songs crakear PCO ees ‘A Figure 4.9 The Gol apparat recevng, processing, ané shiping product The Endomembrane System 654.10 Lysosomes are digestive compartments within a cell "Alysosomeis a membrane-enclosed sac of digestive enzymes, “The name ysosomeis derived from two Greek words meaning “breakdown body.” The enzymes anid membranes of lysosomes are made by rough ERand processed in the Golgi apparatus, Illustrating a key characteristic of eukaryotic cells—compart- ‘mentalization—a lysosome provides an acidic environment for itsenzymes, while safely isolating them from the rst ofthe cell. Lysosomes have several types of digestive functions, Many protists engulf food particles into membranous sacs called food vacuoles. As Figure 4.408 shows, lysosomes fuse with food vacuoles and digest the food. The nutrients are then released into the cytosol. Our white blood cells engulf bacte- sa and then destroy them using lysosomes, Lysosomes also serve as recycling centers. Cells enclose damaged organelles fr small amounts of cytosol in vesicles. lysosome fuses with such a vesicle (Figure 4.108) and dismantles its contents, ‘making organic molecules available for reuse. With the help of lysosomes, a cll continually renews itself. ‘The cells of people with inherited lysosomal storage dis eases lack one or more lysosomal enzymes. The lysosomes ‘become engorged with undigested material, eventually Inteefering with cellular function. In Tay-Sachs disease, for example lipld-digesting enzyme is missing, and brain cells ‘become impaired by an accumulation of lipids, Fortunately, lysosomal storage diseases are rae in the general population, as they ae often fatal in early childhood. owt a ctr a ying etre soon 0 in be ‘A Figure 4.208 tysosome fusing with food vacuole and digesting too, ater wnien nutiente at released tothe eytoeo! “A Figure 4.408 tysosome fusing with a vsile containing # damaged ‘organelle end then cgesting and reyeng its contents 4.11 Vacuoles function in the general maintenance of the cell Vacuoles are large vesicles that havea variety of functions, In Figure 410A, you saw how a food vacuole forms asa cell ingests food. Figure 4.2.18 shows two contractile vacuoles in the protist Paramecium, looking somewhat lke whee! hubs ‘with radiating spokes, The *spokes” collect water from the cell,and the hub expelsit to the outside. Water constantly enters freshwater protist from their environment, Without way to get re of the excess water, ‘the cell would swell and burst In plants and fungi, ce. tain vacuoles have a diges tive function similar to that of lysosomes inanimal cells In the seeds of plants, small vacuolesin storage cellscan hold reserves of proteins, ‘Vacuolesin flower petals contain pig ‘ments that attract Central vacuole. ‘chloroplast. Nucleus > Figure 4.248 conval vacuole Ina plant ct > protect the plant against herbivores by storing compounds | that are poisonous or unpalatable to animals. Examples Include nicotine, caffeine, and various chemicals we use as pharmaceutical drugs. Figure 4.448 shows plant cel’ large central vacuole, ‘which helps the cell grow in size by absorbing water and enlarging. Italso stockpiles vital chemicals and may act asa trash can, safely storing toxic waste products. GB ot vctle part ot ne endememran ater Ex, ‘A Figure 4.1214 Contractile vacuoles In Paramecium, a unesiuar eukaryote pollinating insects. -uoyts suequownapue ayo ‘Vacuoles may also help ied yu susguou uso au ot Sut 19 9 2 66 curren «| A Tour of the Cell4,12 A review of the structures involved in manufacturing and breakdown Figure 4.42 summarizes the relationships with- In the endomembrane system. You can see the direct structural connections between the nucle. arenyelope, rough ER, and smooth ER. The zed arrows show the functional connections, as ‘membranes and proteins produced by the ER travel in transport vesicleso the Golgi and on coother destinations, Some vesicles develop Into lysosomes or vacuoles, Others travel to and fuse with the plasma membrane, secreting thelr contents and adding their membrane to the plasma membrane Peroxisomes (sce Figures 444 and 4.48) are metabolic compartments that do not org {nate from the endomembrane system. Infact, how they ae related to other organelles is still ‘unknown, Some peroxisomes break down fatty acids to be used as celular fuel. In your liver, peroxisomes detoxify harmful compounds. In these processes, enzymes transfer hydrogen 4. Figure 4.12 Review of the endomemarane sytem from the compounds to oxygen, producing hydrogen perox: “canciones one et ‘de Hg0,). Other enzymes in the peroxisome convert this ic by-product