Chapter 6

A Tour of the Cell

Lecture Outline
Overview: The Fundamental Units of Life
All organisms are made of cells. Many organisms are single-celled. The cell is the simplest collection of matter that can live. Even when arranged into higher levels of organization, such as tissues and organs, cells are an organisms asic units of structure and function. !ife at the cellular level arises from structural order, reflecting emergent properties and the correlation etween structure and function. The movement of an animal cell depends on the intricate interplay of the structures that ma"e up a cellular s"eleton. #rganisms interact with their environment$ cells sense and respond to environmental fluctuations. Evolution is the unifying iological theme$ all cells are related y their descent from earlier cells ut have een modified in various ways during the history of life on Earth.

Concept 6.1 To study cells !iolo"ists use microscopes and the tools of !iochemistry.
The discovery and early study of cells progressed with the invention of microscopes in %&'( and their improvement in the %)th century. Microscopes vary in magnification and resolving power, or resolution. Magnification is the ratio of an o *ects image to its real size. Resolution is a measure of image clarity. +t is the minimum distance two points can e separated and still e distinguished as two separate points. ,esolution is limited y the shortest wavelength of the radiation used for imaging. +n a li"ht microscope #L$%, visi le light passes through the specimen and then through glass lenses. The lenses refract light so that the image is magnified into the eye or onto a video screen. The minimum resolution of an !M is a out -(( nanometers .nm/, the size of a small acterium. !Ms can magnify effectively to a out %,((( times the size of the actual specimen. At higher magnifications, the image lurs.

Techni0ues developed in the -(th century have enhanced contrast, staining or la eling cell components so they stand out.
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Although an !M can resolve individual cells, it cannot resolve much of the internal anatomy, especially the or"anelles. To resolve smaller structures, scientists use an electron microscope #&$%, which focuses a eam of electrons through the specimen or onto its surface. 1ecause resolution is inversely related to the wavelength used, EMs .whose electron eams have shorter wavelengths than visi le light/ have finer resolution than !Ms. Theoretically, the resolution of a modern EM could reach (.((- nm, ut the practical limit is closer to a out - nm. The term cell ultrastructure refers to a cells anatomy revealed y an EM. 'cannin" electron microscopes #'&$s% are useful for studying surface structures. The sample surface is covered with a thin film of gold. The eam e2cites electrons on the surface of the sample. These secondary electrons are collected and focused on a screen, producing an image of the topography of the specimen. 3EMs have great depth of field, resulting in an image that seems three-dimensional. Transmission electron microscopes #T&$s% are used to study the internal ultrastructure of cells. A TEM aims an electron eam through a very thin section of the specimen. To enhance contrast, the thin sections are stained with atoms of heavy metals. The image is focused and magnified y electromagnets. Although EMs reveal organelles that are impossi le to resolve with !Ms, the methods used to prepare cells for viewing under an EM "ills them. !Ms do not have as high a resolution as EMs, ut they can e used to study live cells. Microscopes are important tools in cytology, the study of cell structures. 4ytology com ined with biochemistry, the study of molecules and chemical processes in meta olism, produced modern cell iology. 4ell iologists can isolate organelles to study their functions. A centrifuge spins test tu es holding mi2tures of disrupted cells at various speeds. The resulting forces cause a fraction of the cell components to settle to the ottom of the tu e, forming a pellet. At lower speeds, the pellet consists of larger components$ higher speeds yield a pellet with smaller components. An ultracentrifuge can spin at up to %5(,((( revolutions per minute .rpm/ and apply forces greater than % million times gravity .%,(((,((( g/. Cell fractionation prepares isolates of specific cell components so that the functions of these organelles can e determined, especially y the reactions or processes catalyzed y their proteins. 6or e2ample, one cellular fraction was enriched in enzymes that function in cellular respiration. Electron microscopy revealed that this fraction is rich in mitochondria. This evidence helped cell iologists determine that mitochondria are the site of cellular respiration. 4ytology and iochemistry complement each other in correlating cellular structure and function.
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Concept 6.( &u)aryotic cells have internal mem!ranes that compartmentali*e their functions.
Prokaryotic and eukaryotic cells differ in size and complexity. #rganisms of the domains 1acteria and Archaea consist of pro"aryotic cells. 7rotists, fungi, animals, and plants consist of eu"aryotic cells. All cells are surrounded y a selective arrier, the plasma membrane. The semifluid su stance within the mem rane is the cytosol, containing the organelles. All cells contain chromosomes that carry genes in the form of 89A. All cells have ribosomes, tiny comple2es that ma"e proteins ased on instructions contained in genes. A ma*or difference etween pro"aryotic and eu"aryotic cells is the location of the 89A. +n a eu)aryotic cell, most of the 89A is in an organelle ounded y a dou le mem rane, the nucleus. +n a pro)aryotic cell, the 89A is concentrated in the nucleoid without a mem rane separating it from the rest of the cell. The interior of a pro"aryotic cell and the region etween the nucleus and the plasma mem rane of a eu"aryotic cell is the cytoplasm. :ithin the cytoplasm of a eu"aryotic cell are a variety of mem rane- ound organelles with specialized form and function. These mem rane- ound organelles are a sent in pro"aryotes. Eu"aryotic cells are generally much larger than pro"aryotic cells. The logistics of carrying out cellular meta olism set limits on cell size. At the lower limit, the smallest acteria, mycoplasmas, are (.%;%.( <m in diameter. Most acteria are %;%( <m in diameter. Eu"aryotic cells are typically %(;%(( <m in diameter. Meta olic re0uirements also set an upper limit to the size of a single cell. As a cell increases in size, its volume increases faster than its surface area. 3maller o *ects have a higher ratio of surface area to volume. The plasma mem!rane functions as a selective arrier that allows the passage of o2ygen, nutrients, and wastes for the whole volume of the cell. ,ates of chemical e2change across the plasma mem rane may e inade0uate to maintain a cell with a very large cytoplasm. The need for a surface sufficiently large to accommodate the volume e2plains the microscopic size of most cells. !arger organisms do not generally have larger cells than smaller organisms, simply more cells. 4ells that e2change a lot of material with their surroundings, such as intestinal cells, may have long, thin pro*ections from the cell surface called microvilli, which increase the surface area without significantly increasing the cell volume. A eu"aryotic cell has e2tensive and ela orate internal mem ranes, which partition the cell
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Internal membranes compartmentalize the functions of a eukaryotic cell.

