Eukaryote
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Eukaryotes (also spelled "eucaryotes") are organisms with complex cells, in which the genetic material is organized into membrane-bound nuclei. They include the animals, plants, and fungi, which are mostly multicellular, as well as the kingdom of the protists, many of which are unicellular. In contrast, other organisms such as bacteria lack nuclei and other complex cell structures, and are called prokaryotes. Eukaryotes are thought to share a common origin, and are often treated formally as a superkingdom, empire, or domain. The name comes from the Greek eus or true and karyon or nut, referring to the nucleus.
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Structure
Eukaryotic cells are generally much larger than prokaryotes, typically a thousand times by volume. Usually, they have a variety of internal membranes and structures, called organelles, and a cytoskeleton composed of microtubules and microfilaments, which plays an important role in defining the cell's organization. However, some eukaryotes (e.g. diplomonads, microsporidia) do not contain organelles. Eukaryotic DNA is divided into several bundles called chromosomes, which are separated by a microtubular spindle during nuclear division. In addition to asexual cell division, most eukaryotes have some process of sexual reproduction via cell fusion, which is not found among prokaryotes.
Internal membranes
Eukaryotic cells include a variety of membrane-bound structures, collectively referred to as the endomembrane system. Simple compartments, called vesicles or 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.
The nucleus is surrounded by a double membrane, with pores that allow material to move in and out. Various tube- and sheet-like extensions of the nuclear membrane form what is called the endoplasmic reticulum or ER, which is involved in protein transport. It includes rough sections where ribosomes are attached, and the proteins they synthesize enter the interior space or lumen. Subsequently, they generally enter vesicles, which bud off from the smooth section. In most eukaryotes, the proteins may be further modified in stacks of flattened vesicles, called Golgi bodies or dictyosomes.
Vesicles may be specialized for various purposes. For instance, lysosomes contain enzymes that break down the contents of food vacuoles, and peroxisomes are used to break down peroxide which is toxic otherwise. Many protozoa have contractile vacuoles, which collect and expel excess water, and extrusomes, which expel material used to deflect predators or capture prey. In multicellular organisms, hormones are often produced in vesicles. In higher plants, most of a cell's volume is taken up by a central vacuole or tonoplast, which maintains its osmotic pressure.
Mitochondria and plastids
Mitochondria are organelles found in nearly all eukaryotes. They are surrounded by double membranes, the inner of which is folded into invaginations called cristae, where aerobic respiration takes place. They contain their own DNA and are only formed by the fission of other mitochondria. They are now generally held to have developed from endosymbiotic prokaryotes, probably proteobacteria. The few protozoa that lack mitochondria have been found to contain mitochondrion-derived organelles, such as hydrogenosomes and mitosomes.
Plants and various groups of algae also have plastids. Again, these have their own DNA and developed from endosymbiotes, in this case cyanobacteria. They usually take the form of chloroplasts, which like cyanobacteria contain chlorophyll and produce energy through photosynthesis. Others are involved in storing food. Although plastids likely 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.
Endosymbiotic origins have also been proposed for the nucleus and eukaryotic flagella, but this is not generally accepted, both from a lack of cytological evidence and difficulty in reconciling this with cellular reproduction.
Cytoskeletal structures
Many eukaryotes have slender motile projections, usually called flagella when long and cilia when short, that are variously involved in movement, feeding, and sensation. These are entirely distinct from prokaryotic flagella. They are supported by a bundle of microtubules arising from a basal body, also called a kinetosome or centriole, characteristically arranged as nine doublets surrounding two singlets. Flagella also may have hairs or mastigonemes, scales, connecting membranes and internal rods. Their interior is continuous with the cell's cytoplasm.
Centrioles are often present even in cells and groups that do not have flagella. They generally occur in groups of one or two, called kinetids, 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 may also be associated in the formation of a spindle during nuclear division.
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.
Reproduction
Nuclear division is often coordinated with cell division. This generally takes place by mitosis, a process which allows each daughter nucleus to receive one copy of each chromosome. In most eukaryotes there is also a process of sexual reproduction, typically involving an alternation between haploid generations, where only one copy of each chromosome is present, and diploid generations, where two are present, occurring through nuclear fusion (syngamy) and meiosis. There is considerable variation in this pattern, however.
Origin and evolution
The origin of the eukaryotic cell was a milestone in the evolution of life. Although eukaryotes use the same genetic code and metabolic processes as prokaryotes, their higher level of organizational complexity has permitted the development of truly multicellular organisms. Without eukaryotes, the world would lack mammals, birds, fish, invertebrates, mushrooms, plants, and complex single-celled organisms.
Eukaryotes are generally held to have originated around 2 billion years ago, but this has been disputed. Early fossils such as acritarchs are difficult to interpret. Forms that can be related to modern groups start appearing around 800 million years ago, and most fossil lines are known by the end of the Cambrian, around 500 million years ago.
Genetic studies during the 1980s and 1990s left most eukaryotes in an unresolved "crown" group, usually divided by the form of the cristae in their mitochondria. The few amitochondriate groups branched first and so were considered basal. However, newer studies contradict this, and a new picture has been slow to develop. Most eukaryotes may be assigned to several supergroups:
| Opisthokonts | Animals, fungi, choanoflagellates, etc. |
| Amoebozoa | Most lobose amoebae and slime moulds |
| Rhizaria | Various amoeboid protozoa |
| Excavates | Various flagellate protozoa |
| Plantae sensu lato | Land plants, green and red algae |
| Chromista | Brown algae, diatoms, water moulds, etc. |
| Alveolates | Ciliates, Apicomplexa, dinoflagellates, etc. |
However, some protists are not closely related to any of these lines, and the relationships between the different supergroups remain almost entirely uncertain. In particular, there is dispute about where the root of the evolutionary tree belongs, and as a result what the earliest eukaryotes were like.