to water—another example of the impor. tance ofa cell’s compartmental structure How do transport vesicles help te together the endomembrane A cell requires a continuous supply of energy to perform. system? the workof life, Next we consider two organelles that act as cellular power stations—mitochondria and chloroplast, ‘ssopee Ja ssvesan pub seveusu tou SepseedSue 8 eae eto) 4.13 Mitochondria harvest chemical energy from food Mitochondria (singular, mitochondrion) are organelles that carry out celular respiration in nearly all eukaryotic cll. astrating the theme of @XETENENEGED. mitochondria use 0, and release CO, in transforming the chemical energy of foods to a form (ATP) that can be used for cellular work A mitochondrion is enclosed by two membranes, each a phospholipid bilayer with a unique collection of embedded peter membrane tha unique coleti : Pte proteins (Figure 4.13). The mitochondrion has two internal 9, fa . Peed compartments, The fis isthe intermembrane space, the — narrow region between the inner and outer membranes. The ee inner membrane encloses the second compartment, the mitochondrial matrix, which contains mitochondrial DNA and ribosomes a well as enzymes that catalyze some of the reactions of cellular respiration. The inner membrane " ) Pee ey Js highly folded and contains many embedded protein mole cules that function in ATP synthesis The fold, called cristae, Increase the membrante’s surface area, enhancing the mito: chondrion’s ability to produce ATP. at clr espatont 01 somou poo o ious eoiews ow sisies yn sSoD0u1¥@ A Figure 4.13 The mitochondrion, eof cellular eeprtion Energy-Converting rganoies 674.14 Chloroplasts convert solar energy to chemical energy Most of the living world runs on the energy provided by pho- tosynthess, the conversion of light energy from the un to the chemical energy of sugar molecules. Chloroplasts are the photosynthesizing organelles of plants and algae. This organelle carries out complex, multistep processes, soit snot surprising that internal membranes partition the chloroplast into compartments (Figure 4.24) Itisenclosed by an inner and outer membrane separated by a thin inter ‘membrane space. The compartment inside the inner mem. brane holds a thick fluid called stroma, which contains chloroplast DNA and r!bosomes a well as many enzymes. ‘A network of interconnected sacs called thylakoids is sus- pended in the stzoma, The sucs are often stacked like poker chips; each stack is called a granuma (plural, grana). The com= partment inside the thylakoids is called the thylakoid space. The thylakoids are the chloroplasts solar power packs— the sites where the green chlorophyll molecules embedded in thylakoid membranes trap solar energy: In the next module, ‘we explore the origin of mitochondria and chloroplasts. Which membrane in chioroplast appears to be the most ‘extensive? Why might this be 0? ‘A. Figure 4.24 The chloroplast, ste of photosynthesis 4.15 Mitochondria and chloroplasts evolved by endosymbiosis Mitochondria and chorplsscontaina single ‘ieaar DNA noes snain str tow prolrotcchiomosone and ibsomte moe Smiarto prokaryotic ribosomes than ocukayotcone Intsestngy both ones eroduceinacelby spaces tembling that of ean poaryotes “te endosyabiont theory as that miochondsaand htop memes sal prays ht began ng win agercels These joaryotstay hav ele thelr ee esondgese py or press re #2) Vices ypothests how tapos could eben bens wr tha wasbecoming increasing aoc fom the oxygen genetingpo‘orntess of protarote ‘host would hve benefited fom anendomblont tat was thio ure cnygen oles ge amounts fener fom brn mle over thecouse of tuto, te host Cclancisenfosiontmegedinton singe anism —a alata nitechonn, not iesccels sxaueds photoshetepakaryot the poaryotecould provide hse with oust tines pendent ho and rosy net many Renee conldbecome clay cl containing corps All eukaryotes have mitochondtl, but nt ll have chioro- lasts What the evolutionary explanation? Suyerco so a ae ota apyquopan puonar Yeeeah suyyeicn so satan ota dn ase pon soaps 2 © 68 curren | A Tour of the Call ence scm eer { ceasing emma, soars ncostret SSF Sats pesto . vtcrenren erasing eee aan LZ Nenphotseytetc ree tet eomeit Photosynthetic eukaryote ‘A Figure 4.45 Endosymbioti orgn of mitochondria and chloroplastsThe Cytoskeleton and Cell Surfaces 4.16 The cell’s internal skeleton helps organize its structure and activities As you saw in the chapter introduction, networks of protein fibers extend throughout a cel. Collectively called the cytoskeleton, these fibers act like a skeleton In providing for structural support as well as movement. Both the