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    and separates major organelles and other subcellular struc: {ures from one another, The piece of equipment that is used forthis tasks the centrifuge, which spins test tubes holding ‘mixtures of disrupted ces ata series of increasing speeds, «8 process called differential centrifugation. At each speed, the resulting force causes a subset of the cll components to «ttle to the bottom of the tube, forming a pellet At lower iced, the pellet consists of larger components, and higher needs result in a pellet with smaller components ell fractionation enables researchers to prepare spe- « cell components in bulk and identify their functions, sk not usually possible with intact cells. For example, one of the cell fractions, biochemical tests showed w presence of enzymes involved in cellular respiration, ic electron microscopy revealed large numbers of organelles called mitochondria. Together, these data helped biologists determine that mitochondria are the ‘es of cellular respiration. Biochemistry and cytology thos complement each other in correlating cell function, with structure. 1 How stains ued for ight microscopy compare with those Used forelecron microscopy? PM 2. WHAT IF? Which ype of mirorcope would you use 0 study (Jens changesin saps of aang white boa cal 0) che eas ofaurace texture ofa? Forages awe ne appends Eukaryotic cells have internal membranes that compartmentalize their functions CCells—the baste structural and functional units of every organism—are of two distinct types: prokaryotic and eukary- otic. Organisms of the domains Bacteria and Archaea consis ‘of prokaryotic cells. Organisms of the domain Eukarya— protiss, fungi, animals, and plants—all consist of eukary~ otic cells (*Protist” isan informal term referring toa diverse group of mostly unicellular eukaryotes.) Comparing Prokaryotic and Eukaryotic Cells Alcells share certain basic features: They are all bounded by a selective bari, called the plasria membrane (or the cell mem- brane). Inside all cells isa semifluid, jllylike substance called ‘eytosol, in which subcellular components are suspended. All cellscontain chromosomes, which carry genes in the form of DNA. Andall cells have ribosomes, tiny complexes that make proteins according to instructions from the genes. ‘A major difference between prokaryotic and eukaryotic cells isthe location of their DNA. In a eukaryotic cell, ‘most of the DNA isin an organelle called the nucleus, which Isbounded by a double membrane (see Figure 6,8) Ina prokaryotic cell, the DNA is concentrated in a region that isnot membrane enclosed, called the nuteleold (Figure 6.5). Figure 65 A prokaryotic eel. Lacking 2 tue nucleus and the ther membraneenclased organelles the eukaryotic eel the prokaryotic el appears much simpler in intemal structure Prokaryotes include archaea; the general cel structure of these two domain quite sim, Fimbriae: attachment structures on the surface of some prokaryotes | (not visible on TEM) Nucleoid region were the ‘ells DNA is located (nat enclosed by a membrane) Ibosomes compere that yee tas asa ambrane arbre Sein the tone ca wal gid structure outside Sepa ema —oN a | bea: utr coating Stnanyprovaryote —— opm Teenage acne |eoeeraire: (© A thin section through the ‘Corynebacterium gel ocemoton diphtheriae (colorized TEM) organelles some prokaryotes \ CHAPTERS ATouroftheCell 97 Eukaryotic means “true nucleus" from the Greek eu true, and karo, kernel, referring tothe nuciews), an prokaryotic means "before nucleus” (from the Greek po, before, reflec ing the eatlier evolution of prokaryotic cell “The interior of either type of cell iscalled the eytoplasm:; In eukaryotic cell, this term refers only'to the region between the nucleus and the plasma membrane. Within the cyto- suspended in eytosol, area vari plasm ofa eukaryotic cel ‘ety of organelles of specialized form and function, These ‘membrane-bounded structures are absent in almost ll pro: karyotic cells, another distinction between prokaryoticand eukaryotic cells. In spite of the absence of organelles, though, the prokaryotic cytoplasm isnot a formless soup. For exam- ple, some prokaryotes contain regions surrounded by proteins (not membranes), within which specific reactions take place. Eukaryotic cells are generally much larger than prokaryotic cells (ee Figure 6.2), Size isa general feature of cell structure ‘that elates to function, The logistics of carrying out cellular ‘metabolism sets limits on cell size. At the lower limit, the smallest cells known are bacteria called mycoplasmas, which hhave diameters between 0.1 and 1.0 ym. These are perhaps the smallest packages with