717:, after entry into amoebae, in a phagosome the fusion with lysosomes is blocked and these unmatured phagosomes are surrounded by host cell organelles such as mitochondria, vesicles, and a multilayer membrane derived from the rough endoplasmic reticulum (RER) of amoebae. The role of the RER in the intracellular infection is not known, but the RER is not required as a source of proteins for the bacteria. The bacteria reside within these phagosomes during the aggregation and the multicellular development stages. The amoebae preserve their individuality and each amoeba has its own bacterium. During the culmination stage, when the spores are produced, the bacteria pass from the cell to the sorus with the help of a cytoskeletal structure that prevents host cell destruction. Some results suggest the bacteria exploit the exocytosis without killing the cell. Free-living amoebae seem to play a crucial role for persistence and dispersal of some pathogens in the environment. Transient association with amoebae has been reported for a number of different bacteria, including
677:" of the bacteria when the bacteria were included within the slime mold fruiting bodies. The incorporation of the bacteria into the fruiting bodies allows the "seeding" of the food source at the location of the spore dispersal, which is particularly valuable if the new location is low in food resources. Colonies produced from the "farming" spores typically also show the same behavior when sporulating. This incorporation has a cost associated with it: Those colonies that do not consume all of the prey bacteria produce smaller spores that cannot disperse as widely. In addition, much less benefit exists for bacteria-containing spores that land in a food-rich region. This balance of the costs and benefits of the behavior may contribute to the fact that a minority of
353:, 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.
496:, the amoeba secretes the signal, cAMP, out of the cell, attracting other amoebae to migrate toward the source. Every amoeba moves toward a central amoeba, the one dispensing the greatest amount of cAMP secretions. The secretion of the cAMP is then exhibited by all amoebae and is a call for them to begin aggregation. These chemical emissions and amoeba movement occur every six minutes. The amoebae move toward the concentration gradient for 60 seconds and stop until the next secretion is sent out. This behavior of individual cells tends to cause oscillations in a group of cells, and chemical waves of varying cAMP concentration propagate through the group in spirals.
785:. When exposed to a toxic environment created by the use of ethidium bromide, it was shown that the number of sentinel cells per millimeter was lower for farmers than non-farmers. This was concluded by observing the trails left behind as the slugs migrated and counting the number of sentinel cells present in a millimeter. However, the number of sentinel cells does not affect the spore production and viability in farmers. Farmers exposed to a toxic environment produce the same number of spores as farmers in a non-toxic environment, and the spore viability was the same between the farmers and non-farmers. When Clade 2
703:
819:
Mycetozoa, a group that includes the protostelid, dictyostelid, and myxogastrid slime molds. Elongation factor-1α (EF-1α) data analyses support
Mycetozoa as a monophyletic group, though rRNA trees place it as a polyphyletic group. Further, these data support the idea that the dictyostelid and myxogastrid are more closely related to each other than they are the protostelids. EF-1α analysis also placed the Mycetozoa as the immediate outgroup for the animal-fungal clade. Latest phylogenetic data place dictyostelids firmly within supergroup
33:
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541:) rather than following simple probabilistic or continuous behaviour. Bursting transcription now appears to be conserved between bacteria and humans. Another remarkable feature of the organism is that it has sets of DNA repair enzymes found in human cells, which are lacking from many other popular metazoan model systems. Defects in DNA repair lead to devastating human cancers, so the ability to study human repair proteins in a simple tractable model will prove invaluable.
188:
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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.
530:
cellulose coat and extend as a tube through the grex. As they differentiate, they form vacuoles and enlarge, lifting up the prespore cells. The stalk cells undergo apoptosis and die as the prespore cells are lifted high above the substrate. The prespore cells then become spore cells, each one becoming a new myxamoeba upon dispersal. This is an example of how apoptosis is used in the formation of a reproductive organ, the mature fruiting body.
50:
519:
complicated behavior has been analyzed by computer modeling of the behavior and the periodic pattern of temperature changes in soil caused by daily changes in air temperature. The conclusion is that the behavior moves slugs a few centimeters below the soil surface up to the surface. This is an amazingly sophisticated behavior by a primitive organism with no apparent sense of gravity.