internal ‘movement of cell parts and the swimming of crawling motil- ity of some cells usually involve the interaction ofthe cyto- skeleton with motor proteins, Three main kinds of fibers make up the eytoskeleton: microtubules, the thickest fiber; microfilaments, the thin- nest; and termediate filaments, in between in thickness. Figure 4.46 shows three micrographs of cells of the same type, each stained with a different fluorescent dye that selectively highlights one ofthese types of fibers Microtubules ace straight, hollow tubes composed of globular proteins called tubulins. As indicated in the bottom left of Figure 4.16, microtubules elongate by the addition of tubulin proteins, which consist of two subunits. Microtubules are readily disassembled, and their tubulin ‘ean be reused elsewhere in the cell In animal cells, micro- subules grow out from a region called the centrosome, which contains a pair of centrioles, each composed of ring of microtubules (see Figure 4.4A). Plant cells lack centrosomes with centrioles and organize microtubules by other means. Microtubules shape and support the cell and also act as {racks along which organelles equipped with motor proteins move, For example, a lysosome might use Its motor protein “feet” to “walk” along a microtubule to reach a food vacuole. Microtubules aso guide the movement of chromosomes When cells divide, and they are the main components of cia ‘ LAND | Fibrous proteins called together Tubulin rosein Microtubule 4A Figure 4.26 Thee types of ters of he eytskeleton: meotubuls la and flagella, We will return to the structure of these locomo- tive appendages in Module 4.18. Intermediate filaments are found in the cells of most animals, They are made of various Fibrous proteins that supet- coll nto cables. Intermediate flaments reinforce cell shape and anchor some organelles, For example, the nucleus typi cally sts ina cage made of intermediate laments, Whereas ‘microtubules may be disassembled and reassembled elsewhere, intermediate filaments are often more permanent fixtures in the cell, The outer layer of your skin consists of dead skin cells packed fll ofintermediate laments Microfilaments, aso called actin filaments, are solid rods ‘composed mainly of globular proteins called actin, arranged Ina twisted double chain (bottom right of Figure 4.16 Microflaments form a three-dimensional network just inside he plasma membrane that helps support the cell's shape. This !s especially important for animal cells, which lack cell walls Microfllaments are also involved in cell movements. Actin filaments and thicker filaments made ofa type of motor called myosin interact to cause contraction of muscle cells (see Figure 30.98). Localized contractions brought about by actin and myosin are involved in the amoeboid (crawling) movement ‘ofthe protist Amoeba and some of your white biood cells In the next module, we survey some of the techniques that Jed to the discovery of the cytoskeleton, Which component ofthe cytoskeleton is most important in (a) holding the nucleus in place within an animal cl (b) guiding wansport vesicles from the Golgi to the plasma ‘membrane; (e contracting muscle co's? Actin protein Microfilament Intermediate filament 2 with green fucrescont ‘molecules (et), ntermetlate flaments labeled yelowsreen (conte), and mlrfiaments labeled red (rah The Oytoskeleton and Cal Surtacos 694.17 Scientists discovered the cytoskeleton using the tools of biochemistry and microscopy ‘Asyou learned in Module 4.1, improvernents in microscopes and staining techniques ed tothe discovery of organelles. But biologists originally thought that these structures floated freely in the cell, Let’ trace the progressive sequence of new techniques that led to the discovery of microfilaments, ‘the component ofthe cytoskeleton built rom actin, Inthe 1940s, iochemists fst isolated and identified th proteins actin and myosin from muscle cells. In 1954, scientists, sing netly developed techniques of microscopy, established hhow filaments of actin and myosin interac in muscle contrac- tion, In the next decade, researchers developed a technique to stain and identify atin filaments with the electron microscope. Imagine their surprise when they found actin not just in the muscle cells they were studying butalso in other cellspresentin their samples. Purther study identified actin ‘Mlamentsin all types of cel ‘Today we ake for granted our abil- lty to “See” the cytoskeleton (as you saw in the chapter introduction). But intact networks of microfilaments were not visualized in cells until 197 Scientists developed antibody proteins that would bind toactin and attached. fluorescent molecules to them. (When fluorescent molecules absorb light, they “glow” because they emit light of a specific wavelength or color.) These ‘uorescent antibodies revealed, How has our knowledge of cells grown? 