Alveolates: Dinoflagellates, Apicomplexians, and Ciliates
A large body of data supports that the alveolates are derived from a shared common ancestor. The alveolates are named for the presence of an alveolus, or membrane-enclosed sac, beneath the cell membrane. The exact function of the alveolus is unknown, but it may be involved in osmoregulation. The alveolates are further categorized into some of the better-known protists: the dinoflagellates, the apicomplexans, and the ciliates.
Dinoflagellates exhibit extensive morphological diversity and can be photosynthetic, heterotrophic, or mixotrophic. The chloroplast of photosynthetic dinoflagellates was derived by secondary endosymbiosis of a red alga. Many dinoflagellates are encased in interlocking plates of cellulose. Two perpendicular flagella fit into the grooves between the cellulose plates, with one flagellum extending longitudinally and a second encircling the dinoflagellate ((Figure)). Together, the flagella contribute to the characteristic spinning motion of dinoflagellates. These protists exist in freshwater and marine habitats, and are a component of plankton, the typically microscopic organisms that drift through the water and serve as a crucial food source for larger aquatic organisms.
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Dinoflagellates have a nuclear variant called a dinokaryon. The chromosomes in the dinokaryon are highly condensed throughout the cell cycle and do not have typical histones. Mitosis in dinoflagellates is closed, that is, the spindle separates the chromosomes from outside of the nucleus without breakdown of the nuclear envelope.
Some dinoflagellates generate light, called bioluminescence, when they are jarred or stressed. Large numbers of marine dinoflagellates (billions or trillions of cells per wave) can emit light and cause an entire breaking wave to twinkle or take on a brilliant blue color ((Figure)). For approximately 20 species of marine dinoflagellates, population explosions (also called blooms) during the summer months can tint the ocean with a muddy red color. This phenomenon is called a red tide, and it results from the abundant red pigments present in dinoflagellate plastids. In large quantities, these dinoflagellate species secrete an asphyxiating toxin that can kill fish, birds, and marine mammals. Red tides can be massively detrimental to commercial fisheries, and humans who consume these protists may become poisoned.
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The apicomplexan protists are named for a structure called an apical complex ((Figure)), which appears to be a highly modified secondary chloroplast. The apicoplast genome is similar to those of dinoflagellate chloroplasts. The apical complex is specialized for entry and infection of host cells. Indeed, all apicomplexans are parasitic. This group includes the genus Plasmodium, which causes malaria in humans. Apicomplexan life cycles are complex, involving multiple hosts and stages of sexual and asexual reproduction.
The ciliates, which include Paramecium and Tetrahymena, are a group of protists 10 to 3,000 micrometers in length that are covered in rows, tufts, or spirals of tiny cilia. By beating their cilia synchronously or in waves, ciliates can coordinate directed movements and ingest food particles. Certain ciliates have fused cilia-based structures that function like paddles, funnels, or fins. Ciliates also are surrounded by a pellicle, providing protection without compromising agility. The genus Paramecium includes protists that have organized their cilia into a plate-like primitive mouth, called an oral groove, which is used to capture and digest bacteria ((Figure)). Food captured in the oral groove enters a food vacuole, where it combines with digestive enzymes. Waste particles are expelled by an exocytic vesicle that fuses at a specific region on the cell membrane, called the anal pore. In addition to a vacuole-based digestive system, Paramecium also uses contractile vacuoles, which are osmoregulatory vesicles that fill with water as it enters the cell by osmosis and then contract to squeeze water from the cell. Ciliates therefore exhibit considerable structural complexity without having achieved multicellularity.
Paramecium has two nuclei, a macronucleus and a micronucleus, in each cell. The micronucleus is essential for sexual reproduction, and is in many ways a typical eukaryotic nucleus, except that its genes are not transcribed. The transcribed nucleus is the macronucleus, which directs asexual binary fission and all other biological functions. The macronucleus is a multiploid nucleus constructed from the micronucleus during sexual reproduction. Periodic reconstruction of the macronucleus is necessary because the macronucleus divides amitotically, and thus becomes genetically unbalanced over a period of successive cell replications. Paramecium and most other ciliates reproduce sexually by conjugation. This process begins when two different mating types of Paramecium make physical contact and join with a cytoplasmic bridge ((Figure)). The diploid micronucleus in each cell then undergoes meiosis to produce four haploid micronuclei. Three of these degenerate in each cell, leaving one micronucleus that then undergoes mitosis, generating two haploid micronuclei. The cells each exchange one of these haploid nuclei and move away from each other. Fusion of the haploid micronuclei generates a completely novel diploid pre-micronucleus in each conjugative cell. This pre-micronucleus undergoes three rounds of mitosis to produce eight copies, and the original macronucleus disintegrates. Four of the eight pre-micronuclei become full-fledged micronuclei, whereas the other four perform multiple rounds of DNA replication. The copies of the micronuclear chromosomes are severely edited to form hundreds of smaller chromosomes that contain only the protein coding genes. Each of these smaller chromosomes gets new telomeres as the macronucleus differentiates. Two cycles of cell division then yield four new Paramecia from each original conjugative cell.
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