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into compartments. These mem ranes also participate directly in meta olism ecause many enzymes are uilt into mem ranes. The compartments created y mem ranes provide different local environments that facilitate specific meta olic functions, allowing several incompati le processes to go on simultaneously in a cell. The general structure of a iological mem rane is a dou le layer of phospholipids. #ther lipids and diverse proteins are em edded in the lipid ilayer or attached to its surface. Each type of mem rane has a uni0ue com ination of lipids and proteins for its specific functions. 6or e2ample, enzymes em edded in the mem ranes of mitochondria function in cellular respiration.

Concept 6.+ The eu)aryotic cell,s "enetic instructions are housed in the nucleus and carried out !y the ri!osomes.
The nucleus contains most of the genes in a eu"aryotic cell. Additional genes are located in mitochondria and chloroplasts. The nucleus averages a out & <m in diameter. The nucleus is separated from the cytoplasm y a double mem rane called the nuclear envelope. The two mem ranes of the nuclear envelope are separated y -(;=( nm. The envelope is perforated y pores that are a out %(( nm in diameter. At the lip of each pore, the inner and outer mem ranes of the nuclear envelope are fused to form a continuous mem rane. A protein structure called a pore complex lines each pore, regulating the passage of certain large macromolecules and particles. The nuclear side of the envelope is lined y the nuclear lamina, a networ" of protein filaments that maintains the shape of the nucleus. There is evidence that a framewor" of fi ers called the nuclear matrix e2tends through the nuclear interior. :ithin the nucleus, the 89A and associated proteins are organized into discrete units called chromosomes, structures that carry the genetic information. Each chromosome is made up of material called chromatin a comple2 of proteins and 89A. 3tained chromatin appears through light microscopes and electron microscopes as a diffuse mass. As the cell prepares to divide, the chromatin fi ers coil up and condense, ecoming thic" enough to e recognized as the familiar chromosomes. Each eu"aryotic species has a characteristic num er of chromosomes. A typical human cell has => chromosomes. A human se2 cell .egg or sperm/ has only -5 chromosomes. +n the nucleus is a region of densely stained fi ers and granules ad*oining chromatin, the nucleolus.
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+n the nucleolus, ribosomal RNA .r,9A/ is synthesized and assem led with proteins from the cytoplasm to form large and small ri osomal su units. The su units pass through the nuclear pores to the cytoplasm, where they com ine to form ri osomes. ,ecent studies have suggested that the nucleolus also helps regulate cellular processes such as cell division. The nucleus directs protein synthesis y synthesizing messenger ,9A .m,9A/. The m,9A is transported to the cytoplasm through the nuclear pores. #nce in the cytoplasm, ri osomes translate m,9As genetic message into the primary structure of a specific polypeptide. -i!osomes, containing r,9A and protein, are the cellular components that carry out protein synthesis. 4ell types that synthesize large 0uantities of proteins .such as pancreas cells/ have large num ers of ri osomes and prominent nucleoli. Free ribosomes are suspended in the cytosol and synthesize proteins that function within the cytosol. Bound ribosomes are attached to the outside of the endoplasmic reticulum or nuclear envelope. 1ound ri osomes synthesize proteins that are inserted into mem ranes, pac"aged into organelles such as ri osomes, or e2ported .secreted/ from the cell. 4ells that specialize in protein secretion?for instance, the cells of the pancreas that secrete digestive enzymes?fre0uently have a high proportion of ound ri osomes.