4A Figure 4.27 A tuorescence mic {raph of he cyoskeeton mierotubules are een, microfameont re resis oangs) ‘remarkable and beautiful web of microfilaments. Figure 4.27 shows how different fluorescent tags can attach to various components of the cytoskeleton, Researchers then tagged actin proteins themselves ‘with fluorescent molecules and injected them into living cells This technique enabled scientists to visu alize the dynamic behavior of cytoskeletal proteins in living cells. By paring video cameras with microscopes, scien: Uists suddenly could “watch” what was happening in cells ovet {me and follow the changing architecture ofthe cytoskeleton, As scientists develop new techniques, our understanding of the cytoskeleton will continue to grow. Current research {neludes a molecular approach in which ne genes for cytoskeleton proteins are sequenced and compared across diverse organisms. For example, the genes for actin are found to be highly conserved actoss evolutionary time—the actin proteins that facilitate the creeping ‘movement of amoebas are remarkably similar to the actin proteins involved in the “amoeboid” movement of your white blood eels By How does te discovery ofthe cytoskeleton ilustrate the idea that advances in scenttc knowedge often rely on advances in techniques. fan tools? sn soup au poy sieda Kb 4.18 Cilia and flagella move when microtubules bend The role ofthe cytoskeleton in movement is clearly seen in the motile appendages that protrude from certain cells. The short, numerous appendages that propel Panzmecium (see Figure 4.18) are called eilia (singular, cli). Other protists ‘may move using flagella, which are longer than cilia and ‘usualy limited to one ora tie few per cell Some cells of multicellular organisms also have cilia or flagella. For example, Figure 4.484 shows cilia on cells lining the trache (ovindpipe). These cilia sweep ‘mucus containing trapped debris out of your lungs. (This cleaning function is impaired by cigarette smoke, which paralyzes the cilia) Most ani flagellated sperm. A flagellum, shown in Figure 4.488, propels the cell by an undulating whiplike motion. In con: trast, cilia work more like the coordi- nated oars of a rowing team. ‘Though different in length and beating pattern, cilia and flagella have a common structure and mechanism of movement (Figure 4.486, on the facing page) Both are composed of microtubules wrapped in an extension of the plasma membrane. in nearly all eukaryotic cilia and flagella, a ring of nine microtubule doublets sur- rounds a central par of microtubules, This arrangement is called the "9 ~ 2" pattem, The microtubule assembly Flagellum ‘A Figure 4.184 ta on cots ting the respiratory tact 4 Figure 4.188 Undulading ‘mals and some plants have Isanchored in the cell by abasal_—__flagelum on a human sperm cl 70 curren | A Tour of the Callouter microtubule ‘doubiet contrat ‘microtubules Cross-inking proteins Motor proteins (oyneins) Plasma membrane. ‘A Figure 4.28¢ Ieoral structure of 3 eukaryote fags or ela body (not shown in the figure}, which is structurally very similar toa centriole Infact, in humans and many other animals, the basal body of the fertilizing sperm’s flagellum enters the egg and becomes a centeole, How does the microtubule assembly shown in Figure 4.18C produce the movement of cilia and fagella? Large motor proteins called dyneins (red in the figure) are attached along. tach outer microtubule doublet. A dynein protein has two “feet” that “walk” along an adjacent doublet. The walking movement is coordinated so that ithappens on ‘one sideata time. The microtubules are held together by flexible cross-linking proveins {(putplein the diagram). Irthe doublets were not held in place, they would slide past each, ther. Instead, the “walking” of the dynein feet, ‘causes the microtubules—and consequently he cilium or flagellum—to bend, ‘Acilium may also serve asa signal-receiving “antenna” forthe cell. Cilia with this function, are generally nonmotile (they lack the central pair of microtubules), and there is only one per cell. In fact, in vertebrate animals, it appears that almost all cells have what is called a primary cium, Although the primary cilium was discovered more than a century ago, its importance to embryonic evelopment, sensory reception, and cell function ‘is only now being recognized. Defective primary cilia have been linked to polycystic kidney disease and other human disorders, Primary cllary dyskinesia (PCD), also known as Immotle cilia syndrome, is a fary rare disease in which ci and flagella are lacking motor proteins. PCD is characterized by recurrent respiratory tract infections and immotile. sperm. How would you explain these seemingly unrelated symptoms? 