What are the key features of eukaryotic cells? Unlike prokaryotic cells, eukaryotic cells have:
- A membrane-bound nucleus, a central cavity surrounded by membrane that houses the cell’s genetic material.
- A number of membrane-bound organelles, compartments with specialized functions that float in the cytosol. (Organelle means “little organ,” and this name reflects that the organelles, like the organs of our body, have unique functions as part of a larger system.)
- Multiple linear chromosomes, as opposed to the single circular chromosome of a prokaryote.
Eukaryotic cells are much more complicated than those of prokaryotes. They are packed with a fascinating array of subcellular structures that play important roles in energy balance, metabolism, and gene expression.
In the articles and videos that follow, we’ll take a tour through eukaryotic plant and animal cells, exploring the unique structures they contain and the role that each structure plays in the life of the cell.
Already know what part of the cell you want to visit? Use the list below to jump to your region of interest:
- Plasma membrane and cytoplasm
- Nucleus and ribosomes
- Endomembrane system
- Mitochondria and chloroplasts
- Extracellular matrix and cell wall
- Cell junctions
Diagram of a typical animal cell:
Diagram of an animal cell with components lettered.
Diagram of a typical plant cell:
Konstantin Mereschkowski proposed a symbiotic origin for cells with nuclei
The concept of the eukaryote has been attributed to the French biologist Edouard Chatton (1883–1947). The terms prokaryote and eukaryote were more definitively reintroduced by the Canadian microbiologist Roger Stanier and the Dutch-American microbiologist C. B. van Niel in 1962. In his 1937 work Titres et Travaux Scientifiques, Chatton had proposed the two terms, calling the bacteria prokaryotes and organisms with nuclei in their cells eukaryotes. However he mentioned this in only one paragraph, and the idea was effectively ignored until Chatton’s statement was rediscovered by Stanier and van Niel.
In 1905 and 1910, the Russian biologist Konstantin Mereschkowski (1855–1921) argued that plastids were reduced cyanobacteria in a symbiosis with a non-photosynthetic (heterotrophic) host that was itself formed by symbiosis between an amoeba-like host and a bacterium-like cell that formed the nucleus. Plants had thus inherited photosynthesis from cyanobacteria.
In 1967, Lynn Margulis provided microbiological evidence for endosymbiosis as the origin of chloroplasts and mitochondria in eukaryotic cells in her paper, On the origin of mitosing cells. In the 1970s, Carl Woese explored microbial phylogenetics, studying variations in 16S ribosomal RNA. This helped to uncover the origin of the eukaryotes and the symbiogenesis of two important eukaryote organelles, mitochondria and chloroplasts. In 1977, Woese and George Fox introduced a “third form of life”, which they called the Archaebacteria; in 1990, Woese, Otto Kandler and Mark L. Wheelis renamed this the Archaea.
In 1979, G. W. Gould and G. J. Dring suggested that the eukaryotic cell’s nucleus came from the ability of Gram-positive bacteria to form endospores. In 1987 and later papers, Thomas Cavalier-Smith proposed instead that the membranes of the nucleus and endoplasmic reticulum first formed by infolding a prokaryote’s plasma membrane. In the 1990s, several other biologists proposed endosymbiotic origins for the nucleus, effectively reviving Mereschkowski’s theory.
Eukaryotic cells are typically much larger than those of prokaryotes having a volume of around 10,000 times greater than the prokaryotic cell. They have a variety of internal membrane-bound structures, called organelles, and a cytoskeleton composed of microtubules, microfilaments, and intermediate filaments, which play an important role in defining the cell’s organization and shape. Eukaryotic DNA is divided into several linear bundles called chromosomes, which are separated by a microtubular spindle during nuclear division.
The endomembrane system and its components
Eukaryote cells include a variety of membrane-bound structures, collectively referred to as the endomembrane system. Simple compartments, called vesicles and vacuoles, can form by budding off other membranes. Many cells ingest food and other materials through a process of endocytosis, where the outer membrane invaginates and then pinches off to form a vesicle. It is probable that most other membrane-bound organelles are ultimately derived from such vesicles. Alternatively some products produced by the cell can leave in a vesicle through exocytosis.