565:, which is adequate for all stages of the life cycle. When the food supply is diminished, the myxamoebae aggregate to form pseudoplasmodia. Soon, the dish is covered with various stages of the life cycle. Checking the dish often allows for detailed observations of development. The cells can be harvested at any stage of development and grown quickly.
51:
52:
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hinder, stop, or accelerate development. Variations of these parameters can alter the rate and viability of culture growth. Also, the fruiting bodies, being that this is the tallest stage of development, are very responsive to air currents and physical stimuli. It is unknown if there is a stimulus involved with spore release.
741:’s report has made an important advance in the knowledge of amoebic behavior, and the famous Spanish phrase translated as “you are more stupid than an amoeba” is losing the sense because amoebae are an excellent example of social behavior with an amazing coordination and sense of sacrifice for the benefit of the species.
53:
695:, but some of the phagocytized bacteria, including some human pathogens, are able to live in the amoebae and exit without killing the cell. When they enter the cell, where they reside, and when they leave the cell are not known. The research is not yet conclusive but it is possible to draw a general life cycle of
791:, or farmer-associated bacteria, are removed from the farmers, spore production and viability were similar to that of the non-farmers. Thus, it is suggested that bacteria carried by the farmers provide an additional role of protection for the farmers against potential harm due to toxins or pathogens.
503:
was discovered by mathematical biologists Thomas Höfer and Martin
Boerlijst. Mathematical biologist Cornelis J. Weijer has proven that similar equations can model its movement. The equations of these patterns are mainly influenced by the density of the amoeba population, the rate of the production of
458:, which are all normal cellular processes. It is also used to study other aspects of development, including cell sorting, pattern formation, phagocytosis, motility, and signal transduction. These processes and aspects of development are either absent or too difficult to view in other model organisms.
385:
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
756:
are phagocytic cells responsible for removing toxic material from the slug stage of the social cycle. Generally round in shape, these cells are present within the slug sheath where they are found to be circulating freely. The detoxification process occurs when these cells engulf toxins and pathogens
401:
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,
764:
Sentinel cells make up approximately 1% of the total number of slug cells, and the number of sentinel cells remains constant even as they are being released. This indicates a constant regeneration of sentinel cells within the slugs as they are being removed along with toxins and pathogens. Sentinel
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in a laboratory, it is important to take into account its behavioral responses. For example, it has an affinity toward light, higher temperatures, high humidity, low ionic concentrations, and the acidic side of the pH gradient. Experiments are often done to see how manipulations of these parameters
518:
is movement along a gradient of temperature. The slugs have been shown to migrate along extremely shallow gradients of only 0.05 °C/cm, but the direction chosen is complicated; it seems to be away from a temperature about 2 °C below the temperature to which they had been acclimated. This
356:
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
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process occurs when a cell becomes more specialized to develop into a multicellular organism. Changes in size, shape, metabolic activities, and responsiveness can occur as a result of adjustments in gene expression. Cell diversity and differentiation, in this species, involves decisions made from
328:
The migration stage begins once the amoebae have formed a tight aggregate and the elongated mound of cells tips 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
529:
altruistically sacrifice themselves in the formation of the mature fruiting body. During the pseudoplasmodium (slug or grex) stage of its life cycle, the organism has formed three main types of cells: prestalk, prespore, and anterior-like cells. During culmination, the prestalk cells secrete a
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has maintained more of its ancestral genome diversity than plants and animals, although proteome-based phylogeny confirms that amoebozoa diverged from the animal–fungal lineage after the plant–animal split. Subclass
Dictyosteliidae, order Dictyosteliales is a monophyletic assemblage within the
865:
Every genome sequenced plays an important role in identifying genes that have been gained or lost over time. Comparative genomic studies allow for comparison of eukaryotic genomes. A phylogeny based on the proteome showed that the amoebozoa deviated from the animal-fungal lineage after the
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Tandem repeats of trinucleotides are very abundant in this genome; one class of the genome is clustered, leading researchers to believe it serves as centromeres. The repeats correspond to repeated sequences of amino acids and are thought to be expanded by nucleotide expansion. Expansion of
507:
The use of cAMP as a chemotactic agent is not established in any other organism. In developmental biology, this is one of the comprehensible examples of chemotaxis, which is important for an understanding of human inflammation, arthritis, asthma, lymphocyte trafficking, and axon guidance.