19 The extrac: ‘The plasma membrane is usually regarded as the boundary of, the cell, but most cells synthesize and secrete materials that are external to the plasma membrane, Animal cells produce an extracellular matrix (ECM) (Figure 4.18). This elabo- rate layer helps hold cells together in tissues and protects and supports the plasma membrane, The main components of the ECM are glycoproteins, proteins bonded with carbohytrates. The most abundant gly coprotein is collagen, which forms strong fibers outside the cell. In fact, collagen accounts for about 40% of the protein ‘in your body. The collagen fibers are embedded in a network woven from large complexes that include hundreds of small glycoproteins connected to along polysaccharide molecule (own as gzeen in the figure). The ECM may attach to the cell through other glycoproteins that then bind to membrane proteins called imtegrins, Integrins span the membrane, attaching on the other side to proteins connected to microfi ments of the cytoskeleton. 4s their name implies, integrins have the function of. Integration: They transmit signals between the ECM and the cytoskeleton and can communicate changes occurring outside and inside the cell, Current research is revealing new and influential functions ofthe ECM. For example, itcan regulate a cel’s behavior by directing the path along which, embryonic cells move, Researchers have also learned that a jular matrix of animal cells functions in support and regulation polysaccnaride Collagen fiber Connecting stycopretein of cytoskeleton ‘A Figure 4.29 the exracolular mavtx (ECM) of an animal cl cell's ECM can even influence the activity of genes through the signals it relays, Roforing to Figure 4.19, describe the structures that provide support te the plasma membrane. _soqy usByoo pur susnvoatg Bupsowoo oy pus uonawcov eM -unuieqanl rua sumiqus inary pages a subi 941 The Cytoskeleton and call Suraces 7A.4.20 Three types of cell junctions are found in animal tissues Neighboring cellsin animal tissues often adhere, interact, and communicate through specialized junctions between thera, Figure 4.20 uses cell ining the digestive tact to illustrate three types of cll junctions. (The projections atthe top ofthe cell increase the surface area for absorption of nutrients.) At tight junctions, the plasma membranes of neighboring cellsare knit tightly together by proteins. Tight junctions pre- ‘vent leakage of fuld actossa layer of cells. The dotted green arrows show how tight junctions prevent the contents, of the digestive tract from leaking into surrounding tissues. Anchoring junctions function lke rivets, fastening cells together into strong sheets, Intermediate filaments made ofsturdy proteins aichor these junctions inthe eytoplasm. "Anchoring junctions are common in tissues subject to stretching or mechanical stress, such as skin and muscle. Gap junctions, also called communicating junctions, are channels tha allow small molecules to flow through protein-lined pores between cells, The flow of fons through sap junctions in the cells of heart muscle coordinates theit contraction. Gap junctions are common in embryos, where communication between cells s essential for development. A muscle tear injury would probably involve the rupture ‘of whieh typeof ell junction? conaunr moe @ > Tight junctions provent fi fom ‘moving across @ layer ot ces ‘Tight junction ‘Anchoring Junetion | Gap junction F— plasma memoranes of adjacent cols Jone or small molecules ‘A Figure 4.20 Thee types of elunctions in animal tissues Extracellular matrix 4.21. Cell walls enclose and support plant cell ‘The cell walls one ofthe features that distinguishes plant cells from animal cells Ths rigid extracellular structure not only protects the cells but also provides the skeletal support that keeps plants upright on land, Plant cell walls consist of fibers of cellulose (see Figure 3.7) embedded in.a matrix of other polysaccharides and proteins. This fbers-in-i-matrix construction resembles that of stel-einforced concrete, which is also noted for its strength. Figure 4.24 shows the layered structure of plant cell ‘alls. Cells initially lay down a relatively thin and flexible primary wall, which allows the growing cell to continue to enlarge. Between adjacent