Concept 6.. The endomem!rane system re"ulates protein traffic and performs meta!olic functions in the cell.
Many of the internal mem ranes in a eu"aryotic cell are part of the endomem!rane system. The tas"s of the endomem rane system include synthesis of proteins and their transport into mem ranes and organelles or out of the cell, meta olism and movement of lipids, and deto2ification of poisons. These mem ranes are either directly continuous or connected via the transfer of vesicles, sacs of mem rane. +n spite of the connections, these mem ranes are diverse in function and structure. The thic"ness, molecular composition, and types of chemical reactions carried out y proteins in a given mem rane may e modified several times during a mem ranes life. The endomem rane system includes the nuclear envelope, endoplasmic reticulum, @olgi apparatus, lysosomes, vacuoles, and plasma mem rane.

The endoplasmic reticulum manufactures membranes and performs many other biosynthetic functions. The endoplasmic reticulum #&-% accounts for more than half the mem ranes in a eu"aryotic cell. The E, includes a networ" of mem ranous tu ules and sacs called cisternae. The E, mem rane is continuous with the nuclear envelope, and the cisternal space of the E, is continuous with the space etween the two mem ranes of the nuclear envelope.
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Lecture Outline for Campbell/Reece Biology, 8th Edition, Pearson Education, Inc.

There are two connected regions of E, that differ in structure and function. 'mooth &- loo"s smooth ecause it lac"s ri osomes. -ou"h &- loo"s rough ecause ri osomes . ound ri osomes/ are attached to the outside, including the outside of the nuclear envelope. 3mooth E, is rich in enzymes and plays a role in a variety of meta olic processes, including synthesis of lipids, meta olism of car ohydrates, and deto2ification of drugs and poisons. Enzymes of smooth E, synthesize lipids, including oils, phospholipids, and steroids. These include the se2 hormones of verte rates and adrenal steroids. +n the smooth E, of the liver, enzymes help deto2ify poisons and drugs such as alcohol and ar iturates. 6re0uent use of these drugs leads to the proliferation of smooth E, in liver cells, increasing the rate of deto2ification. This proliferation of smooth E, increases tolerance to the target and other drugs, so higher doses are re0uired to achieve the same effect. 3mooth E, stores calcium ions. Muscle cells have a specialized smooth E, that pumps calcium ions from the cytosol into the E, lumen. :hen a nerve impulse stimulates a muscle cell, calcium ions rush from the E, into the cytosol, triggering contraction of the muscle cell. ,ough E, is especially a undant in cells that secrete proteins. 4ertain cells in the pancreas secrete the hormone insulin into the loodstream. As a polypeptide chain grows from a ound ri osome, it is threaded into the E, lumen through a pore formed y a protein comple2 in the E, mem rane. As the new protein enters the E, lumen, it folds into its native shape. Most secretory polypeptides are "lycoproteins, proteins to which a car ohydrate is attached. The car ohydrate is attached to the protein in the E, y specialized molecules uilt into the E, mem rane. 3ecretory proteins are pac"aged in transport vesicles that ud from a specialized region called transitional E,. Transport vesicles carry proteins from one part of the cell to another. ,ough E, is also a mem rane factory for the cell. Mem rane- ound proteins are synthesized directly into the E, mem rane and anchored y their own hydropho ic portions. Enzymes in rough E, also synthesize phospholipids from precursors in the cytosol. As the E, mem rane e2pands, mem rane can e transferred as transport vesicles to other components of the endomem rane system. Many transport vesicles from the E, travel to the /ol"i apparatus for modification of their contents. The @olgi apparatus is a center of manufacturing, warehousing, sorting, and shipping. 3ome products of the E, are modified and stored and then sent on. The @olgi apparatus is especially e2tensive in cells specialized for secretion. The @olgi apparatus consists of flattened mem ranous sacs?cisternae?that loo" li"e a stac"
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The Golgi apparatus is the shipping and receiving center for cell products.