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Note, however, that cAMP oscillations may not be necessary for the collective cell migration at multicellular stages. A study has found that cAMP-mediated signaling changes from propagating waves to a steady state at a multicellular stage of D. discoideum.
232:
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
811:. This was a class of cellular slime molds, which was characterized by the aggregation of individual amoebae into a multicellular fruiting body, making it an important factor that related the acrasids to the dictyostelids.
474:
cell-cell interactions in pathways to either stalk cells or spore cells. These cell fates depend on their environment and pattern formation. Therefore, the organism is an excellent model for studying cell differentiation.
454:. It can be observed at organismic, cellular, and molecular levels primarily because of their restricted number of cell types and behaviors, and their rapid growth. It is used to study cell differentiation, chemotaxis, and
325:. 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.
54:
508:
Phagocytosis is used in immune surveillance and antigen presentation, while cell-type determination, cell sorting, and pattern formation are basic features of embryogenesis that may be studied with these organisms.
491:
is defined as a passage of an organism toward or away from a chemical stimulus along a chemical concentration gradient. Certain organisms demonstrate chemotaxis when they move toward a supply of nutrients. In
920:
761:. Then, the cells clump into groups of five to ten cells, which then attach to the inner sheath of the slug. The sheath is sloughed off as the slug migrates to a new site in search of food bacteria.
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cyclic AMP and the sensitivity of individual amoebas to cyclic AMP. The spiraling pattern is formed by amoebas at the centre of a colony who rotate as they send out waves of cyclic AMP.
537:
research has come from new techniques allowing the activity of individual genes to be visualised in living cells. This has shown that transcription occurs in "bursts" or "pulses" (
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sequencing project was completed and published in 2005 by an international collaboration of institutes. This was the first free-living protozoan genome to be fully sequenced.
585:
has been hampered by large shifts in the protein expression profile between different developmental stages and a general lack of commercially available antibodies for
525:(programmed cell death) is a normal part of species development. Apoptosis is necessary for the proper spacing and sculpting of complex organs. Around 20% of cells in
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strains AC4 and ZA3A are also able to produce macrocysts. Each of these strains, unlike heterothallic strains, likely express both mating type alleles (
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that transitions from a collection of unicellular amoebae into a multicellular slug and then into a fruiting body within its lifetime. Its unique asexual
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consists of a 34-Mb haploid genome with a base composition of 77% and contains six chromosomes that encode around 12,500 proteins. Sequencing of the
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cells are present in the slug even when there are no toxins or pathogens to be removed. Sentinel cells have been located in five other species of
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298:(fruiting body). Myxamoebae hatch from the spores under warm and moist conditions. During their vegetative stage, the myxamoebae divide by
2039:
Grant P.Ottom Mary Y.Wu; Margaret Clarke; Hao Lu; O.Roger
Anderson; Hubert Hilbi; Howard A. Shuman; Richard H. Kessin (11 November 2003).
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sexual development can occur, resulting in the formation of a diploid zygote. Heterothallic mating occurs when two amoebae of different
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1517:"Biomathematics Patterns: Spiral Slime. MATHEMATICAL RECREATIONS by Ian Stewart. Finding mathematics in creatures great and small"
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1778:"The use of strepavidin conjugates as immunoblot loading controls of mitochondrial markers for use with Dictyostelium discoidium"
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cells through which the slug moves. Part of this sheath is left behind as a slimy trail as it moves toward attractants such as
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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.
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This organism's ability to be easily isolated and cultivated in the laboratory adds to its appeal as a model organism. While
2363:"Eumycetozoa = Amoebozoa?: SSUrDNA Phylogeny of Protosteloid Slime Molds and Its Significance for the Amoebozoan Supergroup"
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733:, among others. Agriculture seems to play a crucial role for pathogens' survival, as they can live and replicate inside
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have turned out to be polyphyletic, their stalked fruiting bodies a convergent feature of multiple unrelated lineages.
306:, which attracts the myxamoebae. When the supply of bacteria is depleted, the myxamoebae enter the aggregation stage.
1291:"Variation, sex, and social cooperation: molecular population genetics of the social amoeba Dictyostelium discoideum"
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Tyler M.S. 2000. Developmental
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2041:"Macroautophagy is dispensable for intracellular replication of Legionella pneumophila in Dictyostelium discoideum"
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Brock DA, Douglas TE, Queller DC, Strassmann JE (20 January 2011). "Primitive agriculture in a social amoeba".