cells isa layer of sticky polysac- charides called pectins (shown here in dark brown), which, ‘ue the cells together. (Pectin Is used to thicken jams and jellies) When a cell stops growing, it strengthens its wall, Some cells add a secondary wall deposited in laminated layers next to the plasma membrane. Wood consists mainly of secondary wall, which are strengthened with ecules called lignin, Despite their thickness, plant cell walls do not totally Isolate the cells from each other. Figure 4.21 shows the ‘numerous channels that connect adjacent plant cells, called plasmodesmata (singular, plasmodesma). Cytosol pass- Ing through the plasmodesmata allows water and other small molecules to freely move from cell to cell. Through idmol- 2 ‘A Figure 4.21 Plant call walls and plasmodesmata plasmodesmata, the cells ofa plant tissue share water, nour- Ishment, and chemical messages, GQ) vver sna ct unton snout pasmatese? cone4.22 Review: Eukaryotic cell structures can be grouped on the basis of four main functions Congratulations: You have completed the grand tour ofthe cell. tn the process, you have been introduced tomany important cel structures, To provide s frame- ‘work for this information and reinforce the theme {ha structure's correlated with function, we have srouped the eukaryote cell structures into four cate- goriesby general function, as reviewed in Table 4.22. The fist category i genetic control Here we include themucleus that houses a cells genetic instructions and the ribosomes that produce the proteins coded for inthos instructions. The second category includes, organelles ofthe endomembrane system that are snvolved in the manufacture, dstbution, and break: doven of materials The third category includes the {wo energy-procssing organelles, mitochondria and clloroplasts, And the fourth category—steuctural sup port, movement, and intercellular communication — Inches the cytoskeleton, extracellular structures, and connections between cells. ‘Within most ofthese categories, a structural similarity underlies the general function of each component, Manufacturing depends heavily on 8 network of structually and functionally onnected ‘membranes. All the organelles involved in the breakdown of recycling of materials are membra ‘nous sacs, inside of which enzymatic digestion can safely occur, n the energy-procesing categor expanses of metabolically active membranes and intermembrane compartments within the orga elles enable chloroplasts and mitochondria to per- form the complex energy conversions that power the cell Even in the diverse fourth category, there is 2 common structural theme in the various protein fibers of most ofthese cellular systems. ‘We can summarize further by noting thatthe over- all structure of cells closely related tots specific function. Thus, els that produce proteins fr export contain alarge quantity of ibosomes and rough ER, ‘while muscle cells are packed with microfilaments, ‘myosin motor proteins, and mitechondtia, And, finaly, let us emphasize that these cellular structures form an integrated team-—with the property of ie emerging tthe level ofthe cell from the coordinated functions ofthe team members. A cell beautifully iustates our theme of QUEENS iti ving nit that fgreater than the sum of spars ow do mitochondria, smooth ER, and the cytoskeleton all contribute to the contraction of a muscle cell? “untes Jo oseoys pus arid ou fq uole-tue> evar say si toute au gay 2 wo} 9 Glows ede Nae 11. Genetic Contr! Nucleus DNA opcation, RNA sythess: assembly of ebosemal suri hms) Ribosomes Potpopti eatin syrthosis 2. Manufacturing, Distribution, Rough ER Breakdown ‘Syrtnsis of memtranelipds and proteins, secretory, proteins, and hytevte enzymes; formation of transport vesctes| Lipa syrmesi; detoteton in ver el; elem lon storage In muse cos Mositeation ane sortng of ER prosets formation of Iysosomes and transport wesles Goll apparatus lysosomes tn animal cots and some pts) voouoies Digestion of ingested foo or bacteria art eyeing of cls damaged ones end mseromolecles Digestion ood vacuole: water balance corvactle vacuole storage of hemicals and col enlargement (corral vacuole in plant co's) Diverse metabo processes, with breskdown of tne jogen peroxce byproduct Peroisomes (ot part of fendormembrene sytem) 3. Energy Processing itocnonaia Celular respiration: conversion of chemical energy in ‘ood to nema! energy of ATP Chloropast i pnts Protosyness:conwersion of gt energy to chemical and age} ener of sugars 4, Structural Support, Movement, and Communication Between Cells oytosteleton Maintenance of ci ape; anchorage fr (motiaments, ‘organelles; movement