CAMPBELL BIOLOGY IN FOCUS URRY CAIN WASSERMAN MINORSKY

CAMPBELL BIOLOGY IN FOCUS URRY  CAIN  WASSERMAN  MINORSKY

CAMPBELL BIOLOGY IN FOCUS URRY CAIN WASSERMAN MINORSKY REECE 4 A Tour of the Cell Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge, Simon Fraser University 2016 Pearson Education, Inc. SECOND EDITION Overview: The Fundamental Units of Life All organisms are made of cells The cell is the simplest collection of matter

that can be alive All cells are related by their descent from earlier cells Though cells can differ substantially from one another, they share common features 2016 Pearson Education, Inc. Concept 4.1: Biologists use microscopes and the tools of biochemistry to study cells Microscopy Most cells are too small to be seen by the unaided eye Scientists use microscopes to observe cells too small to be seen with the naked eye In a light microscope (LM), visible light is passed through a specimen and then through glass lenses

Lenses refract (bend) the light, so that the image is magnified 2016 Pearson Education, Inc. Three important parameters of microscopy Magnification, the ratio of an objects image size to its real size Resolution, the measure of the clarity of the image, or the minimum distance between two distinguishable points Contrast, visible differences in parts of the sample 2016 Pearson Education, Inc. LMs can magnify effectively to about 1,000 times the size of the actual specimen Various techniques enhance contrast and enable

cell components to be stained or labeled Most subcellular structures, including organelles (membrane-enclosed compartments), are too small to be resolved by light microscopy 2016 Pearson Education, Inc. 10 m 0.1 m Human height Length of some nerve and muscle cells Chicken egg

1 cm 100 mm 10 mm 1 mm 100 nm 10 nm 1 nm 0.1 nm 2016 Pearson Education, Inc. Frog egg Human egg Most plant and animal cells Nucleus

Most bacteria Mitochondrion EM 1 mm LM 1m Unaided eye Figure 4.2 Smallest bacteria Viruses Ribosomes

Proteins Lipids Small molecules Atoms Superresolution microscopy 10 m 0.1 m Human height Length of some nerve and muscle cells Chicken egg

1 cm 100 mm 10 mm 1 mm 100 nm 10 nm 1 nm 0.1 nm 2016 Pearson Education, Inc. Frog egg Human egg Most plant and animal cells

Nucleus Most bacteria Mitochondrion EM 1 mm LM 1m Unaided eye Figure 4.2 Smallest bacteria Viruses

Ribosomes Proteins Lipids Small molecules Atoms Superresolution microscopy Two basic types of electron microscopes (EMs) are used to study subcellular structures Scanning electron microscopes (SEMs) focus a beam of electrons onto the surface of a specimen, producing images that look three-dimensional Transmission electron microscopes (TEMs) focus a beam of electrons through a specimen TEM is used mainly to study the internal structure

of cells 2016 Pearson Education, Inc. Figure 4.3-1 50 mm Light Microscopy (LM) 2016 Pearson Education, Inc. Brightfield (unstained specimen) Brightfield (stained specimen)

Phase-contrast Differential-interference contrast (Nomarski) Figure 4.3-3 Electron Microscopy (EM) Longitudinal section of cilium Cross section of cilium Cilia

Scanning electron microscopy (SEM) 2016 Pearson Education, Inc. 2 mm Transmission electron 2 mm microscopy (TEM) Figure 4.3-3b Longitudinal section of cilium Cross section of cilium

2 mm Transmission electron microscopy (TEM) 2016 Pearson Education, Inc. Concept 4.2: Eukaryotic cells have internal membranes that compartmentalize their functions The basic structural and functional unit of every organism is one of two types of cells: prokaryotic or eukaryotic Organisms of the domains Bacteria and Archaea consist of prokaryotic cells Protists, fungi, animals, and plants all consist of eukaryotic cells

2016 Pearson Education, Inc. Comparing Prokaryotic and Eukaryotic Cells Basic features of all cells Plasma membrane Semifluid substance called cytosol Chromosomes (carry genes) Ribosomes (make proteins) 2016 Pearson Education, Inc. In a eukaryotic cell most of the DNA is in the

nucleus, an organelle that is bounded by a double membrane Prokaryotic cells are characterized by having No nucleus DNA in an unbound region called the nucleoid region No membrane-bound organelles Both types of cells contain cytoplasm bound by the plasma membrane 2016 Pearson Education, Inc. Figure 4.4 Fimbriae Nucleoid

