| Dictyostelid | |
|---|---|
 | |
| Dictyostelium discoideum | |
Scientific classification  | |
| Domain: | Eukaryota |
| Phylum: | Amoebozoa |
| Subphylum: | Conosa |
| Infraphylum: | Eumycetozoa |
| Class: | Dictyostelia Lister 1909, emend. Olive 1970 |
| Orders | |
The dictyostelids (Dictyostelia/Dictyostelea, ICZN, Dictyosteliomycetes, ICBN) or cellular slime molds are a group of slime molds or social amoebae.
When food (normally bacteria) is readily available dictyostelids behave as individual amoebae, which feed and divide normally. However, when the food supply is exhausted, they aggregate to form a multicellular assembly, called a pseudoplasmodium, grex, or slug (not to be confused with the gastropod mollusc called a slug). The grex has a definite anterior and posterior, responds to light and temperature gradients, and has the ability to migrate. Under the correct circumstances the grex matures forming a sorocarp (fruiting body) with a stalk supporting one or more sori (balls of spores). These spores are inactive cells protected by resistant cell walls, and become new amoebae once food is available.
In Acytostelium, the sorocarp is supported by a stalk composed of cellulose, but in other dictyostelids the stalk is composed of cells, sometimes taking up the majority of the original amoebae. With a few exceptions, these cells die during stalk formation, and there is a definite correspondence between parts of the grex and parts of the fruiting body. Aggregation of amoebae generally takes place in converging streams. The amoebae move using filose pseudopods, and are attracted to chemicals produced by other amoebae. In Dictyostelium discoideum , aggregation is signalled by cAMP, but others use different chemicals. In the species Dictyostelium purpureum , the grouping is by kinship, not just by proximity.
Dictyostelium has been used as a model organism in molecular biology and genetics, and is studied as an example of cell communication, differentiation, and programmed cell death. It is also an interesting example of the evolution of cooperation and cheating. [1] [2] [3] A large body of research data concerning D. discoideum is available on-line at DictyBase.
The mechanism behind the aggregation of the amoebae relies on cyclic adenosine monophosphate (cAMP) as a signal molecule. One cell, the founder of the colony, begins to secrete cAMP in response to stress. Others detect this signal, and respond in two ways:
The effect of this is to relay the signal throughout the nearby population of amoebae and cause inward movement to the area of highest cAMP concentration.
Within an individual cell, the mechanism is as follows:
Because the internal cAMP concentration inactivates the receptor for external cAMP, an individual cell shows oscillatory behaviour. This behaviour produces beautiful spirals seen in converging colonies and is reminiscent of the Belousov–Zhabotinsky reaction and two-dimensional cyclic cellular automata.
The entire genome of Dictyostelium discoideum was published in Nature in 2005 by geneticist Ludwig Eichinger and coworkers. [4] The haploid genome contains approximately 12,500 genes on 6 chromosomes. For comparison, the diploid human genome has 20,000-25,000 genes (represented twice) on 23 chromosome pairs. There is a high level of the nucleotides adenosine and thymidine (~77%) leading to a codon usage that favors more adenosines and thymidines in the third position. Tandem repeats of trinucleotides are abundant in Dictyostelium, which in humans cause Trinucleotide repeat disorders.
Sexual development can occur when amoeboid cells are starved for their bacterial food supply and dark humid conditions are present. [5] Both heterothallic and homothallic strains of Dictostelium can undergo mating. Heterothallic sexual development has been most extensively studied in D. discoideum , and homothallic sexual development has been most well studied in D. mucoroides. [6] Heterothallic matings are initiated by fusion of haploid cells (gametes) from two strains of opposite mating type. This contrasts with homothallic strains that appear to express both mating types. [7]
Mating is initiated by gametogenesis that produces small, motile gametes that fuse to form a small binucleate cell. The volume of the binucleate cell then increases to produce a giant binuclear cell. As growth proceeds, the nuclei swell, and then fuse forming a true diploid zygote giant cell. As this is occurring, amoebae have been undergoing cAMP-induced chemotaxis towards the giant cell surface. This forms a cellular aggregate and at the center of the aggregate the zygote giant cell ingests the surrounding amoebae. Phagocytosis is followed by digestion of the ingested amoebae. Next the zygote forms a macrocyst characterized by a surrounding extracellular cellulose sheath. After the macrocyst is formed it ordinarily remains dormant for a period before germination can occur. [8] Within the macrocyst the diploid zygote undergoes meiosis followed by successive mitotic divisions. When the macrocyst germinates it releases many haploid amoeboid cells.