of erganales within cos lnvamecate cl movement (eawing, muscle contacion, aments, ae bending af ela an ago) reroute) Pasma membrane ‘vaca mati (animals) Regulate trafic in ac out of ca ‘Support regulation of clr acties Communication between cols: ising of cols intssues ‘Support ana protectin; bing of cls In tssues Col juetons cal wala (implores) The Ovtosketeton and Cal Surtees 73For practice quizzes, BioFlx animations, MPS tutorials, video tutors, and more study tools designed for this textbook, go to MasteringBiology” 4 REVIEW REVIEWING THE CONCEPTS Introduction to the Cell (4.1-4.4) ‘4. Microscopes reveal the world ofthe eel. The ight microscope ‘can display living cells. The greater magnification and resolution bf the scanning and transmission electton microscopes reveal the talrastrutuce ofc membrane. Tie microscope sizeof most cells provides large surface to-volume aio. The plasma membrane isa phospholipid blayer vith embeded proteins 43 Prokaryotc coll are structurally simplor ‘han eukaryotic cols Al cellshave plasma mem: brane, DNA, bosom, and eyoso.Prokaryoti cells lackeganeles, ‘44 Eukaryotic calls are partitioned into funetonal ‘compartments. Venbrane enclosed organelles compartmentalizeacel's activities, ‘The Nucleus and Ribosomes (4.5~4.6) ‘45 The nucleus contains the cals genetic Instructions. The ‘hucleus houses the cell's DNA, which directs protein synthesis ia messenger RNA, Subunits of ribosomes are assembled in the pceolis, ‘4 Ribosomes make proteins for use in the cell and for export ‘Composed of ribosomal RNA and proteins, bosomes synthesize proteins according to diectons from DNA ‘The Endomembrane System (4.7-4.12) 4.7 Many organelos are connected In the endomembrane system. 4.8 The endoplasmic reticulum fs a biosynthetic workshop. “The ERs membranous network of tubes and sacs, Smooth ER symtesizes lipids an processes toxins. Rough ER produces, ‘membranes and ribosomes on ts surface make membrane and secretory protelns. ‘4.9 The Golgi apparatus modifies, sorts, and ships cell products. “The Golgi apparatus consists of stack ofsacsin which products ofthe ER are processed and then sent to other argancies orto thecellsurtace 4.20 Lysosomes are digestive compartments within a cof Tysonomes house enzymes that breakdown ingested substances and damaged organeles. ‘4.11 Vacuoles function inthe general maintenance of the eel, Some protists have contract vacuoles, Plant ells contain a large ‘ental vacuole that stores molecules and wastes and faciltates growth. 42 A review of the structures Involved in manufacturing and breakdown. The organelle ofthe endomembrane system are interconnected structurally and functionally Energy-Converting Organelles (4.13-4.15) ‘4.3 Mitochondiia harvest chemical energy from food. 74 cuorers | A Tour of the Cel 4.14 Chloroplasts convert solar energy to chemical energy. 4.18 IMitochondiia and chloroplasts evolved by endosymbiosis. “These organelles onginated trom prokaryotic cell shat became resident a host cl ‘The Cytoskeleton and Cell Surfaces (4.16-4.22) ‘4.6 The co's intemal skeleton helps organize Its structure and settles, The cytoskeleton includes microflaments, intermediate flaments, and microtubules, Tele functions include malntenance of cell shape anchorage and movement of organelles, amoeboid ‘movement, and muscle contraction. ‘4.47 Sclontists discovered tho cytoskeleton using the tools ‘of blochemstry and mleroscopy. ‘418 Cilla and fagetia move when microtubules bond. Eukaryotic ‘iia an flagellaare locomotor appendages made of microtubules 109-2" arrangement {4.29 The extraceluiar matrix of animal eels funetions in support and regulation. The FCM consists mainly of glycoproteins, wie bind ussue cells together, suppor the plasma membrane, 2nd ‘communicate with the cytoskeleton, {4.20 Thvee types of col unetions are found in animal tissues. tight junetons ind els to form leakproo! sheets. Anchoring junctions rivet cell into strong tissues Gap junctions allow ionsand small ‘molecules t flow rom cell 10 cl {4.24 Cell walls enclose and support plant eels Plat cel wallsare made largely of elulse. Plasmadesmata ae connecting chanies between cells 4.22 Review: Eukaryotic eal structures can be grouped onthe basis of four main funetions, These functions ae") genetic con oo 2) manulaeturing,distabution, and breakdown (3) energy processing; ad) stress support; movement, and comm {ation between cell CONNECTING THE CONCEPTS 1. Label the structure in this diagram ofan animal el. Review the fanetions ofeach ofthese organelles,TESTING YOUR KNOWLEDGE Level 4: Knowledge/Comprehension 2. The ultrastructure ofa chloroplasts best studied using 1 light microscope. 