Eukaryotes

The Endosymbiont Theory (Lynn Margulis, 1981)

Provides an explanation for the evolution of the mitochondrion and chloroplast (from symbiotically living bacteria) Does not explain the evolution of other organelles, structures, and processes

Diversity of Eukaryotes
Eukaryotes that are neither plants, animals, or fungi are called protists, or microbial eukaryotes (though not all are microbial). They do not constitute a clade, they are paraphyletic. Phylogeny of microbial eukaryotes is the subject of much research. Electron microscopy and gene sequencing are revealing new information. Most eukaryotes can be divided into five groups: o o o o o Chromalveolates Plantae Excavates Rhizaria Unikonts

Endosymbiosis
Endosymbiosis: in which one organism lives inside another (is common in microbial eukaryotes) Dinoflagellates are common endosymbionts in animals and other microbial eukaryotes; some are photosynthetic. Some dinoflagellates live as endosymbionts in corals.

Evolution of Eukaryotes
Eukaryotic cells arose as the environment was changing dramatically from anaerobic to aerobic. Events that occurred in the evolution of eukaryote cells are still conjectural. The main events: o Origin of a flexible cell surface o Origin of a cytoskeleton o Origin of a nuclear envelope o Appearance of digestive vesicles or vacuoles o Endosymbiotic acquisition of some organelles Flexible cell surface: Prokaryotic cell wall was lost; cells can grow larger. As cell size increases surface area-to-volume ratio decreases, but with a flexible surface, infolding can occur, creating more surface area. A cytoskeleton provided cell support, allowed cells to change shape, and move materials around the cell, including daughter chromosomes. In some cells microtubules gave rise to flagella. The nuclear envelope may have developed from the plasma membrane. The DNA of a prokaryote is attached to the plasma membrane; infolding of the membrane could have been the first step in development of the nucleus.

The next step was probably phagocytosis the ability to engulf and digest other cells. The first true eukaryotes had a cytoskeleton and nuclear envelope; they probably had ER, Golgi apparatus, and perhaps flagella Cyanobacteria were producing oxygen; at some point, some Eukarya incorporated proteobacteria that evolved into mitochondria the endosymbiotic theory. The function of mitochondria initially might have been to detoxify O2 by reducing it to water. Later this became associated with ATP production. Some eukaryotes incorporated a prokaryote related to todays cyanobacteria, which developed into chloroplasts. Evolution of chloroplasts probably occurred in a series of endosymbiotic events Evidence comes from nucleic acid sequencing and electron microscopy.

Primary endosymbiosis: All chloroplasts descended from a gramnegative cyanobacterium (blue/green algae, blue/green bacteria) with an inner and outer membrane. A small amount of peptidoglycan from the bacterial cell wall is found today in the glaucophytes the first group to ranch off. Primary endosymbiosis gave rise to chloroplasts of green algae (chlorophytes and charophytes) and the red algae. Photosynthetic land plants arose from a green algal ancestor Red algal chloroplasts retain some pigments that were present in the original cyanobacterium. Secondary and tertiary endosymbiosis gave rise to chloroplasts in the other microbial eukaryote groups. The uglenid ancestor engulfed a chlorophyte, retaining the chloroplasts. Euglenid chloroplasts have the same pigments as green algae and land plants, and have a third membrane.

Uncertainties remain about the origins of eukaryotic cells. Laterial gene transfer complicates the study of relationships.

Endosymbiosis does not account for all bacterial genes in eukaryotes. A recent suggestion is that Eukarya arose form the fusion of a gram-negative bacterium and an archaean.

Diversity of Microbial Eukaryotes

Microbial eukaryotes have evolved a diversity of lifestyles. Most are aquatic, marine, and freshwater; but also damp soils and decaying organic matter. Some are photosynthetic, some are heterotrophs, some can switch between modes (heterotrophic & autotrophic).

Diversity of the Eukaryotes

Some used to be considered animals, and are called protozoans. But this term lumps phylogenetically unrelated groups. Most protozoans are ingestive heterotrophs. The term algae also lumps many groups of photosynthetic microbial eukaryotes and does not reflect phylogeny.

Eukaryote: Locomotion
Amoeboid motion cells form pseudopods that are extensions of the cell. A network of cytoskeletal microfilaments squeezes the cytoplasm forward. Cilia and flagella developed from microtubules. Cilia beat in a coordinated fashion; move cell forward or backward. Flagella have whip-like movement. Some pull, some push the cell forward. Flagella have a 9+2 arrangement of microtubules.

Eukaryote: Characteristics

Vacuoles increase effective surface area in large cells. Contractile vacuoles in freshwater microbial eukaryotes such as Paramecium are used to excrete excess water.

Food vacuoles are formed by Paramecium and others when solid food particles are ingested by endocytosis. The food is digested in the vacuole. Small vesicles pinch off increasing surface area for products of digestion to be absorbed by the rest of the cell.

Eukaryote: Reproduction
Most microbial eukaryotes have both sexual and asexual reproduction. Asexual processes o Binary fission equal splitting; mitosis followed by cytokinesis. o Multiple fission splitting into more than two cells.

o Budding outgrowth of a new cell from the surface of an old cell. o Spores specialized cells that are capable of growing into a new individual. The ciliate clade (such as Paramecium) have a single macronucleus and one to several micronuclei. The macronucleus contains many copies of the genetic information, packaged into units; regulates the life of the cell. Micronuclei are essential for genetic recombination.

In alternation of generations, a diploid spore-forming organism gives rise a haploid gamete-forming organism. When haploid gametes fuse (fertilization or syngamy) a diploid individual is formed. The haploid or diploid organism, or both, may reproduce asexually.

Eukaryotes: Alveolates
Alveolates: synapomorphy that distinguishes this clade is presence of alveoli or sacs beneath surface of plasma membrane. All unicellular; includes dinoflagellates, apicomplexans, and ciliates. Most dinoflagellates are marine and are important primary producers. Mixture of pigments give them a golden brown colour. Some are endosymbionts in corals and other invertebrates and microbial eukaryotes. Some are nonphotosynthetic parasites. Ciliates have numerous cilia, the structure is identical to flagella. Most are heterotrophic; very diverse group. Have complex body form; two types of nuclei.