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of pita read. The mem rane of each cisterna separates its internal space from the cytosol. #ne side of the @olgi apparatus, the cis face, is located near the E,. The cis face receives material y fusing with transport vesicles from the E,. The other side, the trans face, uds off vesicles that travel to other sites. 7roducts from the E, are usually modified during their transit from the cis to the trans region of the @olgi apparatus. Aarious @olgi enzymes modify the car ohydrate portions of glycoproteins. 4ar ohydrates are first added to proteins in rough E,, often during the process of polypeptide synthesis. The car ohydrate on the resulting glycoprotein is modified as it passes through the rest of the E, and the @olgi. The @olgi removes some sugar monomers and su stitutes others, producing a large variety of car ohydrates. The @olgi can also manufacture its own macromolecules, including pectin and other noncellulose polysaccharides. @olgi products that will e secreted depart from the trans face of the @olgi inside transport vesicles that eventually fuse with the plasma mem rane. The @olgi apparatus is a dynamic structure. The @olgi manufactures and refines its products in stages, with different cisternae etween the cis and trans regions containing uni0ue teams of enzymes. According to the cisternal maturation model, the cisternae of the @olgi progress from the cis to the trans face, carrying and modifying their protein cargo as they move. 6inally, the @olgi sorts and pac"ages materials into transport vesicles. Molecular identification tags such as phosphate groups are added to products to aid in sorting. These tags act li"e B+7 codes on mailing la els to identify the products final destination. Transport vesicles udded from the @olgi may have e2ternal molecules on their mem ranes that recognize Cdoc"ing sitesD on the surface of specific organelles or on the plasma mem rane, thus targeting them appropriately. A lysosome is a mem rane- ound sac of hydrolytic enzymes that an animal cell uses to digest macromolecules. !ysosomal enzymes wor" est at acidic pE. The rupture of one or a few lysosomes has little impact on a cell ecause the lysosomal enzymes are not very active at the neutral pE of the cytosol. Massive rupture of many lysosomes can destroy a cell y autodigestion, however. !ysosomal enzymes and mem rane are synthesized y rough E, and then transferred to the @olgi apparatus for further modification. 3ome lysosomes arise y udding from the trans face of the @olgi apparatus. 7roteins on the inner surface of the lysosomal mem rane are spared y digestion y their three-dimensional conformations, which protect vulnera le onds from hydrolysis. !ysosomes carry out intracellular digestion in a variety of circumstances.
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Lysosomes are digestive compartments.

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Amoe as eat y engulfing smaller organisms y pha"ocytosis. The food vacuole formed y phagocytosis fuses with a lysosome, whose enzymes digest the food. As the polymers are digested, monomers pass to the cytosol to ecome nutrients for the cell. Euman macrophages help defend the ody y phagocytosing and destroying invasive acteria.

!ysosomes can play a role in recycling the cells organelles and macromolecules. This recycling, or autophagy, renews the cell. 8uring autophagy, a damaged organelle or region of cytosol ecomes surrounded y a dou le mem rane of un"nown origin. A lysosome fuses with the outer mem rane of the vesicle, digesting the macromolecules and returning the organic monomers to the cytosol for reuse. The cells of people who have inherited lysosomal storage diseases lac" a functioning hydrolytic enzyme normally present in lysosomes. The lysosomes ecome engorged with indigesti le su strates, which egin to interfere with other cellular activities. +n people who have Tay-3achs disease, a lipid-digesting enzyme is missing or inactive, and the rain ecomes impaired y an accumulation of lipids in the cells. acuoles have diverse functions in cell maintenance. Aesicles and vacuoles .larger versions/ are mem rane- ound sacs with varied functions. Food vacuoles are formed y phagocytosis and fuse with lysosomes. Contractile vacuoles, found in freshwater protists, pump e2cess water out of the cell to maintain the appropriate concentration of salts. A large central vacuole is found in many mature plant cells. The vacuolar mem rane is selective in its transport of solutes into the central vacuole. As a result, the solution inside the vacuole, called cell sap, differs in composition from the cytosol. The functions of the central vacuole include stoc"piling proteins or inorganic ions, disposing of meta olic y-products, holding pigments, and storing defensive compounds that protect the plant against her ivores. The vacuole has a ma*or role in the growth of plant cells, which enlarge as their vacuoles a sor water, ena ling the cell to ecome larger with little investment in new cytoplasm. 1ecause of the large vacuole, the cytosol occupies only a thin layer etween the plasma mem rane and the central vacuole. The presence of a large vacuole increases the ratio of surface area to volume for the cell.