The nucleus is surrounded by a double membrane (commonly referred to as a nuclear membrane or nuclear envelope), with pores that allow material to move in and out. Various tube- and sheet-like extensions of the nuclear membrane form the endoplasmic reticulum, which is involved in protein transport and maturation. It includes the rough endoplasmic reticulum where ribosomes are attached to synthesize proteins, which enter the interior space or lumen. Subsequently, they generally enter vesicles, which bud off from the smooth endoplasmic reticulum. In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles (cisternae), the Golgi apparatus.
Vesicles may be specialized for various purposes. For instance, lysosomes contain digestive enzymes that break down most biomolecules in the cytoplasm. Peroxisomes are used to break down peroxide, which is otherwise toxic. Many protozoans have contractile vacuoles, which collect and expel excess water, and extrusomes, which expel material used to deflect predators or capture prey. In higher plants, most of a cell’s volume is taken up by a central vacuole, which mostly contains water and primarily maintains its osmotic pressure.
Mitochondria and plastids:
Simplified structure of a mitochondrion
Mitochondria are organelles found in all but one eukaryote. Mitochondria provide energy to the eukaryote cell by converting sugars into ATP. They have two surrounding membranes, each a phospholipid bi-layer; the inner of which is folded into invaginations called cristae where aerobic respiration takes place.
The outer mitochondrial membrane is freely permeable and allows almost anything to enter into the intermembrane space while the inner mitochondrial membrane is semi permeable so allows only some required things into the mitochondrial matrix.
Mitochondria contain their own DNA, which has close structural similarities to bacterial DNA, and which encodes rRNA and tRNA genes that produce RNA which is closer in structure to bacterial RNA than to eukaryote RNA. They are now generally held to have developed from endosymbiotic prokaryotes, probably proteobacteria.
Some eukaryotes, such as the metamonads such as Giardia and Trichomonas, and the amoebozoan Pelomyxa, appear to lack mitochondria, but all have been found to contain mitochondrion-derived organelles, such as hydrogenosomes and mitosomes, and thus have lost their mitochondria secondarily. They obtain energy by enzymatic action on nutrients absorbed from the environment. The metamonad Monocercomonoides has also acquired, by lateral gene transfer, a cytosolic sulfur mobilisation system which provides the clusters of iron and sulfur required for protein synthesis. The normal mitochondrial iron-sulfur cluster pathway has been lost secondarily.
Plants and various groups of algae also have plastids. Plastids also have their own DNA and are developed from endosymbionts, in this case cyanobacteria. They usually take the form of chloroplasts which, like cyanobacteria, contain chlorophyll and produce organic compounds (such as glucose) through photosynthesis. Others are involved in storing food. Although plastids probably had a single origin, not all plastid-containing groups are closely related. Instead, some eukaryotes have obtained them from others through secondary endosymbiosis or ingestion. The capture and sequestering of photosynthetic cells and chloroplasts occurs in many types of modern eukaryotic organisms and is known as kleptoplasty.
Endosymbiotic origins have also been proposed for the nucleus, and for eukaryotic flagella.
Longitudinal section through the flagellum of Chlamydomonas reinhardtii
Many eukaryotes have long slender motile cytoplasmic projections, called flagella, or similar structures called cilia. Flagella and cilia are sometimes referred to as undulipodia, and are variously involved in movement, feeding, and sensation. They are composed mainly of tubulin. These are entirely distinct from prokaryotic flagellae. They are supported by a bundle of microtubules arising from a centriole, characteristically arranged as nine doublets surrounding two singlets. Flagella also may have hairs, or mastigonemes, and scales connecting membranes and internal rods. Their interior is continuous with the cell’s cytoplasm.
Microfilamental structures composed of actin and actin binding proteins, e.g., α-actinin, fimbrin, filamin are present in submembraneous cortical layers and bundles, as well. Motor proteins of microtubules, e.g., dynein or kinesin and actin, e.g., myosins provide dynamic character of the network.
Centrioles are often present even in cells and groups that do not have flagella, but conifers and flowering plants have neither. They generally occur in groups that give rise to various microtubular roots. These form a primary component of the cytoskeletal structure, and are often assembled over the course of several cell divisions, with one flagellum retained from the parent and the other derived from it. Centrioles produce the spindle during nuclear division.