2. scanning electron microscope. © transmission electron microscope. light microscope and fuorescet dyes. '3, The cello an ant and an elephant are, on average, these small ‘nw an elephant just has more of them. What Is the main ada tage of small cell sz? (Explain your rvsoning) 4 Asmall eel hasa larger plasma membrane surface area than does large cel ‘by, Smal cll can better take upsuficlentnutsents and oxygen to service ther cell volume. ¢Ietakes les energy to make an organism out of small ces 4 Smallcalls equi less oxygen than do large cls 4, Which of the folowing clues would ell you whether acellis prokaryoticoreularyotc? 4 the presence orabsence ofa gi cell all >, whether or not the cells partitioned by internal membranes © thepresence or absence of ribosomes Both band eareimportant cues. ‘5, Which ofthe followings one ofthe major components of the plasma membrane of plant cll? | phospholipids >, cause Sere «collagen fibers 4. pectine 6. What four cellular components are shared by prokaryotic and ceakaroticeels? 7, Describe two diferent waysin which aca funtion in organisms, Level 2: Application/Analysis (Choose from the following cel for questions 6-1: ‘a pancraticell that secretes digestive enzymes ', ovarian cll that produces estogen (a sterois hormone) {© mmsclecellinthe thigh of along-stance runner 4. Whileblood cel that engullsbacteria §, Inwhict cell would you find the mostysosomes? 5, In which cell would you find the most smoot ER? 10, Inwhich cell would you find the most ough ER? 1. Inwhieh cell would you find the most mitochondria? 12, In what ways do the internal membranes ofa cukazyotie cell contribute to the functioning ofthe cell? 1, Isthisstatement treo ase Animal cells have mitochond plant cells have chioroplats” Explain yout answer, and desrbe the functions ofthese organelles, 14, Describe the structure othe plasma membrane ofa animal cell. What would be found diet inside and outside the ‘membrane? 1, Imagine a sparial ell with radi of 10 pm, Whats the cll’ ‘surface afea num Is volume, tn um? (Note: For aspheze of ‘adi, suface aea = 2? and volume 321, Remember that the alue of ris 314) Whatis the ratio of surface area to volume for this cell Now do the same calculations fora second cel thi fone with atadius of 20 um. Compare the sutfact-to-volume ratios ‘ofthe two cells. How itis comparison signifcant tothe fun oningof els? 16, Descibe the pathway of the protein hormone inslin from i gene toitsexport from acellof your pancreas, Level 3: Synthesis/Evaluation 1, ow might the phrase “ingested but not digested” bewsed ina description ofthe endosymbiotic theory? 18, Claarefound on celsin almost evry oxgan of te human bod, and the malfunction of cia involved in several human dsor ers, Duringembryolgical development, for example cia genee ate aleltward low oflu that niates the eesight organization ‘ofthe body organs. Some individuals with primary cary cys resi (sce Module 4.18 checkpoint question exhibita condition (sts vesu) in which internal organs suchas the heart are on the wrong side ofthe body Explain why thi revered arrangement may bea symptom of PCD. 19, QERUEERININD Microrabutesoften produce movement ‘ough theirinteraction wth motor proteins. Buti some cases, ‘microtobules move cell components wien the lengths ofthe miro {ubule changes, Through series of experiment, researchers deter ‘mined that microtubules grow andshorten as tubulin proteins are ‘added or removed fom their ends, Other experiments showed that Imicrotabues make up the spindle apparatus that pals” chromo ‘Somes toward opposite ends(poes) ofa dividing cell. The figures, ‘below describe a clever experiment donein 1987 to determine ‘whether spindle microtubule shortens (depolymerizes)3¢the end holding a chromosome oat the poleend faving eel Experimentes labeled the microtubules of dividing cell rom ‘pig Kidney with a yellow Suocescent dye. As shown on the et Iualfot te diagram belo, hey then marked region halivay along the microtubules by using laser fo eliminate the Mures- ‘cence from that region. hey dd not mak tne other side ofthe Spindle right side ofthe igure), sta \ZE= == | “Thefigure below lustates the results they observed as the cho _mosomes moved toward the opposte poles ofthe call 1 Su ecient a mlyco hn ie Peo cea ee temic tient Imenters determine wheter hiss the mechanism of remasome movement al cll? Souce: GJ. Got ta, Chctntomnes mv polemasdin nape slang Statlonayrlrtubuls that ootaatey dase om hee ds, oral of Ce gy 104918, 198, ‘Answers to all questions can be found in Appendix 4, 7s i Chapter 4 Review