Concept 6.0 $itochondria and chloroplasts chan"e ener"y from one form to another.
Mitochondria and chloroplasts are the organelles that convert energy to forms that cells can use for wor". $itochondria are the sites of cellular respiration, generating AT7 y e2tracting energy from sugars, fats, and other fuels with the help of o2ygen.
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Chloroplasts, found in plants and algae, are the sites of photosynthesis. 4hloroplasts convert solar energy to chemical energy and synthesize new organic compounds such as sugars from 4#- and E-#. Mitochondria and chloroplasts are not part of the endomem rane system. +n contrast to organelles of the endomem rane system, each mitochondrion or chloroplast has two mem ranes separating the innermost space from the cytosol. Their mem rane proteins are not made y ri osomes ound to the E,, ut rather y free ri osomes in the cytosol and y ri osomes within the organelles themselves. 1oth mitochondria and chloroplasts have small 0uantities of 89A that direct the synthesis of the polypeptides produced y these internal ri osomes. Mitochondria and chloroplasts grow and reproduce as semiautonomous organelles. The pero1isome is an o2idative organelle that is not part of the endomem rane system. !i"e mitochondria and chloroplasts, the pero2isome imports its proteins primarily from the cytosol. Almost all eu"aryotic cells have mitochondria. Even in e2ceptions, such as the human intestinal parasite Giardia and some other protists, recent studies have identified closely related organelles that may have evolved from mitochondria. 4ells may have one very large mitochondrion or hundreds to thousands of individual mitochondria. The num er of mitochondria is correlated with aero ic meta olic activity. A typical mitochondrion is %;%( <m long. Mitochondria are dynamicF moving, changing shape, fusing, and dividing. Mitochondria have a smooth outer mem rane and a convoluted inner mem rane with infoldings called cristae. The inner mem rane divides the mitochondrion into two internal compartments. The first compartment is the intermem rane space, a narrow region etween the inner and outer mem ranes. The inner mem rane encloses the mitochondrial matri1, a fluid-filled space with mitochondrial 89A, ri osomes, and enzymes. 3ome of the meta olic steps of cellular respiration are catalyzed y enzymes in the matri2. #ther proteins that function in respiration, including the enzyme that ma"es AT7, are uilt into the inner mem rane. The highly folded cristae present a large surface area for these enzymes. The chloroplast is one of several mem ers of a related family of plant structures called plastids. Amyloplasts are colorless plastids that store starch in roots and tu ers. Chromoplasts store pigments for fruits and flowers. Chloroplasts contain the green pigment chlorophyll as well as enzymes and other molecules that function in the photosynthetic production of sugar. 4hloroplasts are -;& <m in diameter and are found in leaves and other green organs of plants and algae. The contents of the chloroplast are separated from the cytosol y an envelope consisting of two mem ranes separated y a narrow intermem rane space.
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+nside the innermost mem rane is a fluid-filled space, the stroma, in which float mem ranous sacs, the thyla)oids. The stroma contains chloroplast 89A, ri osomes, and enzymes. The thyla"oids are flattened sacs that play a critical role in converting light to chemical energy. +n some regions, thyla"oids are stac"ed li"e po"er chips into "rana. The mem ranes of the chloroplast divide the chloroplast into three compartmentsF the intermem rane space, the stroma, and the thyla"oid space. !i"e mitochondria, chloroplasts are dynamic structures. The shape of mitochondria is plastic, and they can reproduce themselves y pinching in two. Mitochondria and chloroplasts are mo ile and move around the cell along trac"s of the cytos"eleton. 7ero2isomes, ound y a single mem rane, contain enzymes that transfer hydrogen from various su strates to o2ygen. A y-product of this process is hydrogen pero2ide .E-#-/. 3ome pero2isomes rea" fatty acids down to smaller molecules that are transported to mitochondria as fuel for cellular respiration. 7ero2isomes in the liver deto2ify alcohol and other harmful compounds y transferring hydrogen from these molecules to o2ygen. The E-#- formed y pero2isomes is itself to2ic, ut pero2isomes also contain an enzyme that converts E-#- to water. The enzymes that produce hydrogen pero2ide and those that dispose of it are se0uestered in the same space, away from other cellular components that could otherwise e damaged. 3pecialized pero2isomes, glyoxysomes, convert the fatty acids in seeds to sugars, which the seedling can use as a source of energy and car on until it is capa le of photosynthesis. 7ero2isomes do not ud from the endomem rane system, ut y incorporating proteins and lipids from the cytosol, as well as lipids synthesized within the pero2isome itself. 7ero2isomes split in two when they reach a certain size.

Peroxisomes generate and degrade !"O" in performing various metabolic functions.

Concept 6.6 The cytos)eleton is a networ) of fi!ers that or"ani*es structures and activities in the cell.
The cytoskeleton provides support# motility# and regulation. The cytos)eleton provides mechanical support to the cell and maintains the cells shape. This function is especially important in animal cells, which lac" walls. The cytos"eleton provides anchorage for many organelles and cytosolic enzymes. The cytos"eleton is dynamic and can e dismantled in one part and reassem led in another to change the shape of the cell. The cytos"eleton also plays a ma*or role in cell motility, including changes in cell location and limited movements of parts of the cell. The cytos"eleton interacts with motor proteins to produce motility. 4ytos"eleton elements and motor proteins wor" together with plasma mem rane molecules to move the whole cell along fi ers outside the cell.
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Motor proteins ring a out movements of cilia and flagella y gripping cytos"eletal components such as microtu ules and moving them past each other. A similar mechanism causes muscle cells to contract.