enough DNA to program metabo- lism and enough enzymes and other cellular equipment to ‘carry out the activities necessary fora cello sustain itself and, reproduce. Typical bacteria are 1-5 wm in diameter, about ter times the sizeof mycoplasmas. Eukaryotic cells are typically 10-100 um in diameter. Metabolic requirements also impose theoretical uppe its om the siz that is practical fora single cell.At the boundary of every cell, the plasma membrane functions asa selective barrier that allows passage of enough oxygen, nutrients, and wastes to service the entie cell Figure 6.6). For cach square ‘micrometer of membrane, only a limited amount of a particu lar substance can cross per second, so the ratio of surface area to volumes critical. Asa cell (or any other objec) increases, In size its surface area grows proportionately less than its volume. (Area Is proportional toa linear dimension squared, ‘whereas volume is proportional tothe linear dimension cubed.) Thus, a smaller cell asa greater ratio of surface area to volume: Compare the calculations fr the fst two “cells” in Figure 6.7. The Scientific Skils Exercise gives you a chance to ‘calculate the volumes and surface areas of two actual cells—a ‘mature yeast cll and a cell budding from It. To explore differ ent ways that the surface area of cellsis maximized in various ‘organisms, see Make Connections Figure 33.8. ‘The need for a surface area large enough to accommodate the volume helps explain the microscopic size of most cells. and the narrow, elongated shapes of some cells, such as nerve cells. Larger organisms do not generally have lager cells than smaller organisms—they simply have more cells sce the far right of Figure 67). A sufficiently high ratio of surface area to volume is especially important in cells that exchange a lot ‘of material with their surroundings, such as intestinal cll, Such cells may have many long, thin projections from their 98 uniTTWo The Cell 1 Figure 6.6 The plasma mambrans. The plasma membrane See oe re tot organs cont of double jer je Sposa ou rte ached too ene een tpl paso phospho ard merxane vrei ar ouns inher ofthe membrane white the Potency are contact wth aqueous sluts on ether ae rare dechmsmey be ataches 9 poten Fils nti cute surac ofthe lama membrane fo) Clore TEM ofa laa membrane The ase nena ‘appears as a pair of dark bands separated by a gold band, Outside of el mee a. Insdeotcel — bogart (Gjoplasm) 04H Hydrophilic region Hydroohobic region Hydophile region (6) Structure ofthe plasma membrane VISUAL SKILLS What parts of the membrane diagram in (t) correspond tothe dark bandsin the TEM in (a)? What pas Cavrespand to the gold band? (Review Figure 5.11) © Mastering Biology BioFix® Animation: Membranes Y Figure 67 Geometric relationships between surface a2 and volume, inthis gram, cel are represented as cubes Use array unt ot ength we cn caelate te cals surface are Ginsquare units, oF uns) 3s oc Re volume ibis or Snsort Uris) andra of surface ——— ateato volume. high surface ates-o-volume ratio ‘aciltates the exchange of i materials between a cell i and its environment. y 18 ibenenan Sera ol i) 19, XbdeXrumbeacedh) unt??? 6 2 6 © Mastering Biology Figure Walkthrough Using a Scale Bar to Calculate Volume and Surface Area of a Cell sow Much Now Cytoplasm and Plasma Membrane IreMade by Growing east al The uncelsa yeast ‘Schrom es cerevsie dhades by budding of esa socal Steno ie ect tom fue 68) uring ts growth, te new el sess her Flom whichinceates te volime, and ew plasma meme {hich nesses surface aren nts exerse you wif use {ial barto determine theses fs mature parent yeast cel and SEeibudding rom Yu wl hen calculate he volume an tfc are ofeach cel Yu se your calculations ede rane how muchojoplaim and plasma membrane he nec eds tosymeie to row to fla How the Experiment Was Done Yeast cells were grown under conditions that promoted divsion by budding. The cells were then viewed witha differential interference convas ight microscope and photographed. bata from the Experiment This light micrograph shows a bus! ding yeast cll about tobe released from the mature parent cel Mature parent en fuding a vam egg ram ne anges ron wom rors he mater nckn Sacco eteson, MOC BBY tm ca 65495-34660) Surface called microvilli, which increase surface area without ‘nappreciable increase in volume. The evolutionary relationships between prokaryotic and ‘haryotc cells will be discussed late in this chapter, and Prokaryotic ces will be described in detail elsewhere (see (Chapter 27). Most ofthe discussion of cell structure that ‘ollows in this chapter applies to eukaryotic cells. Panoramic View of the Eukaryotic Cell In tion tothe plasma