The Dictyostelium phylogeny tree has undergone multiple changes in the past decades. The first dictyostelid to be described was Dictyostelium mucoroides in 1869 by Osker Brefeld, and the original discovery of Dictyostelium discoideum occurred in 1935, [9] with further discoveries from Kenneth Raper, followed by global efforts led by James Cavender and collaborators. Dictyostelium discoideum was initially classified under ‘lower fungi,’ but the classification has since shifted the classification to under the Amoebozoa phylum where it is currently. [4]
The groupings within the dictyostelid phylogeny tree has undergone frequent reordering due to availability new evidence. Most currently accepted phylogenies of dictyostelids utilize genome sequencing and small subunit ribosomal DNA (ssu-rDNA). The dictyostelids can be further subdivided into four groups. In particular, group 4 contains the Dictyostelium discoideum species and differs from the other groups with its usage of cAMP as an attractant emitted during aggregation. [10]
Fossil calibrations indicate that dictyostelids class originally diverged into two major branches approximately 0.52 billion years ago. Current theories speculate that dictyostelid stalk and spore formation originally evolved as an adaption to global glacial formations. Further subdivision of dictyostelids species likely arose as most glacial formations melted. Most species in major groups 1, 2, and 3 display an encystment ability that allows individuals to survive low temperatures, but spores have shown an increased ability in resisting lower temperatures. Group 4 differs from other major groups in a general lacking ability to encyst, but its spores have shown better resistances against lower temperatures relative to spores from other groups. [11]
The internal phylogeny of the Dictyostelids is shown in the cladogram. [12]
Class DictyosteliaLister 1909 em. Olive 1970 [13]
Dictyostelium shares many molecular features with macrophages, the human host of Legionella . The cytoskeletal composition of D. discoideum is similar to that of mammalian cells as are the processes driven by these components, such as phagocytosis, membrane trafficking, endocytic transit and vesicle sorting. Like leukocytes, D. discoideum possess chemotactic capacity. Hence, D. discoideum represents a suitable model system to ascertain the influence of a variety of host cell factors during Legionella infections. [14]
Slime mold or slime mould is an informal name given to a polyphyletic assemblage of unrelated eukaryotic organisms in the Stramenopiles, Rhizaria, Discoba, Amoebozoa and Holomycota clades. Most are microscopic; those in the Myxogastria form larger plasmodial slime molds visible to the naked eye.
Mycetozoa is a polyphyletic grouping of slime molds. It was originally thought to be a monophyletic clade, but recently it was discovered that protostelia are a polyphyletic group within Conosa.
Dictyostelium is a genus of single- and multi-celled eukaryotic, phagotrophic bacterivores. Though they are Protista and in no way fungal, they traditionally are known as "slime molds". They are present in most terrestrial ecosystems as a normal and often abundant component of the soil microflora, and play an important role in the maintenance of balanced bacterial populations in soils.
Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional classification schemes, Amoebozoa is usually ranked as a phylum within either the kingdom Protista or the kingdom Protozoa. In the classification favored by the International Society of Protistologists, it is retained as an unranked "supergroup" within Eukaryota. Molecular genetic analysis supports Amoebozoa as a monophyletic clade. Modern studies of eukaryotic phylogenetic trees identify it as the sister group to Opisthokonta, another major clade which contains both fungi and animals as well as several other clades comprising some 300 species of unicellular eukaryotes. Amoebozoa and Opisthokonta are sometimes grouped together in a high-level taxon, named Amorphea. Amoebozoa includes many of the best-known amoeboid organisms, such as Chaos, Entamoeba, Pelomyxa and the genus Amoeba itself. Species of Amoebozoa may be either shelled (testate) or naked, and cells may possess flagella. Free-living species are common in both salt and freshwater as well as soil, moss and leaf litter. Some live as parasites or symbionts of other organisms, and some are known to cause disease in humans and other organisms.
The family Acrasidae is a family of slime molds which belongs to the excavate group Percolozoa. The name element acrasio- comes from the Greek akrasia, meaning "acting against one's judgement". This group consists of cellular slime molds.
