Dictyostelium discoideum

Dictyostelium discoideum is a species of soil-dwelling amoeba belonging to the phylum Amoebozoa, infraphylum Mycetozoa. Commonly referred to as slime mold, D. discoideum is a eukaryote that transitions from a collection of unicellular amoebae into a multicellular slug and then into a fruiting body within its lifetime. Its unique asexual lifecycle consists of four stages: vegetative, aggregation, migration, and culmination. The lifecycle of D. discoideum is relatively short, which allows for timely viewing of all stages. The cells involved in the lifecycle undergo movement, chemical signaling, and development, which are applicable to human cancer research. The simplicity of its lifecycle makes D. discoideum a valuable model organism to study genetic, cellular, and biochemical processes in other organisms. In the wild, D. discoideum can be found in soil and moist leaf litter. Its primary diet consists of bacteria, such as Escherichia coli, found in the soil and decaying organic matter. Uninucleate amoebae of D. discoideum consume bacteria found in its natural habitat, which includes deciduous forest soil and decaying leaves. The life cycle of D. discoideum begins as spores are released from a mature sorocarp (fruiting body). Myxamoebae hatch from the spores under warm and moist conditions. During their vegetative stage, the myxamoebae divide by mitosis as they feed on bacteria. The bacteria secrete folic acid, attracting the myxamoebae. When the supply of bacteria is depleted, the myxamoebae enter the aggregation stage. During aggregation, starvation initiates the creation of a biochemical machinery that includes glycoproteins and adenylyl cyclase. The glycoproteins allow for cell-cell adhesion, and adenylyl cyclase creates cyclic AMP. Cyclic AMP is secreted by the amoebae to attract neighboring cells to a central location. As they move toward the signal, they bump into each other and stick together by the use of glycoprotein adhesion molecules. The migration stage begins once the amoebae have formed a tight aggregate and the elongated mound of cells tip over to lie flat on the ground. The amoebae work together as a motile pseudoplasmodium, also known as a slug. The slug is about 2–4 mm long, composed of up to 100,000 cells, and is capable of movement by producing a cellulose sheath in its anterior cells through which the slug moves. Part of this sheath is left behind as a slimy trail as it moves toward attractants such as light, heat, and humidity in a forward-only direction. Cyclic AMP and a substance called differentiation-inducing factor, help to form different cell types. The slug becomes differentiated into prestalk and prespore cells that move to the anterior and posterior ends, respectively. Once the slug has found a suitable environment, the anterior end of the slug forms the stalk of the fruiting body and the posterior end forms the spores of the fruiting body. Anterior-like cells, which have only been recently discovered, are also dispersed throughout the posterior region of the slug. These anterior-like cells form the very bottom of the fruiting body and the caps of the spores. After the slug settles into one spot, the posterior end spreads out with the anterior end raised in the air, forming what is called the 'Mexican hat', and the culmination stage begins. The prestalk cells and prespore cells switch positions in the culmination stage to form the mature fruiting body. The anterior end of the Mexican hat forms a cellulose tube, which allows the more posterior cells to move up the outside of the tube to the top, and the prestalk cells move down. This rearrangement forms the stalk of the fruiting body made up of the cells from the anterior end of the slug, and the cells from the posterior end of the slug are on the top and now form the spores of the fruiting body. At the end of this 8– to 10-hour process, the mature fruiting body is fully formed. This fruiting body is 1–2 mm tall and is now able to start the entire cycle over again by releasing the mature spores that become myxamoebae. In general, although D. discoideum generally reproduces asexually, D. discoideum is still capable of sexual reproduction if certain conditions are met. D. discoideum has three different mating types and studies have identified the sex locus that specifies these three mating types. Type I strains are specified by the gene called MatA, Type II strains have three different genes: MatB (homologous to Mat A), Mat C, and Mat D, and Type III strains have Mat S and Mat T genes (which are homologous to Mat C and Mat D). These sexes can only mate with the two different sexes and not with its own. When incubated with their bacterial food supply, heterothallic or homothallic sexual development can occur, resulting in the formation of a diploid zygote. Heterothallic mating occurs when two amoebae of different mating types are present in a dark and wet environment, where they can fuse during aggregation to form a giant zygote cell. The giant cell then releases cAMP to attract other cells, then engulfs the other cells cannibalistically in the aggregate. The consumed cells serve to encase the whole aggregate in a thick, cellulose wall to protect it. This is known as a macrocyst. Inside the macrocyst, the giant cell divides first through meiosis, then through mitosis to produce many haploid amoebae that will be released to feed as normal amoebae would. Homothallic D. discoideum strains AC4 and ZA3A are also able to produce macrocysts. Each of these strains, unlike heterothallic strains, likely express both mating type alleles (matA and mata). While sexual reproduction is possible, it is very rare to see successful germination of a D. discoideum macrocyst under laboratory conditions. Nevertheless, recombination is widespread within D. discoideum natural populations, indicating that sex is likely an important aspect of their lifecycle.