The significance of cytoskeletal structures is underlined in the determination of shape of the cells, as well as their being essential components of migratory responses like chemotaxis and chemokinesis. Some protists have various other microtubule-supported organelles. These include the radiolaria and heliozoa, which produce axopodia used in flotation or to capture prey, and the haptophytes, which have a peculiar flagellum-like organelle called the haptonema.
Eukaryotic Chromosome Structure:
The length of DNA in the nucleus is far greater than the size of the compartment in which it is contained. To fit into this compartment the DNA has to be condensed in some manner. The degree to which DNA is condensed is expressed as its packing ratio.
Packing ratio – the length of DNA divided by the length into which it is packaged
For example, the shortest human chromosome contains 4.6 x 107 bp of DNA (about 10 times the genome size of E. coli). This is equivalent to 14,000 µm of extended DNA. In its most condensed state during mitosis, the chromosome is about 2 µm long. This gives a packing ratio of 7000 (14,000/2).
To achieve the overall packing ratio, DNA is not packaged directly into final structure of chromatin. Instead, it contains several hierarchies of organization. The first level of packing is achieved by the winding of DNA around a protein core to produce a “bead-like” structure called a nucleosome. This gives a packing ratio of about 6. This structure is invariant in both the euchromatin and heterochromatin of all chromosomes. The second level of packing is the coiling of beads in a helical structure called the 30 nm fiber that is found in both interphase chromatin and mitotic chromosomes. This structure increases the packing ratio to about 40. The final packaging occurs when the fiber is organized in loops, scaffolds and domains that give a final packing ratio of about 1000 in interphase chromosomes and about 10,000 in mitotic chromosomes.
Eukaryotic chromosomes consist of a DNA-protein complex that is organized in a compact manner which permits the large amount of DNA to be stored in the nucleus of the cell. The subunit designation of the chromosome is chromatin. The fundamental unit of chromatin is the nucleosome.
Chromatin – the unit of analysis of the chromosome; chromatin reflects the general structure of the chromosome but is not unique to any particular chromosome
Nucleosome – simplest packaging structure of DNA that is found in all eukaryotic chromosomes; DNA is wrapped around an octamer of small basic proteins called histones; 146 bp is wrapped around the core and the remaining bases link to the next nucleosome; this structure causes negative supercoiling
The nucleosome consists of about 200 bp wrapped around a histone octamer that contains two copies of histone proteins H2A, H2B, H3 and H4. These are known as the core histones. Histones are basic proteins that have an affinity for DNA and are the most abundant proteins associated with DNA. The amino acid sequence of these four histones is conserved suggesting a similar function for all.
The length of DNA that is associated with the nucleosome unit varies between species. But regardless of the size, two DNA components are involved. Core DNA is the DNA that is actually associated with the histone octamer. This value is invariant and is 146 base pairs. The core DNA forms two loops around the octamer, and this permits two regions that are 80 bp apart to be brought into close proximity. Thus, two sequences that are far apart can interact with the same regulatory protein to control gene expression. The DNA that is between each histone octamer is called the linker DNA and can vary in length from 8 to 114 base pairs. This variation is species specific, but variation in linker DNA length has also been associated with the developmental stage of the organism or specific regions of the genome.
The next level of organization of the chromatin is the 30 nm fiber. This appears to be a solenoid structure with about 6 nucleosomes per turn. This gives a packing ratio of 40, which means that every 1 µm along the axis contains 40 µm of DNA. The stability of this structure requires the presence of the last member of the histone gene family, histone H1. Because experiments that strip H1 from chromatin maintain the nucleosome, but not the 30 nm structure, it was concluded that H1 is important for the stabilization of the 30 nm structure.
The final level of packaging is characterized by the 700 nm structure seen in the metaphase chromosome. The condensed piece of chromatin has a characteristic scaffolding structure that can be detected in metaphase chromosomes. This appears to be the result of extensive looping of the DNA in the chromosome.
The last definitions that need to be presented are euchromatin and heterochromatin. When chromosomes are stained with dyes, they appear to have alternating lightly and darkly stained regions. The lightly-stained regions are euchromatin and contain single-copy, genetically-active DNA. The darkly-stained regions are heterochromatin and contain repetitive sequences that are genetically