Many of the protist species classified into the supergroup Excavata are asymmetrical, single-celled organisms with a feeding groove “excavated” from one side. This supergroup includes heterotrophic predators, photosynthetic species, and parasites. Its subgroups are the diplomonads, parabasalids, and euglenozoans. The group includes a variety of modified mitochondria, as well as chloroplasts derived from green algae by secondary endosymbiosis. Many of the euglenozoans are free-living, but most diplomonads and parabasalids are symbionts or parasites.

Diplomonads

Figure 1. The mammalian intestinal parasite Giardia lamblia, visualized here using scanning electron microscopy, is a waterborne protist that causes severe diarrhea when ingested. (credit: modification of work by Janice Carr, CDC; scale-bar data from Matt Russell)

Among the Excavata are the diplomonads, which include the intestinal parasite, Giardia lamblia (Figure 1). Until recently, these protists were believed to lack mitochondria. Mitochondrial remnant organelles, called mitosomes, have since been identified in diplomonads, but although these mitosomes are essentially nonfunctional as respiratory organelles, they do function in iron and sulfur metabolism. Diplomonads exist in anaerobic environments and use alternative pathways, such as glycolysis, to generate energy. Each diplomonad cell has two similar, but not identical haploid nuclei. Diplomonads have four pairs of locomotor flagella that are fairly deeply rooted in basal bodies that lie between the two nuclei.

Parabasalids

A second Excavata subgroup, the parabasalids, are named for the parabasal apparatus, which consists of a Golgi complex associated with cytoskeletal fibers. Other cytoskeletal features include an axostyle, a bundle of fibers that runs the length of the cell and may even extend beyond it. Parabasalids move with flagella and membrane rippling, and these and other cytoskeletal modifications may assist locomotion. Like the diplomonads, the parabasalids exhibit modified mitochondria. In parabasalids these structures function anaerobically and are called hydrogenosomes because they produce hydrogen gas as a byproduct.

The parabasalid Trichomonas vaginalis causes trichomoniasis, a sexually transmitted disease in humans, which appears in an estimated 180 million cases worldwide each year. Whereas men rarely exhibit symptoms during an infection with this protist, infected women may become more susceptible to secondary infection with human immunodeficiency virus (HIV) and may be more likely to develop cervical cancer. Pregnant women infected with T. vaginalis are at an increased risk of serious complications, such as pre-term delivery.

Some of the most complex of the parabasalids are those that colonize the rumen of ruminant animals and the guts of termites. These organisms can digest cellulose, a metabolic talent that is unusual among eukaryotic cells. They have multiple flagella arranged in complex patterns and some additionally recruit spirochetes that attach to their surface to act as accessory locomotor structures.

Euglenozoans

Euglenozoans includes parasites, heterotrophs, autotrophs, and mixotrophs, ranging in size from 10 to 500 µm. Euglenoids move through their aquatic habitats using two long flagella that guide them toward light sources sensed by a primitive ocular organ called an eyespot. The familiar genus, Euglena, encompasses some mixotrophic species that display a photosynthetic capability only when light is present. The chloroplast of Euglena descends from a green alga by secondary endosymbiosis. In the dark, the chloroplasts of Euglena shrink up and temporarily cease functioning, and the cells instead take up organic nutrients from their environment. Euglena has a tough pellicle composed of bands of protein attached to the cytoskeleton. The bands spiral around the cell and give Euglena its exceptional flexibility.

The human parasite, Trypanosoma brucei, belongs to a different subgroup of Euglenozoa, the kinetoplastids. The kinetoplastid subgroup is named after the kinetoplast, a large modified mitochondrion carrying multiple circular DNAs. This subgroup includes several parasites, collectively called trypanosomes, which cause devastating human diseases and infect an insect species during a portion of their life cycle. T. brucei develops in the gut of the tsetse fly after the fly bites an infected human or other mammalian host. The parasite then travels to the insect salivary glands to be transmitted to another human or other mammal when the infected tsetse fly consumes another blood meal. T. brucei is common in central Africa and is the causative agent of African sleeping sickness, a disease associated with severe chronic fatigue, coma, and can be fatal if left untreated since it leads to progressive decline of the function of the central nervous system.

Figure 2. Trypanosoma brucei, the causative agent of sleeping sickness, spends part of its life cycle in the tsetse fly and part in humans. (credit: modification of work by CDC)