Ribosomes Plasma membrane Bacterial chromosome Cell wall Capsule 0.5 mm Flagella (a) A typical rod-shaped bacterium 2016 Pearson Education, Inc. (b) A thin section through the bacterium Corynebacterium

diphtheriae (colorized TEM) Eukaryotic cells are generally much larger than prokaryotic cells Typical bacteria are 15 mm in diameter Eukaryotic cells are typically 10100 mm in diameter 2016 Pearson Education, Inc. The plasma membrane is a selective barrier that allows sufficient passage of oxygen, nutrients, and waste to service the volume of every cell The general structure of a biological membrane is a double layer of phospholipids 2016 Pearson Education, Inc.

**Figure 4.5 Outside of cell Inside of cell 0.1 mm (a) TEM of a plasma membrane Carbohydrate side chains Hydrophilic region Hydrophobic region

Hydrophilic region Phospholipid Proteins (b) Structure of the plasma membrane 2016 Pearson Education, Inc. A Panoramic View of the Eukaryotic Cell A eukaryotic cell has internal membranes that divide the cell into compartmentsorganelles The plasma membrane and organelle membranes participate directly in the cells metabolism 2016 Pearson Education, Inc.

**Figure 4.7-1 ENDOPLASMIC RETICULUM (ER) Smooth ER Flagellum Rough ER Nuclear envelope Nucleolus NUCLEUS Chromatin Centrosome Plasma membrane

CYTOSKELETON: Microfilaments Intermediate filaments Microtubules Microvilli Ribosomes (small brown dots) Golgi apparatus Peroxisome Lysosome Mitochondrion 2016 Pearson Education, Inc.

**Figure 4.7-2 Nuclear envelope Nucleolus Chromatin NUCLEUS Rough endoplasmic reticulum Smooth endoplasmic reticulum Ribosomes (small brown dots) Golgi apparatus

Central vacuole Microfilaments CYTOSKELETON Microtubules Mitochondrion Peroxisome Plasma membrane Chloroplast Cell wall Wall of adjacent cell 2016 Pearson Education, Inc. Plasmodesmata

Concept 4.3: The eukaryotic cells genetic instructions are housed in the nucleus and carried out by the ribosomes The nucleus contains most of the DNA in a eukaryotic cell Ribosomes use the information from the DNA to make proteins 2016 Pearson Education, Inc. The Nucleus: Information Central The nucleus contains most of the cells genes and is usually the most conspicuous organelle The nuclear envelope encloses the nucleus, separating it from the cytoplasm The nuclear membrane is a double membrane; each membrane consists of a lipid bilayer

2016 Pearson Education, Inc. Nuclear pores regulate the entry and exit of molecules The shape of the nucleus is maintained by the nuclear lamina, which is composed of protein 2016 Pearson Education, Inc. Figure 4.8 Nucleus 1 mm Nucleus

Nucleolus Chromatin Nuclear envelope: Inner membrane Outer membrane Nuclear pore Rough ER Surface of nuclear envelope (TEM) Pore complex Ribosome Pore complexes (TEM) 0.5 mm

0.25 mm Close-up of nuclear envelope Nuclear lamina (TEM) 2016 Pearson Education, Inc. Chromatin **Figure 4.8-1 Nucleus Nucleolus Chromatin

Nuclear envelope: Inner membrane Outer membrane Nuclear pore Rough ER Pore complex Ribosome Close-up of nuclear envelope 2016 Pearson Education, Inc. Chromatin Figure 4.8-2

1 mm Nuclear envelope: Inner membrane Outer membrane Nuclear pore Surface of nuclear envelope (TEM) 2016 Pearson Education, Inc. In the nucleus, DNA is organized into discrete units called chromosomes Each chromosome is one long DNA molecule associated with proteins The DNA and proteins of chromosomes together are called chromatin

Chromatin condenses to form discrete chromosomes as a cell prepares to divide The nucleolus is located within the nucleus and is the site of ribosomal RNA (rRNA) synthesis 2016 Pearson Education, Inc. Ribosomes: Protein Factories Ribosomes are complexes of ribosomal RNA and protein Ribosomes carry out protein synthesis in two locations In the cytosol (free ribosomes) On the outside of the endoplasmic reticulum or the nuclear envelope (bound ribosomes) 2016 Pearson Education, Inc.