+nside the cell, vesicles can travel along CmonorailsD provided y the cytos"eleton. This is how vesicles containing neurotransmitter molecules migrate to the tips of a2ons. The vesicles that ud off from the E, travel to the @olgi apparatus along trac"s uilt of cytos"eletal elements. The cytos"eleton manipulates the plasma mem rane to form food vacuoles during phagocytosis. 4ytoplasmic streaming in plant cells is caused y the cytos"eleton. The cytos"eleton also plays a role in the regulation of iochemical activities in the cell in response to mechanical stimulation. 4ytos"eletal elements transmit forces e2erted y e2tracellular molecules via cell surface proteins into the cell?and even into the nucleus. +n an e2periment, investigators used a micromanipulation device to pull on certain plasma mem rane proteins attached to the cytos"eleton, leading to almost instantaneous rearrangements of nucleoli and other structures in the nucleus. +n this way, cytos"eletal transmission of naturally occurring mechanical signals may help regulate and coordinate the cells response. Three main types of fibers make up the cytoskeleton$ microtubules# microfilaments# and intermediate filaments.

Microtubules are the thic"est of the three types of fi ers$ microfilaments .or actin filaments/ are the thinnest$ and intermediate filaments are fi ers with diameters in a middle range. $icrotu!ules are hollow rods a out -& nm in diameter and -(( nm to -& <m in length. Microtu ule fi ers are constructed of the glo ular protein tu ulin. Each tu ulin molecule is a dimer consisting of two su units. A tu ulin dimer consists of two slightly different polypeptidesF -tu ulin and -tu ulin. A microtu ule changes in length y adding or removing tu ulin dimers. The two ends of a microtu ule are slightly different. The CplusD end can accumulate or release tu ulin dimers at a much higher rate than the other end. Microtu ules shape and support the cell and serve as trac"s to guide motor proteins carrying organelles to their destination. Microtu ules guide secretory vesicles from the @olgi apparatus to the plasma mem rane. Microtu ules are also responsi le for the separation of chromosomes during cell division. +n many animal cells, microtu ules grow out from a centrosome near the nucleus. This region is considered to e a Cmicrotu ule-organizing center.D These microtu ules resist compression to the cell. :ithin the centrosome is a pair of centrioles, each with nine triplets of microtu ules arranged in a ring. 1efore an animal cell divides, the centrioles replicate. Although centrosomes with centrioles may help organize microtu ule assem ly in animal cells, they are not essential for this function.
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Geast cells and plant cells lac" centrosomes with centrioles ut have well-organized microtu ules. #ther microtu ule-organizing centers must play the role of centrosomes in these cells.

A specialized arrangement of microtu ules is responsi le for the eating of cilia and fla"ella. Many unicellular eu"aryotic organisms are propelled through water y cilia and flagella. The sperm of animals, algae, and some plants have flagella. :hen cilia or flagella e2tend from cells within a tissue layer, they can eat to move fluid over the surface of the tissue. 6or e2ample, cilia lining the windpipe sweep mucus carrying trapped de ris out of the lungs. +n the reproductive tract, cilia lining the oviducts help move an egg toward the uterus. 4ilia usually occur in large num ers on the cell surface. 4ilia are a out (.-& <m in diameter and -;-( <m long. There are usually *ust one or a few flagella per cell. 6lagella are the same diameter as cilia ut are %(;-(( <m long. 4ilia and flagella differ in their eating patterns. A flagellum has an undulatory movement that generates force in the same direction as the flagellums a2is. 4ilia move more li"e oars, with alternating power and recovery stro"es that generate force perpendicular to the ciliums a2is. 4ilia may also act as a "ind of CantennaD for the cell. Mem rane proteins on the cilium transmit molecular signals from the environment outside the cell to the cells nucleus, resulting in changes in the cells ehavior. 3uch pathways involving cilia- ased signaling appear to e crucial to rain function and to em ryonic development. A growing num er of human genetic diseases have een traced ac" to malfunctioning cilia. +n spite of their differences, oth cilia and flagella have the same ultrastructure. 1oth have a core of microtu ules sheathed y an e2tension of the plasma mem rane. 9ine dou lets of microtu ules are arranged in a ring around a pair at the center. This C' H -D pattern is found in nearly all eu"aryotic cilia and flagella. 6le2i le cross-lin"ing proteins connect outer dou lets to each other and to the two central microtu ules. The cilium or flagellum is anchored in the cell y a !asal !ody, whose structure is identical to that of a centriole. +n many animals .including humans/, the asal ody of the fertilizing sperms flagellum enters the egg and ecomes a centriole. The ending of cilia and flagella is driven y the arms of a motor protein, dynein. A dynein performs a comple2 cycle of movements caused y changes in the shape of the protein, powered y AT7. The mechanics of dynein- ased ending involve a process that resem les wal"ing. A typical dynein protein has two CfeetD that Cwal"D along the microtu ule of the ad*acent dou let$ one foot maintains contact while the other releases and reattaches one step farther along the microtu ule.
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Lecture Outline for Campbell/Reece Biology, 8th Edition, Pearson Education, Inc.

