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Semiautonomous Organelles

We now turn to those organelles in eukaryotic cells that are considered semiautonomous: mitochondria and chloroplasts. These organelles can grow and divide to reproduce themselves, but they are not completely autonomous because they depend on other parts of the cell for their internal components (Figure 4.23). For example, most of the proteins found in mitochondria are imported from the cytosol. In this section, we will survey the structures and functions of the semiautonomous organelles in eukaryotic cells and consider their evolutionary origins. In Chapters 7 and 8, we will explore the functions of mitochondria and chloroplasts in greater depth.

Figure 4.23
Semiautonomous organelles.These are the mitochondria and chloroplasts.
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Mitochondria Supply Cells with Most of Their ATP

Mitochondrion (plural, mitochondria) literally means thread granule, which is what mitochondria look like under a light microscope. They are similar in size to bacteria. Depending on a cell's function, it may contain a few hundred to a few thousand mitochondria. Cells with particularly heavy energy demands, such as muscle cells, have more mitochondria than other cells. Research has shown that regular exercise increases the number and size of mitochondria in human muscle cells to meet the expanded demand for energy.

A mitochondrion has an outer membrane and an inner membrane separated by a region called the intermembrane space (Figure 4.24). The inner membrane is highly invaginated (folded) to form projections called cristae. These invaginations greatly increase the surface area of the inner membrane, which is the site where ATP is made. The compartment enclosed by the inner membrane is the mitochondrial matrix.

Figure 4.24
Structure of a mitochondrion.This organelle is enclosed in two membranes. The invaginations of the inner membrane are called cristae. The mitochondrial matrix lies inside the inner membrane. The micrograph is a colorized TEM.

Concept check: What is the advantage of having a highly invaginated inner membrane?


The primary role of mitochondria is to make ATP. Even though mitochondria produce most of a cell's ATP, mitochondria do not create energy. Rather, their primary function is to convert chemical energy that is stored within the covalent bonds of organic molecules into a form that can be readily used by cells. Covalent bonds in sugars, fats, and amino acids store a large amount of energy. The breakdown of these molecules into simpler molecules releases energy that is used to make ATP. Many proteins in living cells utilize ATP to carry out their functions, such as muscle contraction, uptake of nutrients, cell division, and many other cellular processes.

Mitochondria perform other functions as well. They are involved in the synthesis, modification, and breakdown of several types of cellular molecules. For example, the synthesis of certain hormones requires enzymes that are found in mitochondria. Another interesting role of mitochondria is to generate heat in specialized fat cells known as brown fat cells. Groups of brown fat cells serve as “heating pads” that help to revive hibernating animals and protect sensitive areas of young animals from the cold.

Chloroplasts Carry Out Photosynthesis

Chloroplasts are organelles that can capture light energy and use some of that energy to synthesize organic molecules such as glucose. This process, called photosynthesis, is described in Chapter 8. Chloroplasts are found in nearly all species of plants and algae. Figure 4.25 shows the structure of a typical chloroplast. Like the mitochondrion, a chloroplast contains an outer and inner membrane. An intermembrane space lies between these two membranes. A third system of membranes, the thylakoid membrane, forms many flattened, fluid-filled tubules that enclose a single, convoluted compartment. These tubules tend to stack on top of each other to form a structure called a granum (plural, grana). The stroma is the compartment of the chloroplast that is enclosed by the inner membrane but outside the thylakoid membrane. The thylakoid lumen is enclosed by the thylakoid membrane.

Figure 4.25
Structure of a chloroplast.Like a mitochondrion, a chloroplast is enclosed in a double membrane. In addition, it has an internal thylakoid membrane system that forms flattened compartments. These compartments stack on each other to form grana. The stroma is located inside the inner membrane but outside the thylakoid membrane. This micrograph is a colorized TEM.

Chloroplasts are a specialized version of plant organelles that are more generally known as plastids. All plastids are derived from unspecialized proplastids. The various types of plastids are distinguished by their synthetic abilities and the types of pigments they contain. Chloroplasts, which carry out photosynthesis, contain the green pigment chlorophyll. The abundant number of chloroplasts in the leaves of plants gives them their green color. Chromoplasts, a second type of plastid, function in synthesizing and storing the yellow, orange, and red pigments known as carotenoids. Chromoplasts give many fruits and flowers their colors. In autumn, the chromoplasts also give many leaves their yellow, orange, and red colors. A third type of plastid, leucoplasts, typically lacks pigment molecules. An amyloplast is a leucoplast that synthesizes and stores starch. Amyloplasts are common in underground structures such as roots and tubers.

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Mitochondria and Chloroplasts Contain Their Own Genetic Material and Divide by Binary Fission

To fully appreciate the structure and organization of mitochondria and chloroplasts, we also need to briefly examine their genetic properties. In 1951, Y. Chiba exposed plant cells to Feulgen, a DNA-specific dye, and discovered that the chloroplasts became stained. Based on this observation, he was the first to suggest that chloroplasts contain their own DNA. Researchers in the 1970s and 1980s isolated DNA from both chloroplasts and mitochondria. These studies revealed that the DNA of these organelles resembled smaller versions of bacterial chromosomes.