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Robson GE, Williams KL (April 1980). "The mating system of the cellular slime mould
Dictyostelium discoideum".
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In the picture, one can see the different stages. First, in the starvation stage, bacteria are enclosed within
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is closely related to higher metazoans. It carries similar genes and pathways, making it a good candidate for
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Hudson J. J.; Hsu D. W.; Guo K.; Zhukovskaya N.; Liu P. H.; Williams J. G.; Pears C. J.; Lakin N. D. (2005).
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Flowers JM, Li SI, Stathos A, Saxer G, Ostrowski EA, Queller DC, Strassmann JE, Purugganan MD (July 2010).
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genome is noteworthy because its many encoded proteins are commonly found in fungi, plants, and animals.
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Bloomfield, Gareth; Skelton, Jason; Ivens, Alasdair; Tanaka, Yoshimasa; Kay, Robert R. (2010-12-10).
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a valuable model organism to study genetic, cellular, and biochemical processes in other organisms.
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An elegant set of mathematical equations that reproduces the spirals and the streaming patterns of
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reported an antibody-free protein visualization standard for immunoblotting based on detection of
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Cavender J.C.; Spiegl F.; Swanson A. (2002). "Taxonomy, slime molds, and the questions we ask".
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consists of four stages: vegetative, aggregation, migration, and culmination. The life cycle of
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trinucleotide repeats also occurs in humans, in general leading to many diseases. Learning how
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1555:"Collective cell migration of Dictyostelium without cAMP oscillations at multicellular stages"
2016:
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and the surfaces are kept moist. The cultures grow best at 22–24 °C (room temperature).
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cells endure these amino acid repeats may provide insight to allow humans to tolerate them.
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natural populations, indicating that sex is likely an important aspect of their life cycle.
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shares with mammalian host cells a similar cytoskeleton and cellular processes relevant to
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Brock, Debra A.; Callison, W. Éamon; Strassmann, Joan E.; Queller, David C. (2016-04-27).
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8:
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769:, which suggests that sentinel cells can be described as a general characteristic of the
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macrocyst under laboratory conditions. Nevertheless, recombination is widespread within
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can be grown in liquid culture, it is usually grown in Petri dishes containing nutrient
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Because many of its genes are homologous to human genes, yet its life cycle is simple,
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During aggregation, starvation initiates the production of protein compounds such as
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genome provides a more intricate study of its cellular and developmental biology.
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Kay R.R.; Garrod D.; Tilly R. (1978). "Requirements for cell differentiation in
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whose boundary is colored by curvature, scale bar: 5 µm, duration: 22 seconds
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1975:"Primitive Amoebae as Training Grounds for Intracellular Bacterial Pathogens"
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426:). While sexual reproduction is possible, it is very rare to see successful
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The number of sentinel cells varies depending on the farming status of wild
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and is a suitable model for studying the infection process. Specifically,
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amoebae that will be released to feed as normal amoebae would. Homothallic
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Shadwick, LL; Spiegel, FW; Shadwick, JD; Brown, MW; Silberman, JD (2009).
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can be found in soil and moist leaf litter. Its primary diet consists of
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1669:"DNA-PKcs-dependent signaling of DNA damage in Dictyostelium discoideum"
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published findings that demonstrated a "primitive farming behaviour" in
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adapted for farmer clones to better understand this symbiotic process.
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Over sneeuwkristallen en zebrastrepen. De wereld volgens de wiskunde
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in a forward-only direction. Cyclic AMP and a substance called
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consume bacteria found in their natural habitat, which includes
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406:. Inside the macrocyst, the giant cell divides first through
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Biology: A Guide for Experimental Study.2nd ed
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at MetaMicrobe: taxonomy, facts, ontologies, and references
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Chen, Guokai; Zhuchenko, Olga; Kuspa, Adam (2007-08-03).
1850:"Amoebas show primitive farming behaviour as they travel"
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and bacterial prey, about one-third of wild-collected
2304:"Origin and evolution of the slime molds (Mycetozoa)"
2090:"Immune-like Phagocyte Activity in the Social Amoeba"
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Cavender, James C.; Raper, Kenneth B. (March 1965).
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Chubb, JR; Trcek, T; Shenoy, SM; Singer, RH (2006).