:ithout restraint on the movement of the microtu ule dou lets, one dou let continues to Cwal"D along and slide past the surface of the other, elongating the cilium or flagellum rather than ending it. 6or lateral movement of a cilium or flagellum, the dynein Cwal"ingD must have something to pull against, as when the muscles in your leg pull against your ones to move your "nee. +n cilia and flagella, the microtu ule dou lets seem to e held in place y the cross-lin"ing proteins *ust inside the outer dou lets and y the radial spo"es and other structural elements. Thus, neigh oring dou lets cannot slide past each other very far. +nstead, the forces e2erted y dynein Cwal"ingD cause the dou lets to curve, ending the cilium or flagellum. $icrofilaments are solid rods a out ) nm in diameter. Microfilaments are also called actin filaments ecause they are uilt from molecules of actin, a glo ular protein. Each microfilament is uilt as a twisted dou le chain of actin su units. Microfilaments can form structural networ"s ecause of their a ility to ranch. Microfilaments are present in all eu"aryotic cells. The structural role of microfilaments in the cytos"eleton is to ear tension, resisting pulling forces within the cell. Cortical microfilaments form a three-dimensional networ" *ust inside the plasma mem rane to help support the cells shape, giving the cell corte1 the semisolid consistency of a gel. This consistency contrasts with the more fluid .sol/ state of the interior cytoplasm. +n animal cells specialized for transporting materials across the plasma mem rane, such as intestinal cells, undles of microfilaments ma"e up the core of microvilli. Microfilaments are important in cell motility, especially as part of the contractile apparatus of muscle cells. +n muscle cells, thousands of actin filaments are arranged parallel to one another. Thic"er filaments composed of myosin interdigitate with the thinner actin fi ers. !i"e dynein with microtu ules, myosin acts as a microfilament- ased motor protein, wal"ing along the actin filaments to shorten the cell. +n other cells, actin-myosin aggregates are less organized ut still cause localized contraction. A contracting elt of microfilaments forms a cleavage furrow that divides the cytoplasm of animal cells during cell division. !ocalized contraction rought a out y actin and myosin also drives amoe oid movement. 2seudopodia, cellular e2tensions, e2tend and contract through the reversi le assem ly and contraction of actin su units into microfilaments. Microfilaments assem le into networ"s that convert sol to gel. According to a widely accepted model, filaments near the cells trailing edge interact with myosin, causing contraction. The contraction forces the interior fluid into the pseudopodium, where the actin networ" has een wea"ened. The pseudopodium e2tends until the actin reassem les into a networ". Many cells in the animal ody, including some white lood cells, also move y crawling. +n plant cells, actin-myosin interactions and sol-gel transformations drive cytoplasmic
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Lecture Outline for Campbell/Reece Biology, 8th Edition, Pearson Education, Inc.

streamin" which creates a circular flow of cytoplasm in the cell, speeding the distri ution of materials within the cell. 3ntermediate filaments range in diameter from I to %- nm, larger than microfilaments ut smaller than microtu ules. +ntermediate filaments are a diverse class of cytos"eletal units, uilt from a family of proteins that includes the "eratins. !i"e microfilaments, intermediate filaments are specialized for earing tension. +ntermediate filaments are more permanent fi2tures of the cytos"eleton than are the other two classes. Even after cells die, intermediate filament networ"s often persist. The networ"s reinforce cell shape and fi2 organelle location. The nucleus sits within a cage made of intermediate filaments, fi2ed in location y ranches of the filaments that e2tend into the cytoplasm. #ther intermediate filaments ma"e up the nuclear lamina that lines the interior of the nuclear envelope. :hen the shape of the entire cell is correlated with function, intermediate filaments support that shape. The a2ons of nerve cells are strengthened y one class of intermediate filament. Aarious "inds of intermediate filaments function as the framewor" of the entire cytos"eleton.