Figure 4.9 0.25 mm Ribosomes ER Free ribosomes in cytosol Endoplasmic reticulum (ER) Ribosomes bound to ER Large subunit

TEM showing ER and ribosomes 2016 Pearson Education, Inc. Small subunit Diagram of a ribosome Computer model of a ribosome Figure 4.9-1 0.25 mm Free ribosomes in cytosol Endoplasmic reticulum (ER)

Ribosomes bound to ER Large subunit Small subunit TEM showing ER and ribosomes 2016 Pearson Education, Inc. Diagram of a ribosome Concept 4.4: The endomembrane system regulates protein traffic and performs metabolic functions in the cell Components of the endomembrane system

Nuclear envelope Endoplasmic reticulum Golgi apparatus Lysosomes Vacuoles Plasma membrane These components are either continuous or connected through transfer by vesicles 2016 Pearson Education, Inc.

The Endoplasmic Reticulum: Biosynthetic Factory The endoplasmic reticulum (ER) accounts for more than half of the total membrane in many eukaryotic cells The ER membrane is continuous with the nuclear envelope There are two distinct regions of ER Smooth ER: lacks ribosomes Rough ER: surface is studded with ribosomes 2016 Pearson Education, Inc. Figure 4.10 Smooth ER Rough ER

ER lumen Cisternae Ribosomes Transport vesicle 2016 Pearson Education, Inc. Nuclear envelope Transitional ER Smooth ER Rough ER

0.2 mm Functions of Smooth ER The smooth ER Synthesizes lipids Metabolizes carbohydrates Detoxifies drugs and poisons Stores calcium ions 2016 Pearson Education, Inc. Functions of Rough ER

The rough ER Has bound ribosomes, which secrete glycoproteins (proteins covalently bonded to carbohydrates) Distributes transport vesicles, proteins surrounded by membranes Is a membrane factory for the cell 2016 Pearson Education, Inc. The Golgi Apparatus: Shipping and Receiving Center The Golgi apparatus consists of flattened membranous sacs called cisternae Functions of the Golgi apparatus Modifies products of the ER Manufactures certain macromolecules Sorts and packages materials into transport vesicles

2016 Pearson Education, Inc. Figure 4.11 Golgi apparatus cis face (receiving side of Golgi apparatus) 0.1 mm Cisternae trans face (shipping side of Golgi apparatus)

2016 Pearson Education, Inc. TEM of Golgi apparatus Lysosomes: Digestive Compartments A lysosome is a membranous sac of hydrolytic enzymes that can digest macromolecules Lysosomal enzymes work best in the acidic environment inside the lysosome The three-dimensional shape of lysosomal proteins protects them from digestion by lysosomal enzymes 2016 Pearson Education, Inc. Some types of cell can engulf another cell by phagocytosis; this forms a food vacuole A lysosome fuses with the food vacuole and digests

the molecules Lysosomes also use enzymes to recycle the cells own organelles and macromolecules, a process called autophagy 2016 Pearson Education, Inc. Figure 4.12 Nucleus 1 mm Lysosome Digestive enzymes

Lysosome Plasma membrane Digestion Food vacuole 2016 Pearson Education, Inc. Figure 4.13 Vesicle containing two damaged organelles 1 mm Mitochondrion

fragment Peroxisome fragment Lysosome Peroxisome Vesicle 2016 Pearson Education, Inc. Mitochondrion Digestion Vacuoles: Diverse Maintenance Compartments Vacuoles are large vesicles derived from the endoplasmic reticulum and Golgi apparatus

The solution inside a vacuole differs in composition from the cytosol 2016 Pearson Education, Inc. Food vacuoles are formed by phagocytosis Contractile vacuoles, found in many freshwater protists, pump excess water out of cells Central vacuoles, found in many mature plant cells, hold organic compounds and water Certain vacuoles in plants and fungi carry out enzymatic hydrolysis like lysosomes 2016 Pearson Education, Inc. Figure 4.14