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1612:"Transcriptional pulsing of a developmental gene"
1382:Dictybase, About Dictyostelium. (1, May, 2009).
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402:cellulose wall to protect it. This is known as a
389:When incubated with their bacterial food supply,
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186:
2295:
1718:"The Acrasieae in Nature. I. Isolation"
410:, then through mitosis to produce many
2646:
2427:. Sinauer Associates. pp. 31–34.
1928:
1822:, a tractable model host organism for
1089:
646:
591:Beatson West of Scotland Cancer Centre
360:
2488:
2487:
2302:Baldauf S.L.; Doolittle W.F. (1997).
1861:
915:
1826:. In: Heuner K, Swanson M, editors.
1173:
1171:
643:, vesicle sorting, and chemotaxis.
589:antigens. In 2013, a group at the
329:capable of movement by producing a
13:
2416:
2241:Eichenger L.; et al. (2005).
2216:The Mycological Society of America
1734:10.1002/j.1537-2197.1965.tb06788.x
893:single-pass transmembrane proteins
681:colonies engage in this practice.
544:
14:
2690:
2461:
2243:"The genome of the social amoeba
1168:
882:- general genomic database about
744:
615:includes the species that causes
533:A recent major contribution from
2058:10.1046/j.1365-2958.2003.03826.x
1948:10.1111/j.1462-5822.2009.01413.x
1257:10.1111/j.1469-185X.2011.00200.x
1025:"Crawling in to a new era – the
1023:Eichinger L; Noegel, AA (2003).
929:Journal of Agricultural Research
75:
2354:
2146:
2100:(5838). New York, NY: 678–681.
2081:
1769:
1748:
1709:
1660:
1603:
1546:
1527:
1508:
1388:
1376:
381:if certain conditions are met.
351:differentiation-inducing factor
321:, and adenylyl cyclase creates
2679:Taxa named by Kenneth B. Raper
1083:
950:
909:
200:is a species of soil-dwelling
1:
2444:Developmental Biology. 8th ed
1515:Ian Stewart (November 2000).
903:
2388:10.1371/journal.pone.0006754
1472:Dusenbery, David B. (1996).
1308:10.1371/journal.pgen.1001013
1113:Cells As Experimental Models
873:
795:Classification and phylogeny
7:
2446:. Sinauer. pp. 36–39.
1991:10.1128/AEM.71.1.20-28.2005
899:and several other organisms
891:provides information about
294:are released from a mature
278:Life cycle and reproduction
10:
2695:
1722:American Journal of Botany
1536:What Shape is a Snowflake?
803:was placed in the defunct
799:In older classifications,
661:colonies. Described as a "
577:Protein expression studies
274:soil and decaying leaves.
212:. Commonly referred to as
2496:
2442:Scott F. Gilbert (2006).
1929:Clarke, Margaret (2010).
1694:10.1016/j.cub.2005.09.039
1636:10.1016/j.cub.2006.03.092
1571:10.1038/s42003-018-0273-6
935:: 135–147. Archived from
834:
673:colonies engaged in the "
167:
160:
72:Scientific classification
70:
61:
47:
39:
30:
23:
2498:Dictyostelium discoideum
2470:Dictyostelium discoideum
2329:10.1073/pnas.94.22.12007
2245:Dictyostelium discoideum
2157:Dictyostelium discoideum
1416:Dictyostelium discoideum
1245:Biol Rev Camb Philos Soc
923:Dictyostelium discoideum
884:Dictyostelium discoideum
866:plant-animal split. The
757:within the slug through
754:Dictyostelium discoideum
639:, membrane trafficking,
539:transcriptional bursting
241:Natural habitat and diet
204:belonging to the phylum
197:Dictyostelium discoideum
171:Dictyostelium discoideum
25:Dictyostelium discoideum
2114:10.1126/science.1143991
1204:10.1126/science.1197423
442:Use as a model organism
2423:Mary S. Tyler (2000).
2175:10.1098/rspb.2015.2727
2045:Molecular Microbiology
1979:Appl Environ Microbiol
1559:Communications Biology
727:Francisella tularensis
719:Legionella pneumophila
710:
693:Gram-negative bacteria
485:
450:is commonly used as a
192:
16:Species of slime mould
2654:Developmental biology
1935:Cellular Microbiology
1384:http://dictybase.org/
737:, making husbandry.