Concept 6.4 &1tracellular components and connections !etween cells help coordinate cellular activities.
Plant cells are encased by cell %alls. Cell walls are found in pro"aryotes, fungi, and some protists. +n plants, the cell wall protects the cell, maintains its shape, prevents e2cessive upta"e of water, and supports the plant against the force of gravity. The thic"ness of plant cell walls ranges from (.% <m to several micrometers. The chemical composition of the cell wall differs with species and among cell types within a plant. The asic design of cell walls consists of microfi rils of cellulose synthesized y an enzyme called cellulose synthase. The cellulose microfi rils are secreted to the e2tracellular space, where they are em edded in a matri2 of proteins and other polysaccharides. This is the asic design of steel-reinforced concrete or fi erglass. A young plant cell secretes a relatively thin and fle2i le wall called the primary cell wall. +n actively growing cells, the cellulose fi rils are oriented at right angles to the direction of cell e2pansion, possi ly affecting the growth pattern. 8avid Ehrhardt and colleagues found that cortical microtu ules guide cellulose synthase as it synthesizes and deposits the fi rils. 1y orienting the deposition of cellulose, microtu ules affect the growth pattern of the cells. 1etween the primary walls of ad*acent cells is a middle lamella, a thin layer with stic"y
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Lecture Outline for Campbell/Reece Biology, 8th Edition, Pearson Education, Inc.

polysaccharides called pectins that glues cells together. :hen a plant cell stops growing, it strengthens its wall y secreting hardening su stances into the primary wall or y adding a secondary cell wall etween the plasma mem rane and the primary wall. The secondary wall may e deposited in several layers. +t has a strong and dura le matri2 that provides support and protection. :ood consists mainly of secondary walls. 7lant cell walls are perforated y channels etween ad*acent cells called plasmodesmata.

The extracellular matrix of animal cells provides support# adhesion# movement# and regulation. Though lac"ing cell walls, animal cells do have an ela orate e1tracellular matri1 #&C$%. The primary constituents of the E4M are glycoproteins, especially colla"en fi ers, em edded in a networ" of glycoprotein proteo"lycans. 4ollagen accounts for a out half the total protein in the human ody. A proteoglycan molecule consists of a small core protein with many car ohydrate chains covalently attached, so that it may e up to '&J car ohydrate. !arge proteoglycan comple2es can form when hundreds of proteoglycans ecome noncovalently attached to a single long polysaccharide molecule. +n many cells, fi!ronectins in the E4M connect to inte"rins cell-surface receptor proteins that span the mem rane and ind on their cytoplasmic side to proteins attached to microfilaments of the cytos"eleton. The interconnections from the E4M to the cytos"eleton via the fi ronectin-integrin lin" permit the integration of changes inside and outside the cell. The E4M can regulate cell ehavior. Em ryonic cells migrate along specific pathways y matching the orientation of their microfilaments to the CgrainD of fi ers in the E4M. The E4M can influence the activity of genes in the nucleus via a com ination of chemical and mechanical signaling pathways. Mechanical signaling involves fi ronectin, integrins, and microfilaments of the cytos"eleton. 4hanges in the cytos"eleton may in turn trigger chemical signaling pathways inside the cell, leading to changes in the set of proteins eing made y the cell and thus changes in the cells function. This may coordinate the ehavior of all the cells within a tissue. 9eigh oring cells in tissues, organs, and organ systems often adhere, interact, and communicate through direct physical contact. 7lant cells are perforated with plasmodesmata, channels allowing cytosol to pass etween cells. :ater and small solutes can pass freely from cell to cell. +n certain circumstances, proteins and ,9A can e e2changed. Macromolecules eing transported to neigh oring cells reach the plasmodesmata y moving along fi ers of the cytos"eleton. Animals have three main types of intercellular lin"sF tight unctions, desmosomes, and gap
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Intercellular &unctions help integrate cells into higher levels of structure and function.

Lecture Outline for Campbell/Reece Biology, 8th Edition, Pearson Education, Inc.

unctions. All three types of intercellular *unctions are especially common in epithelial tissue, which lines the e2ternal and internal surfaces of the ody. +n ti"ht 5unctions mem ranes of ad*acent cells are fused, forming continuous elts around cells that prevent lea"age of e2tracellular fluid. 6esmosomes .or anchoring *unctions/ fasten cells together into strong sheets, much li"e rivets. +ntermediate filaments of "eratin reinforce desmosomes. /ap 5unctions .or communicating *unctions/ provide cytoplasmic channels etween ad*acent cells. 3pecial mem rane proteins surround these pores. +ons, sugars, amino acids, and other small molecules can pass. +n em ryos, gap *unctions facilitate chemical communication during development.

Lecture Outline for Campbell/Reece Biology, 8th Edition, Pearson Education, Inc.

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