membrane atts ote surface a ‘skaryotic cell has extensive, elaborately arranged internal ‘embranes that divide the cell into compartments—the "nels mentioned ear. The cell's compartments ro- {dierent oa envionment that suppor specific mets bal functions, xo incompatible processes can occur simula lyin asingl cel. The plasma membrane and organelle 41 Examine the micrograph ofthe yeast calls The scale bar under thepot abel jm. The sale bar works in the same way Sabon on map, vere for example, 1 inch equal 1 mile hese the bar represents one thousandth of miller thing the salebar as bese unt determine te cameter of the rbatre parent cl and the new cel Start by measuring the esl bar andthe dometer of each cll The units you use are = ‘Geran but working nilimeterss convenient Die each “amter bythe length ofthe caleba and then mutpy bythe Scalebersength vous to ge you the diameter in micrometes. 2. Theshopeofayeast cell can be approximated by asphere. jesse volume of each eal sing the formule forthe {lume of sphere: 4p) v= fe ‘Note that (the Greek letter pi isa constant with an approx imate value of 314d stands for diameter, and rstands for radi, ‘which half the diameter. (b) What volume of new cytoplasm Wille new cel have to synthesize 35 it matures? To determine this calculate the difference between the volume ofthe fullsized Cell and the volume of the new ce ‘As the new cel gros ts plasma membrane needs to expand to ‘Contain the increased volume ofthe cell (a Calculate the sur ace area ofeach cell using the formula forthe surface area of a sphere: A = 4x75 (b) How much area of new plasma membrane tulhe new celhave to synthesize ast matures? 4. When the new cell matures, it willbe approximately how many times gresterin volume and how. many times greater in Surface area than its current size? © Instructors: A version of this Scientific Sls Exercise can be signed n Mastering Biology SCIENCE PRACTICES? 5 ‘membranes aso participate directly in the cell's metabolism because many enzymes are built right into the membranes, The basic fabric of most biological membranes isa double layer of phospholipids and other lips. Embedded in this lipid bilayer or attached to its surfaces are diverse proteins (see Figure 6.6). However, each type of membrane has unique composi- tion of lipids and proteins suited to that membrane's specific functions. For example, enzymes embedded in the membranes ‘of the organelles called mitochondria function in cellular respl- ration, Because membranes are so fundamental tothe organi zation ofthe cell, Chapter 7 will discuss them in detail Before continuing with this chapter, examine the eukai oti cells in Figure 6.8, The generalized diagrams of an animal cell and a plant cell introduce the various organelles and show the key differences between animal and plant cells. The _micrographs atthe bottom of the figure give you. glimpse of cells from different types of eukaryotic organisms. CaPTERG ATourofthe Cell 99 rs oie uke Animal Cell (cutaway view of generalized cel) Nuclear envelope: double) ‘membrane enclosing the rucleus; perforated by | pores; continuous with ER ENDOPLASMIC RETICULUM (ER): network ‘of membranous sacs and tubes; active in ‘membrane synthesis and other synthetic / and metabolic processes; has rough (ribosome studded) and smooth regions Flagellum: motility strueture present in some animal cell composed of a custer of microtubules within an extension of the plasma membrane Nudleolus: nonmembranous structure involved in production | ff oF tivosomes; a nucleus has P NUCLEUS JA} \ one oF more nucleo Rough ER Smooth ER erat er ‘of DNA and proteins visible in a dividing cel as individual // | Chromatin: material consis / | atin: mate isting / /\ condensed chromosomes case: where the cell's \ omens Pama neice memibane enclosing the cl CyTosKELETON: Feinforces eels shape; functions in cell movement components are made of protein Includes: ( Microftaments df } Intermediate |_ ff S foments Cag | icrotubules MO . AF |Rivosomes small brown Microtubules YF | cots: complexes that FE \ | make proteins: freein Microw {jtosal or bound to rough ER or nudeat o envelope \ | olai apparatus: organelle actie Yinsynihess, moditestion, sr | and secretion o el products projections that increase the celts | \ surface area hic ea Peroxisome: organelle with Various specialized metabolic functions produces hydrogen “ Peroride 2s a by-product and then converts i 2 water 4 Mitochondtion: organelle where cellular respiration occurs and most ATP 8 generated organelle where macromolecules are hydrolyzed \ sem digestive © mastering Biology BioFlix® Animation: Tour of an Animal Cl Parent cal a Buds Nucleus Nucleolus ‘east calls reproducing by budding (Sbove colored SEM) anda Se cel (ooh, coorzes TEM) Pn umn cel from ining of wers (colored TEM) 100 uniTrwo The Cell Ee een ‘Nuclear envelope Rough Nucteus Nucleolus ndoniasmic reticulum chromatin Central vacuole: prominent organelle Inolder plant cells; functions include storage, [breakdown of waste products, and hydrolysis | of macromolecules; enlargement of the Golgi apparatus | Yacuole i 2 major mechanism of plant growth Miroflomen’) vrosgeieToN Microtubules _/ Mitochondrion '~ oe ‘ Peroxisome — Chloroplast: photosynthetic organelle converts energy of Plasma membrane I Sunlight to chemical energy stored in sugar molecules Cell wall: outer layer that maintains cells shape and protects cell from ‘mechanical damage; made of cellulose, af Plasmodesmata: cytoplasmic © Mastering Biology ee eee fi ‘channels through cell walls. BioFlix? Animatior al of cent cal” Ee osm Pagel Nucleus Nucieolus Vacuole Chioroolas nd Cel from duckweed (Spitodela olgothize, 4 floating plat (cotezed TEM) (abo, colorized SEM right, colorined Te) col wall charteR6 ATouroftheCell 101 1. Briefly describe the structure and function of the nucleus, the mitochondrion, the chloroplast, and the endoplasmic reticulum. 2. DRAW IT Draw a simplified elongated cell that measures 125 1 1 arbitrary units. nerve cell would be roughly this shape. Predict how Its surface-to-volume ratio would com- pare with those in Figure 67. Then calculate the ratio and check your prediction. For suggested answers, see Apandix et 6.3 The eukaryotic cell's genetic instructions are housed in the nucleus and carried out by the ribosomes On the firs stop of our detailed tour ofthe eukaryotic cell, let's look at two cellular components involved in the genetic control ofthe cel: the nucleus, which houses most of the cell’s DNA, and the ribosomes, which use information from the DNA to make proteins. The Nucleus: Information Central ‘The mucleus contains most of the genes in the eukaryotic cell. Some genes are located in mitochondria and chloroplasts) Itis usu ally the most conspicuous organelle (see the purple structure in the fluorescence micrograph), averaging about 5 pm in diameter. The nuclear envelope encloses the nucleus (Figure 6.9), separating its con- tents from the cytoplasm. The nuclear envelope isa double mem brane. The two membranes, each a lipid bilayer with associated proteins, are separated by a space of 20-40 nm. The envelope is perforated. by pore structures that are about 100 nm in diameter. At the lip of each pore, the inner and outer membranes of the nuclear envelope are continuous. An intricate protein structure called a pore complex lines each pore and plays an important role in the cell by regulating the entry and exit of proteins and RNAs, as well as large complexes of macromolecules. Except at the pores, the nuclear side of the envelope is lined by the nuclear lamina, a netlike array of protein filaments (in animal cells, called intermediate fla- ‘ments) that maintains the shape of the nucleus by mechani- cally supporting the nuclear envelope. There is also much evidence for a nuclear matrix, a framework of protein fibers extending throughout the nuclear interior. The nuclear Jamina and matrix may help organize the genetic material so it functions efficiently 102 unit rwo The Cell Within the nucleus, the DNA is organized into discrete units called chromosomes, structures that carry the genetic information, Each chromosome contains one long DNA, ‘molecule associated with many proteins, including small basic proteins called histones. Some of the proteins help cil, the DNA molecule of each chromosome, reducing its length and allowing it to fit into the nucleus. The complex of DNA and proteins making up chromosomes s called chromatin, ‘When aces not dividing, stained chromatin appearsasa diffuse mass in micrographs, and the chromosomes cannot be distinguished from one another, even though discrete chro 'mosomes are present. Asa cell prepares to divide, however, the ‘chromosomes form loops and coil, condensing and becoming thick enough to be distinguished under a microscope as epe- rate structures (se Figure 16.23). Each eukaryotic species hasa characteristic number of chromosomes, For example, atypical ‘human cell has 46 chromosomes in its nucleus; the excep- tions are human sex cells (eggs and sperm), which have only 23 chromosomes. A fruit ly cell has 8 chromosomes in most cellsand 4 the sex ells A prominent structure within the nondividing nucleusis the nucleolus (plural, nucleoli), which appears through the ‘electron microscope asa mass of densely stained granules ‘and fibers adjoining part of the chromatin. Herea