706:Life cycle of farmer
705:
635:infection, including
617:legionnaire's disease
605:Legionnaire's disease
480:
369:generally reproduces
190:
1534:Ian Stewart (2000).
1045:10.1093/emboj/cdg214
976:10.1099/mic.0.001040
771:innate immune system
687:is known for eating
609:The bacterial genus
471:cell differentiation
377:is still capable of
2379:2009PLoSO...4.6754S
2320:1997PNAS...9412007B
2314:(22): 12007–12012.
2271:10.1038/nature03481
2263:2005Natur.435...43E
2106:2007Sci...317..678C
1897:10.1038/nature09668
1889:2011Natur.469..393B
1685:2005CBio...15.1880H
1628:2006CBio...16.1018C
1521:Scientific American
1476:Life at Small Scale
1432:1978Natur.271...58K
1196:2010Sci...330.1533B
1190:(6010): 1533–1536.
889:Membranome database
773:in social amoebae.
623:is also a host for
379:sexual reproduction
361:Sexual reproduction
40:Fruiting bodies of
2169:(1829): 20152727.
1765:(1): 9. July 2013.
1354:10.1007/BF00390948
1109:Sinauer Associates
1091:Cooper, Geoffrey M
711:
568:While cultivating
561:feed primarily on
486:
193:
153:D. discoideum
2669:Amoebozoa species
2641:
2640:
2626:Open Tree of Life
2490:Taxon identifiers
2453:978-0-87893-250-4
2434:978-0-87893-843-8
1883:(7330): 393–396.
1858:, 19 January 2011
1836:978-1-904455-26-4
1797:10.2144/000114054
1487:978-0-7167-5060-4
1122:978-0-87893-106-4
651:A 2011 report in
185:
184:
55:
2686:
2634:
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2608:
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2605:NBNSYS0000015009
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2085:
2079:
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2020:
2012:
2002:
1970:
1961:
1960:
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1440:10.1038/271058a0
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1175:
1166:
1156:
1127:
1126:
1087:
1081:
1078:
1067:
1066:
1056:
1039:(9): 1941–1946.
1033:The EMBO Journal
1020:
1005:
1004:
978:
954:
948:
947:
945:
944:
913:
731:Escherichia coli
315:adenylyl cyclase
272:deciduous forest
256:Escherichia coli
173:
80:
79:
57:
56:
35:
21:
20:
2694:
2693:
2689:
2688:
2687:
2685:
2684:
2683:
2659:Model organisms
2644:
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2492:
2464:
2454:
2435:
2419:
2417:Further reading
2414:
2359:
2355:
2300:
2296:
2257:(7038): 34–57.
2239:
2235:
2212:
2208:
2163:Proc. R. Soc. B
2151:
2147:
2086:
2082:
2037:
2030:
2014:
2013:
1971:
1964:
1927:
1920:
1873:
1862:
1848:
1841:
1818:Bruhn H. 2008.
1817:
1813:
1774:
1770:
1754:
1753:
1749:
1714:
1710:
1665:
1661:
1622:(10): 1018–25.
1616:Current Biology
1608:
1604:
1551:
1547:
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1528:
1513:
1509:
1493:
1492:
1488:
1470:
1463:
1426:(5640): 58–60.
1412:
1408:
1393:
1389:
1381:
1377:
1338:
1334:
1301:(7): e1001013.
1287:
1280:
1241:
1237:
1176:
1169:
1157:
1130:
1123:
1088:
1084:
1079:
1070:
1029:genome project"
1021:
1008:
955:
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942:
940:
914:
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779:
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649:
607:
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547:
545:Lab cultivation
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363:
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129:Dictyosteliidae
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49:
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2463:
2462:External links
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2222:(6): 968–979.
2206:
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2080:
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1962:
1941:(3): 283–291.
1918:
1860:
1839:
1811:
1780:. Benchmarks.
1768:
1747:
1728:(3): 294–296.
1708:
1679:(20): 1880–5.