Central vacuole Cytosol Nucleus Central vacuole Cell wall Chloroplast 5 mm 2016 Pearson Education, Inc. The Endomembrane System: A Review The endomembrane system is a complex and

dynamic player in the cells compartmental organization 2016 Pearson Education, Inc. **Figure 4.15 Nucleus Rough ER Smooth ER cis Golgi trans Golgi 2016 Pearson Education, Inc. Plasma

membrane Concept 4.5: Mitochondria and chloroplasts change energy from one form to another Mitochondria are the sites of cellular respiration, a metabolic process that uses oxygen to generate ATP Chloroplasts, found in plants and algae, are the sites of photosynthesis Peroxisomes are oxidative organelles 2016 Pearson Education, Inc. The Evolutionary Origins of Mitochondria and Chloroplasts Mitochondria and chloroplasts display similarities with bacteria

Enveloped by a double membrane Contain ribosomes and multiple circular DNA molecules Grow and reproduce somewhat independently in cells 2016 Pearson Education, Inc. The endosymbiont theory An early ancestor of eukaryotic cells engulfed a nonphotosynthetic prokaryotic cell, which formed an endosymbiont relationship with its host The host cell and endosymbiont merged into a single organism, a eukaryotic cell with a mitochondrion At least one of these cells may have taken up a photosynthetic prokaryote, becoming the ancestor of cells that contain chloroplasts

2016 Pearson Education, Inc. Figure 4.16 Endoplasmic reticulum Nucleus Nuclear envelope Engulfing of oxygenusing nonphotosynthetic prokaryote, which, becomes a mitochondrion

Ancestor of eukaryotic cells (host cell) Engulfing of photosynthetic prokaryote Mitochondrion Mitochondrion Chloroplast At least one cell Photosynthetic eukaryote 2016 Pearson Education, Inc. Nonphotosynthetic eukaryote

Mitochondria: Chemical Energy Conversion Mitochondria are in nearly all eukaryotic cells They have a smooth outer membrane and an inner membrane folded into cristae The inner membrane creates two compartments: intermembrane space and mitochondrial matrix Some metabolic steps of cellular respiration are catalyzed in the mitochondrial matrix Cristae present a large surface area for enzymes that synthesize ATP 2016 Pearson Education, Inc. Figure 4.17 Mitochondrion

10 mm Intermembrane space Mitochondria Outer membrane DNA Free ribosomes in the mitochondrial matrix Inner membrane

Cristae Matrix (a) Diagram and TEM of mitochondrion 2016 Pearson Education, Inc. Mitochondrial DNA 0.1 mm Nuclear DNA (b) Network of mitochondria in Euglena (LM) Chloroplasts: Capture of Light Energy Chloroplasts contain the green pigment chlorophyll,

as well as enzymes and other molecules that function in photosynthesis Chloroplasts are found in leaves and other green organs of plants and in algae 2016 Pearson Education, Inc. Chloroplast structure includes Thylakoids, membranous sacs, stacked to form a granum Stroma, the internal fluid Stomata, pores needed for gas exchange The chloroplast is one of a group of plant organelles called plastids 2016 Pearson Education, Inc.

Figure 4.18 Chloroplast Stroma Ribosomes DNA Thylakoid Intermembrane space (a) Diagram and TEM of chloroplast 2016 Pearson Education, Inc. 50 mm Inner and outer membranes

Granum 1 mm Chloroplasts (red) (b) Chloroplasts in an algal cell Figure 4.18-1 Stroma Ribosomes Inner and outer membranes Granum

DNA Thylakoid Intermembrane space (a) Diagram and TEM of chloroplast 2016 Pearson Education, Inc. 1 mm Peroxisomes: Oxidation Peroxisomes are specialized metabolic compartments bounded by a single membrane Peroxisomes produce hydrogen peroxide and then convert it to water

Peroxisomes perform reactions with many different functions 2016 Pearson Education, Inc. Concept 4.6: The cytoskeleton is a network of fibers that organizes structures and activities in the cell The cytoskeleton is a network of fibers extending throughout the cytoplasm It organizes the cells structures and activities 2016 Pearson Education, Inc. 10 mm Figure 4.20

2016 Pearson Education, Inc. Roles of the Cytoskeleton: Support and Motility The cytoskeleton helps to support the cell and maintain its shape It provides anchorage for many organelles and molecules It interacts with motor proteins to produce motility Inside the cell, vesicles and other organelles can walk along the tracks provided by the cytoskeleton 2016 Pearson Education, Inc. Figure 4.21 ATP