Eumycetozoa, or true slime molds, is a diverse group of protists that behave as slime molds and develop fruiting bodies, either as sorocarps or as sporocarps. It is a monophyletic group or clade within the phylum Amoebozoa that contains the myxogastrids, dictyostelids and protosporangiids.
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 life cycle consists of four stages: vegetative, aggregation, migration, and culmination. The life cycle of D. discoideum is relatively short, which allows for timely viewing of all stages. The cells involved in the life cycle undergo movement, chemical signaling, and development, which are applicable to human cancer research. The simplicity of its life cycle makes D. discoideum a valuable model organism to study genetic, cellular, and biochemical processes in other organisms.
Homothallic refers to the possession, within a single organism, of the resources to reproduce sexually; i.e., having male and female reproductive structures on the same thallus. The opposite sexual functions are performed by different cells of a single mycelium.
Conosa is a grouping of Amoebozoa. It is subdivided into three groups: Archamoeba, Variosea and Mycetozoa.
Protosteliomycetes/Protosteliales (ICBN) or Protostelea/Protostelia/Protosteliida (ICZN) is a grouping of slime molds from the phylum Mycetozoa. The name can vary depending upon the taxon used. Other names include Protostelea, Protostelia, and Protostelida. When not implying a specific level of classification, the term protostelid or protosteloid amoeba is sometimes used.
Differentiation-inducing factor (DIF) is one of a class of effector molecules that induce changes in cell chemistry, inhibiting growth and promoting differentiation of cell type. This name has been given to several factors before it was clear if they were the same or different effectors. DIFs have garnered interest with their potential tumor inhibiting properties. DIFs have also been used to help regulate plant growth.
Dictyostelium purpureum is a species of Dictyostelium.
Acytostelium is a genus of dictyostelid.
Polysphondylium is a genus of cellular slime mold, including the species Polysphondylium pallidum. The genus was circumscribed by German mycologist Julius Oscar Brefeld in 1884.
An amoeba, often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopods. Amoebae do not form a single taxonomic group; instead, they are found in every major lineage of eukaryotic organisms. Amoeboid cells occur not only among the protozoa, but also in fungi, algae, and animals.
Tupanvirus is a genus of viruses first described in 2018. The genus is composed of two species of virus that are in the giant virus group. Researchers discovered the first isolate in 2012 from deep water sediment samples taken at 3,000 m depth off the coast of Brazil. The second isolate was collected from a soda lake in Southern Nhecolândia, Brazil in 2014. They are named after Tupã (Tupan), a Guaraní thunder god, and the places they were found. These are the first viruses reported to possess genes for amino-acyl tRNA synthetases for all 20 standard amino acids.
Pauline Schaap is a Dutch cell biologist and evolutionary biologist. She is a professor of Developmental Signalling at the University of Dundee., a corresponding member of the Royal Netherlands Academy of Arts and Sciences, a Fellow of the Royal Society of Biology, and a Fellow of the Royal Society of Edinburgh.
Amoebozoa of the free living genus Acanthamoeba and the social amoeba genus Dictyostelium are single celled eukaryotic organisms that feed on bacteria, fungi, and algae through phagocytosis, with digestion occurring in phagolysosomes. Amoebozoa are present in most terrestrial ecosystems including soil and freshwater. Amoebozoa contain a vast array of symbionts that range from transient to permanent infections, confer a range of effects from mutualistic to pathogenic, and can act as environmental reservoirs for animal pathogenic bacteria. As single celled phagocytic organisms, amoebas simulate the function and environment of immune cells like macrophages, and as such their interactions with bacteria and other microbes are of great importance in understanding functions of the human immune system, as well as understanding how microbiomes can originate in eukaryotic organisms.
Copromyxa protea is a cellular slime mold belonging to the supergroup Amoebozoa. The taxonomical history of C. protea starts as Guttulina protea and ultimately ends with becoming its own genus of protists. Its morphological features like tubular mitochondrial cristae help pinpoint its placement in its supergroup, as well as its pseudopodia. The life cycle of this cellular slime mold is outlined and expanded upon, delving into specific characteristics of each component in the stages. The specific environments where C. protea was gathered for experimental purposes is also detailed.
Acrasis kona is a eukaryotic microorganism within the family Acrasidae, notable for its life cycle that alternates between unicellular and multicellular stages.