1659:
1602:
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1401:2012-07-29 at
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1105:NCBI Bookshelf
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827:. Meanwhile,
809:Acrasiomycetes
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750:Sentinel cells
746:
745:Sentinel cells
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648:
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606:
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452:model organism
443:
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333:sheath in its
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1820:Dictyostelium
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1783:BioTechniques
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1403:archive.today
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1027:Dictyostelium
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939:on 2017-12-08
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897:Dictyostelium
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869:
868:D. discoideum
863:
861:
860:D. discoideum
855:
853:
852:D. discoideum
849:
848:D. discoideum
845:
842:
841:D. discoideum
832:
830:
826:
823:, along with
822:
817:
816:Dictyostelium
812:
810:
806:
802:
801:Dictyostelium
792:
790:
789:
784:
783:D. discoideum
774:
772:
768:
762:
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755:
751:
742:
740:
736:
735:D. discoideum
732:
728:
724:
723:Mycobacterium
720:
716:
715:D. discoideum
709:
708:D. discoideum
704:
700:
698:
697:D. discoideum
694:
691:, as well as
690:
689:Gram-positive
686:
685:D. discoideum
682:
680:
679:D. discoideum
676:
672:
671:D. discoideum
668:
667:D. discoideum
664:
660:
659:D. discoideum
656:
655:
644:
642:
638:
634:
630:
629:D. discoideum
626:
622:
621:D. discoideum
618:
614:
613:
602:
600:
596:
592:
588:
587:Dictyostelium
584:
583:Dictyostelium
574:
571:
570:D. discoideum
566:
564:
560:
559:D. discoideum
556:
552:
551:D. discoideum
542:
540:
536:
535:Dictyostelium
531:
528:
527:D. discoideum
524:
520:
517:
513:
509:
505:
502:
501:D. discoideum
497:
495:
494:D. discoideum
490:
483:
482:D. discoideum
479:
475:
472:
467:
465:
464:gene knockout
461:
460:D. discoideum
457:
453:
449:
448:D. discoideum
439:
437:
436:D. discoideum
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432:D. discoideum
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421:
417:
416:D. discoideum
413:
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405:
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392:
391:heterothallic
387:
384:
383:D. discoideum
380:
376:
375:D. discoideum
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367:D. discoideum
358:
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344:
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332:
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311:glycoproteins
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288:D. discoideum
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269:
268:D. discoideum
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248:
247:D. discoideum
245:In the wild,
238:
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235:D. discoideum
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230:D. discoideum
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218:D. discoideum
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163:
162:Binomial name
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64:D. discoideum
60:
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43:
42:D. discoideum
38:
34:
29:
26:
22:
19:
2497:
2476:
2469:
2443:
2424:
2373:(8): e6754.
2370:
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2148:
2097:
2093:
2083:
2051:(1): 63–72.
2048:
2044:
2017:cite journal
1985:(1): 20–28.
1982:
1978:
1938:
1934:
1880:
1876:
1853:
1827:
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1814:
1790:(1): 39–41.
1787:
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966:
963:Microbiology
962:
952:
941:. Retrieved
937:the original
932:
928:
922:
911:
896:
883:
867:
864:
859:
856:
851:
847:
840:
838:
829:protostelids
815:
813:
805:polyphyletic
800:
798:
788:Burkholderia
786:
782:
780:
767:Dictyostelia
766:
763:
759:phagocytosis
753:
748:
738:
734:
730:
726:
722:
718:
714:
712:
707:
696:
684:
683:
678:
670:
666:
658:
652:
650:
637:phagocytosis
632:
628:
624:
620:
619:in humans.
610:
608:
601:conjugates.
599:streptavidin
586:
582:
580:
569:
567:
562:
558:
550:
548:
534:
532:
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431:
423:
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399:mating types
388:
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374:
366:
364:
355:
327:
308:
290:begins when
287:
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267:
254:
246:
244:
234:
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196:
195:
194:
170:
168:
152:
151:
139:
109:Dictyostelia
63:
62:A migrating
41:
24:
18:
2674:Eumycetozoa
1342:Curr. Genet
917:Raper, K.B.
825:myxomycetes
641:endocytosis
516:Thermotaxis
428:germination
395:homothallic
261:Uninucleate
2648:Categories
1828:Legionella
1824:Legionella
1295:PLOS Genet
943:2016-01-20
904:References
665:" between
633:Legionella
625:Legionella
612:Legionella
489:Chemotaxis
323:cyclic AMP
304:folic acid
284:life cycle
253:, such as
226:life cycle
214:slime mold
191:Life cycle
2664:Mycetozoa
2183:0962-8452
2122:0036-8075
1786:(paper).