type ‘of RNA called ribosomal RNA (cRNA) is synthesized ‘rom genes in the DNA. Also in the nucleo- |us, proteins imported from the cytoplasm are assembled with rRNA into large and small subunits of ribosomes. These sb units then exit the nucteus through the ‘nuclear pores to the cytoplasm, where large and a small subunit can assemble {ntoa ribosome. Sometimes there are two or more nucleoli the number depends on the species and the stagein the cel’s reproductive cycle. The nucleo ‘may also play a role in controlling cell div sion and the lifespan ofa cel. Sum As we saw in Figure 5.22, the nucleus directs protein synthesisby synthesizing messenger RNA (mRNA) ‘that caries information from the DNA. The mRNAs then transported to the cytoplasm via nuclear pores. Once an ‘mRNA molecule reaches the cytoplasm, ribosomes translate the mRNA’s genetic message into the primary structure of@ specific polypeptide. (This process of transcribing and transl {ng genetic information is described in detail in Chapter 17) © Mastering Biology BioFlix* Animation: Nucleus and Ribosome* Ribosomes: Protein Factories Ribosomes, which are complexes made of ribosomal RNAS and proteins, are the cellular components that carry out protein synthesis (igure 6.10). (Note that ribosomes are not ‘membrane bounded and thus are not considered organell®») Y Figure 6: The nucleus and its envelop. Within the nucleus are the chromosomes, which 2ppe3! as mas of cvomatn(ONA and astocated protein) andone or moe nucle angus, iota nutes), which function in bosome synthesis The nuclear envelope which cna of te Outer membrane membranes separated by narow space, perforated with pores on inedby the aces amin fort manbrane Nuclear envelope: Outer membrane — Inner membrane — —~ > Nucesr pores = N rough ER Pore eee “G mks ae, | segs: Saco e We SMa ea ‘eeaing bon oe Pate Histone Hs > Nuclear lamina (TEM). The neice lamina lines the inner surface ofthe nuclear envelope The ight cela spots are nudes pores) 1 Figure 6.10 Ribosomes. Ths electron micrograph of a pancreas © mastering Blotogy ‘el shows both fee and bound ribosomes. The simplified diagram and eaia an ea hocks computer model show the two subunits ofa ribosome. [Saxivive orcmeceeicnire Free ribosomes in eyosol Endoplasmic reveulum (ER) Large subunit mall subunit ‘TEM showing ER and ribosomes Diagram of aribosome Computer model ofa ribosome 9"20h that might be making a protein that vl be secreted, chaPTER6 ATouroftheCell 103 Celis with high rates of protein synthesis have particularly large numbers of ribosomes as well as prominent nucleoli ‘which makes sense, given the role of nucleoli in nbosome ‘assembly. For example, a human pancreas cell, which makes ‘many digestive enzymes, has few million slbosomes. Ribosomes build proteins in two cytoplasmic regions: Atany given time, fee ribosomes are suspended in the cyto- sol, while Bound ribosomes are attached to the outside of the sndoplasmic reticulum or nuclear envelope (see Figute 6.10). Bound and free ribosomes are structurally identical, and ribosomes can play ether role at different times. Most of the proteins made on free ribosomes function within the cytosol; ‘examples are enzymes that catalyze the first steps of sugar breakdown. Bound ribosomes generally make proteins that are destined for insertion into membranes, for packaging within certain organelles such as lysosomes (see Figure 68) or for export from the cell (secretion). Cells that specialize in protein secretion—for instance, the cells ofthe pancreas that secrete digestive enaymes—frequently havea high propor tion of bound ribosomes. (You will Ian more about rbo- ‘some structure and function in Concept 17.4) +1. Whatroledorbosomes play in carving out geneticinstuctions? 2. Describe the molecular composition of nucleo and explain their function, 13, WHAT IF? As 2 cel begins the process of dividing, ts chromozomes become shorter, thicker, and individually ‘aie in an LM (ight micrograph), Explain what happening at the molecular level For suggested answers 38 Appenac A Be eee The endomembrane system regulates protein traffic and performs metabolic functions “Many ofthe different membrane-bounded organelle of the cearyotic cell are part ofthe endomembrane system, ‘which includes the nuclear envelope, the endoplasmic reticu- tum, the Gol apparatus, lysosomes, various kinds of vesicles and vacuoles, and the plasma membrane. This system cates, cout variety of tasks in the cll, including synthesis of pro- teins, transport of proteins into membranes and organelles ‘rout ofthe cell, metabolism and movement of lipids, and ) Givean example of

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