Vesicle Receptor for motor protein Motor protein Microtubule (ATP powered) of cytoskeleton (a) Motor proteins walk vesicles along cytoskeletal fibers. Microtubule Vesicles 0.25 mm (b) Two vesicles move toward the tip of a nerve cell extension called an axon (SEM) 2016 Pearson Education, Inc. Components of the Cytoskeleton

Three main types of fibers make up the cytoskeleton Microtubules are the thickest of the three components of the cytoskeleton Microfilaments, also called actin filaments, are the thinnest components Intermediate filaments are fibers with diameters in a middle range 2016 Pearson Education, Inc. Figure 4.T01 2016 Pearson Education, Inc. Microtubules Microtubules are hollow rods constructed from globular protein dimers called tubulin

Functions of microtubules Shape and support the cell Guide movement of organelles Separate chromosomes during cell division 2016 Pearson Education, Inc. Centrosomes and Centrioles In animal cells, microtubules grow out from a centrosome near the nucleus The centrosome is a microtubule-organizing center The centrosome has a pair of centrioles, each with nine triplets of microtubules arranged in a ring 2016 Pearson Education, Inc.

Video: Flagellum Microtubule 2016 Pearson Education, Inc. Cilia and Flagella Microtubules control the beating of cilia and flagella, microtubule-containing extensions projecting from some cells Flagella are limited to one or a few per cell, while cilia occur in large numbers on cell surfaces Cilia and flagella also differ in their beating patterns 2016 Pearson Education, Inc. Cilia and flagella share a common structure A group of microtubules sheathed by the plasma membrane

A basal body that anchors the cilium or flagellum A motor protein called dynein, which drives the bending movements of a cilium or flagellum 2016 Pearson Education, Inc. Figure 4.23 Outer microtubule doublet Motor proteins (dyneins) Central microtubule 0.1 mm

Radial spoke Microtubules Plasma membrane Basal body 0.5 mm Cross-linking (b) Cross section protein between of motile cilium outer doublets 0.1 mm Triplet (a) Longitudinal section

of motile cilium (c) Cross section of basal body 2016 Pearson Education, Inc. Plasma membrane How dynein walking moves flagella and cilia Dynein arms alternately contact, move, and release the outer microtubules The outer doublets and central microtubules are held together by flexible cross-linking proteins Movements of the doublet arms cause the cilium or flagellum to bend

2016 Pearson Education, Inc. Microfilaments (Actin Filaments) Microfilaments are thin solid rods, built from molecules of globular actin subunits The structural role of microfilaments is to bear tension, resisting pulling forces within the cell Bundles of microfilaments make up the core of microvilli of intestinal cells 2016 Pearson Education, Inc. Microfilaments that function in cellular motility interact with the motor protein myosin For example, actin and myosin interact to cause muscle contraction, amoeboid movement of white blood cells, and cytoplasmic streaming in plant cells

2016 Pearson Education, Inc. Intermediate Filaments Intermediate filaments are larger than microfilaments but smaller than microtubules Intermediate filaments are only found in the cells of some animals, including vertebrates They support cell shape and fix organelles in place Intermediate filaments are more permanent cytoskeleton elements than the other two classes 2016 Pearson Education, Inc. Concept 4.7: Extracellular components and connections between cells help coordinate cellular activities

Most cells synthesize and secrete materials that are external to the plasma membrane These extracellular materials are involved in many cellular functions 2016 Pearson Education, Inc. Cell Walls of Plants The cell wall is an extracellular structure that distinguishes plant cells from animal cells The cell wall protects the plant cell, maintains its shape, and prevents excessive uptake of water Plant cell walls are made of cellulose fibers embedded in other polysaccharides and protein 2016 Pearson Education, Inc.

Plant cell walls may have multiple layers Primary cell wall: relatively thin and flexible Middle lamella: thin layer between primary walls of adjacent cells Secondary cell wall (in some cells): added between the plasma membrane and the primary cell wall Plasmodesmata are channels between adjacent plant cells 2016 Pearson Education, Inc. Figure 4.25 Central vacuole Cytosol Plasma membrane

Plant cell walls Plasmodesmata Secondary cell wall Primary cell wall Middle lamella 1 mm 2016 Pearson Education, Inc. The Extracellular Matrix (ECM) of Animal Cells Animal cells lack cell walls but are covered by an elaborate extracellular matrix (ECM)