1761:(paper).
1742:0002-9122
1673:Curr Biol
1579:2399-3642
1565:(1): 34.
1496:cite book
1273:205599638
1212:0036-8075
1103:(Work in
1001:232092012
985:1350-0872
880:DictyBase
874:Databases
821:Amoebozoa
725:species,
675:husbandry
663:symbiosis
647:"Farming"
523:Apoptosis
456:apoptosis
404:macrocyst
371:asexually
365:Although
331:cellulose
222:eukaryote
210:Mycetozoa
206:Amoebozoa
147:Species:
99:Amoebozoa
89:Eukaryota
2587:MycoBank
2579:11022455
2548:Fungorum
2507:Wikidata
2407:19707546
2367:PLOS ONE
2289:15875012
2228:21156570
2201:27097923
2140:17673666
2075:22801290
2067:14651611
2009:15640165
1957:19919566
1905:21248849
1855:BBC News
1806:23834384
1703:16243037
1654:16713960
1597:30701199
1399:Archived
1370:23172357
1362:24189663
1327:20617172
1265:21929567
1230:21148389
1100:The Cell
1093:(2000).
1063:12727861
993:33646931
919:(1935).
347:humidity
335:anterior
319:adhesion
296:sorocarp
251:bacteria
125:Family:
95:Phylum:
85:Domain:
2566:3212434
2513:Q134351
2398:2727795
2375:Bibcode
2348:9342353
2316:Bibcode
2280:1352341
2259:Bibcode
2192:4855374
2131:3291017
2102:Bibcode
2094:Science
1913:4333826
1885:Bibcode
1681:Bibcode
1645:4764056
1624:Bibcode
1588:6345914
1456:4160546
1428:Bibcode
1318:2895654
1221:3648785
1192:Bibcode
1184:Science
721:, many
563:E. coli
412:haploid
408:meiosis
300:mitosis
264:amoebae
135:Genus:
115:Order:
105:Class:
2631:160850
2592:263306
2553:263306
2540:197896
2450:
2431:
2405:
2395:
2346:
2336:
2287:
2277:
2251:Nature
2226:
2199:
2189:
2181:
2138:
2128:
2120:
2073:
2065:
2007:
2000:544274
1997:
1955:
1911:
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1877:Nature
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1228:
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1061:
1054:156086
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991:
983:
844:genome
835:Genome
807:class
739:Nature
729:, and
654:Nature
597:using
345:, and
292:spores
202:amoeba
180:, 1935
2618:44689
2574:IRMNG
2527:35VPX
2339:23686
2071:S2CID
1909:S2CID
1538:[
1452:S2CID
1366:S2CID
1269:S2CID
997:S2CID
969:(3).
895:from
595:MCCC1
430:of a
339:light
220:is a
178:Raper
2613:NCBI
2561:GBIF
2448:ISBN
2429:ISBN
2403:PMID
2344:PMID
2308:PNAS
2285:PMID
2224:PMID
2197:PMID
2179:ISSN
2136:PMID
2118:ISSN
2063:PMID
2023:link
2005:PMID
1953:PMID
1901:PMID
1832:ISBN
1802:PMID
1738:ISSN
1699:PMID
1650:PMID
1593:PMID
1575:ISSN
1502:link
1482:ISBN
1444:PMID
1358:PMID
1323:PMID
1261:PMID
1226:PMID
1208:ISSN
1160:ISBN
1117:ISBN
1059:PMID
989:PMID
981:ISSN
839:The
555:agar
469:The
424:mata
422:and
420:matA
343:heat
313:and
282:The
2600:NBN
2535:EoL
2522:CoL
2393:PMC
2383:doi
2334:PMC
2324:doi
2275:PMC
2267:doi
2255:435
2187:PMC
2171:doi
2167:283
2126:PMC
2110:doi
2098:317
2053:doi
1995:PMC
1987:doi
1943:doi
1893:doi
1881:469
1792:doi
1730:doi
1689:doi
1640:PMC
1632:doi
1583:PMC
1567:doi
1436:doi
1424:211
1418:".
1350:doi
1313:PMC
1303:doi
1253:doi
1216:PMC
1200:doi
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