Around 3.5 Billion B.C.T., the first primitive life forms emerged on Earth. The fossil record contains prokaryotes (cells without nuclei) from up to 3.5 billion years ago, and eukaryotes (cells with nuclei) from up to 2.7 billion years ago.
The prokaryotes are a group of organisms whose cells lack a cell nucleus (karyon), or any other membrane-bound organelles. The organisms whose cells do have a nucleus are called eukaryotes. Most prokaryotes are unicellular organisms, although a few such as myxobacteria have multi-cellular stages in their life cycles or create large colonies like cyanobacteria. The word prokaryote comes from the Greek (pro-) "before" + (karyon) "nut or kernel".
Prokaryotes do not have a nucleus, mitochondria, or any other membrane-bound organelles. In other words, all their intra-cellular water-soluble components (proteins, DNA and metabolites) are located together in the same area enclosed by cell membrane, rather than separated in different cellular compartments.
The division to prokaryotes and eukaryotes reflects two distinct levels of cellular organization rather than biological classification of species. Prokaryotes include two major classification domains: the bacteria and the archaea. Archaea were recognized as a domain of life in 1990. These organisms were originally thought to live only in inhospitable conditions such as extremes of temperature, pH, and radiation but have since been found in all types of habitats.
The division to prokaryotes and eukaryotes is usually considered the most important distinction among organisms. The distinction is that eukaryotic cells have a "true" nucleus containing their DNA, whereas prokaryotic cells do not have a nucleus. One criticism of this classification points out that the word "prokaryote" is based on what these organisms are not (they are not eukaryotic), rather than what they are (either archaea or bacteria). Another difference is that ribosomes in prokaryotes are smaller than in eukaryotes. However, two organelles found in many eukaryotic cells, mitochondria and chloroplasts, contain ribosomes similar in size and makeup to those found in prokaryotes. This is one of many pieces of evidence that mitochondria and chloroplasts are themselves descended from free-living bacteria.
The genome in a prokaryote is held within a DNA/protein complex in the cytosol called the nucleoid, which lacks a nuclear envelope. The complex contains a single, cyclic, double-stranded molecule of stable chromosomal DNA, in contrast to the multiple linear, compact, highly organized chromosomes found in eukaryotic cells. In addition, many important genes of prokaryotes are stored in separate circular DNA structures called plasmids.
Prokaryotes lack distinct mitochondria and chloroplasts. Instead, processes such as oxidative phosphorylation and photosynthesis take place across the prokaryotic cell membrane. However, prokaryotes do possess some internal structures, such as prokaryotic cytoskeletons, and the bacterial order Planctomycetes have a membrane around their nucleoid and contain other membrane-bound cellular structures. Both eukaryotes and prokaryotes contain large RNA/protein structures called ribosomes, which produce protein.
Prokaryotic cells are usually much smaller than eukaryotic cells. Therefore, prokaryotes have a larger surface-area-to-volume ratio, giving them a higher metabolic rate, a higher growth rate, and, as a consequently, a shorter generation time than eukaryotes.
In 1977, Carl Woese proposed dividing prokaryotes into the bacteria and archaea (originally eubacteria and archaebacteria) because of the major differences in the structure and genetics between the two groups of organisms. This arrangement of eukaryota (also called "eukarya"), bacteria, and archaea is called the three-domain system, replacing the traditional two-empire system.
The oldest known fossilized prokaryotes were laid down approximately 3.5 billion years ago, only about 1 billion years after the formation of the Earth's crust. Even today, prokaryotes are perhaps the most successful and abundant life-forms. Eukaryotes only appear in the fossil record later, and may have formed from endosymbiosis of multiple prokaryote ancestors. The oldest known fossil eukaryotes are about 1.7 billion years old. However, some genetic evidence suggests eukaryotes appeared as early as 3 billion years ago.
While Earth is the only place in the universe where life is known to exist, some have suggested that there is evidence on Mars of fossil or living prokaryotes; but this possibility remains the subject of considerable debate and skepticism.
Prokaryotes have diversified greatly throughout their long existence. The metabolism of prokaryotes is far more varied than that of eukaryotes, leading to many highly distinct prokaryotic types. For example, in addition to using photosynthesis or organic compounds for energy, as eukaryotes do, prokaryotes may obtain energy from inorganic compounds such as hydrogen sulfide. This enables prokaryotes to thrive in harsh environments as cold as the snow surface of Antarctica, and as hot as undersea hydrothermal vents and land-based hot springs.
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A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear envelope, within which the genetic material is carried. The presence of a nucleus gives eukaryotes their name, which comes from the Greek eu ("good") and karyon ("nut" or "kernel"). Most eukaryotic cells also contain other membrane-bound organelles such as mitochondria, chloroplasts and the Golgi apparatus. All large complex organisms are eukaryotes, including animals, plants and fungi. The group also includes many unicellular organisms.
Cell division in eukaryotes is different from that in organisms without a nucleus (Prokaryote). It involves separating the duplicated chromosomes, through movements directed by microtubules. There are two types of division processes. In mitosis, one cell divides to produce two genetically identical cells. In meiosis, which is required in sexual reproduction, one diploid cell (a cell having two instances of each chromosome, one from each parent) undergoes recombination of each pair of parental chromosomes, and then two stages of cell division, resulting in four haploid cells (gametes). Each gamete has just one complement of chromosomes, each a unique mix of the corresponding pair of parental chromosomes.
Eukaryotes appear to be monophyletic, and thus make up one of the three domains of life. The two other domains, bacteria and archaea, are prokaryotes and have none of the above features. Eukaryotes represent a tiny minority of all living things; even in a human body there are 10 times more microbes than human cells. However, due to their much larger size their collective worldwide biomass is estimated at about equal to that of prokaryotes.
The origin of the eukaryotic cell is considered a milestone in the evolution of life, since they include all complex cells and almost all multicellular organisms. The timing of this series of events is hard to determine. Some acritarchs are known from at least 1.65 billion years ago, and the possible algae Grypania has been found as far back as 2.1 billion years ago.
Organized living structures have been found in black shales of the Palaeoproterozoic Francevillian B Formation in Gabon, dated at 2.1 billion years old. Eukaryotic life could have evolved at that time. Fossils that are clearly related to modern groups start appearing an estimated 1.2 billion years ago, in the form of a red algae, though recent work suggests the existence of fossilized filamentous algae in the Vindhya basin dating back perhaps to 1.6 to 1.7 billion years ago.
Biomarkers suggest that at least stem eukaryotes arose even earlier. The presence of steranes in Australian shales indicates that eukaryotes were present in these rocks dated at 2.7 billion years old.
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