The ECM is made up of glycoproteins such as collagen, proteoglycans, and fibronectin ECM proteins bind to receptor proteins in the plasma membrane called integrins 2016 Pearson Education, Inc. **Figure 4.26-1 Collagen A proteoglycan complex: EXTRACELLULAR FLUID Fibronectin

Integrins Plasma membrane Microfilaments 2016 Pearson Education, Inc. CYTOPLASM Figure 4.26-2 Polysaccharide molecule Carbohydrates

Core protein Proteoglycan molecule 2016 Pearson Education, Inc. Cell Junctions Neighboring cells in an animal or plant often adhere, interact, and communicate through direct physical contact There are several types of intercellular junctions that facilitate this

Plasmodesmata Tight junctions Desmosomes Gap junctions 2016 Pearson Education, Inc. Plasmodesmata in Plant Cells Plasmodesmata are channels that perforate plant cell walls Through plasmodesmata, water and small solutes (and sometimes proteins and RNA) can pass from cell to cell 2016 Pearson Education, Inc.

Tight Junctions, Desmosomes, and Gap Junctions in Animal Cells Animal cells have three main types of cell junctions Tight junctions Desmosomes Gap junctions All are especially common in epithelial tissue 2016 Pearson Education, Inc. Figure 4.27 Tight junctions prevent fluid from moving across a layer of cells.

Tight junction TEM 0.5 mm Tight junction Intermediate filaments Desmosome Gap junction TEM 1 mm

Plasma membranes of adjacent cells 2016 Pearson Education, Inc. Space between cells Extracellular matrix TEM Ions or small molecules 0.1 mm

Figure 4.27-1 Tight junctions prevent fluid from moving across a layer of cells. Tight junction Intermediate filaments Desmosome Gap junction

Ions or small molecules Plasma membranes of adjacent cells 2016 Pearson Education, Inc. Space between cells Extracellular matrix The Cell: A Living Unit Greater Than the Sum of Its Parts None of the components of a cell work alone

For example, a macrophages ability to destroy bacteria involves the whole cell, coordinating components such as the cytoskeleton, lysosomes, and plasma membrane Cellular functions arise from cellular order 2016 Pearson Education, Inc. Figure 4.UN01-1 Mature parent cell Budding cell 1 mm

2016 Pearson Education, Inc. **Figure 4.UN03 Cell Component Nucleus Structure Function Surrounded by nuclear envelope (double membrane) perforated by nuclear pores; nuclear envelope continuous with endoplasmic reticulum (ER)

Houses chromosomes, which are made of chromatin (DNA and proteins); contains nucleoli, where ribosomal subunits are made; pores regulate entry and exit of materials Two subunits made of ribosomal RNA and proteins; can be free in cytosol or bound to ER Protein synthesis (ER)

Ribosome 2016 Pearson Education, Inc. **Figure 4.UN04 Cell Component Structure Extensive network of Endoplasmic reticulum membrane-bounded tubules (Nuclear envelope) and sacs; membrane separates lumen from cytosol; continuous with

nuclear envelope Function Smooth ER: synthesis of lipids, metabolism of carbohydrates, Ca2+ storage, detoxification of drugs and poisons Rough ER: aids in synthesis of secretory and other proteins from bound ribosomes; adds carbohydrates to proteins to make glycoproteins; produces new membrane Golgi apparatus

Stacks of flattened membranous sacs; has polarity (cis and trans faces) Modification of proteins, carbohydrates on proteins, and phospholipids; synthesis of many polysaccharides; sorting of Golgi products, which are then released in vesicles Lysosome Membranous sac of hydrolytic

enzymes (in animal cells) Breakdown of ingested substances, cell macromolecules, and damaged organelles for recycling Digestion, storage, waste disposal, water balance, plant cell growth and protection Vacuole 2016 Pearson Education, Inc. Large membrane-bounded vesicle

**Figure 4.UN05 Cell Component Structure Function Mitochondrion Bounded by double membrane; inner membrane has infoldings (cristae) Cellular respiration

Chloroplast Typically two membranes around fluid stroma, which contains thylakoids stacked into grana (in cells of photosynthetic eukaryotes, including plants) Photosynthesis Peroxisome Specialized metabolic compartment bounded by a single membrane Contains enzymes that transfer hydrogen atoms from certain

molecules to oxygen, producing hydrogen peroxide (H2O2) as a by-product; H2O2 is converted to water by another enzyme 2016 Pearson Education, Inc.

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