The chromosomes found in mitochondria and chloroplasts are referred to as the mitochondrial genome and chloroplast genome, respectively, and the chromosomes found in the nucleus of the cell constitute the nuclear genome. Like bacteria, the genomes of most mitochondria and chloroplasts are composed of a single circular chromosome. Compared to the nuclear genome, they are very small. For example, the amount of DNA in the human nuclear genome (about 3 billion base pairs) is about 200,000 times greater than the mitochondrial genome. In terms of genes, the human genome has approximately 20,000 to 25,000 different genes, whereas the human mitochondrial genome has only a few dozen. Chloroplast genomes tend to be larger than mitochondrial genomes, and they have a correspondingly greater number of genes. Depending on the particular species of plant or algae, a chloroplast genome is about 10 times larger than the mitochondrial genome of human cells.

Just as the genomes of mitochondria and chloroplasts resemble bacterial genomes, the production of new mitochondria and chloroplasts bears a striking resemblance to the division of bacterial cells. Like their bacterial counterparts, mitochondria and chloroplasts increase in number via binary fission, or splitting in two. Figure 4.26 illustrates the process for a mitochondrion. The mitochondrial genome, which is found in a region called the nucleoid, is duplicated, and the organelle divides into two separate organelles. Mitochondrial and chloroplast division are needed to maintain a full complement of these organelles when cell growth occurs following cell division. In addition, environmental conditions may influence the sizes and numbers of these organelles. For example, when plants are exposed to more sunlight, the number of chloroplasts in leaf cells increases.

Figure 4.26
Division of mitochondria by binary fission.
Mitochondria and Chloroplasts Are Derived from Ancient Symbiotic Relationships

The observation that mitochondria and chloroplasts contain their own genetic material may seem puzzling. Perhaps you might think that it would be simpler for a eukaryotic cell to have all of its genetic material in the nucleus. The distinct genomes of mitochondria and chloroplasts can be traced to their evolutionary origin, which involved an ancient symbiotic association.

A symbiotic relationship occurs when two different species live in direct contact with each other. Endosymbiosis describes a symbiotic relationship in which the smaller species—the symbiont—actually lives inside the larger species. In 1883, Andreas Schimper proposed that chloroplasts were descended from an endosymbiotic relationship between cyanobacteria (a bacterium capable of photosynthesis) and eukaryotic cells. In 1922, Ivan Wallin also hypothesized an endosymbiotic origin for mitochondria.

In spite of these interesting ideas, the question of endosymbiosis was largely ignored until the discovery that mitochondria and chloroplasts contain their own genetic material. In 1970, the issue of endosymbiosis as the origin of mitochondria and chloroplasts was revived by Lynn Margulis in her book Origin of Eukaryotic Cells. During the 1970s and 1980s, the advent of molecular genetic techniques allowed researchers to analyze genes from mitochondria, chloroplasts, bacteria, and eukaryotic nuclear genomes. Researchers discovered that genes in mitochondria and chloroplasts are very similar to bacterial genes. Likewise, mitochondria and chloroplasts are strikingly similar in size and shape to certain bacterial species. These observations provided strong support for the endosymbiosis theory, which proposes that mitochondria and chloroplasts originated from bacteria that took up residence within a primordial eukaryotic cell (Figure 4.27). Over the next 2 billion years, the characteristics of these intracellular bacterial cells gradually changed to those of a mitochondrion or chloroplast. A more in-depth discussion of the origin of eukaryotic cells is found in Chapter 22.

Figure 4.27
A simplified view of the endosymbiosis theory.(a) According to this concept, modern mitochondria were derived from purple bacteria, also called α-proteobacteria. Over the course of evolution, their characteristics changed into those found in mitochondria today. (b) A similar phenomenon occurred for chloroplasts, which were derived from cyanobacteria, a bacterium that is capable of photosynthesis.


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Concept check: Discuss the similarities and differences between modern bacteria and mitochondria.


Symbiosis occurs because the relationship is beneficial to one or both species. According to the endosymbiosis theory, this relationship provided eukaryotic cells with useful cellular characteristics. Chloroplasts, which were derived from cyanobacteria, have the ability to carry out photosynthesis. This benefits plant cells by giving them the ability to use the energy from sunlight. By comparison, mitochondria are thought to have been derived from a different type of bacteria known as purple bacteria or α-proteobacteria. In this case, the endosymbiotic relationship enabled eukaryotic cells to synthesize greater amounts of ATP. How the relationship would have been beneficial to a cyanobacterium or purple bacterium is less clear, though the cytosol of a eukaryotic cell may have provided a stable environment with an adequate supply of nutrients.

During the evolution of eukaryotic species, many genes that were originally found in the genome of the primordial purple bacteria and cyanobacteria have been transferred from the organelles to the nucleus. This has occurred many times throughout evolution, so modern mitochondria and chloroplasts have lost most of the genes that still exist in present-day purple bacteria and cyanobacteria. Some researchers speculate that the movement of genes into the nucleus makes it easier for the cell to control the structure, function, and division of mitochondria and chloroplasts. In modern cells, hundreds of different proteins that make up these organelles are encoded by genes that have been transferred to the nucleus. These proteins are made in the cytosol and then taken up into mitochondria or chloroplasts. We will discuss this topic next.

Section 4.5 Quiz
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