Reticulomyxa

Last updated

Reticulomyxa
Reticulomyxa filosa - Cell body and food particle.jpg
darkfield microscopic photo of the cell body (Z) of Reticulomyxa filosa. In the lower right of the picture you see some food particles (F) (wheat germ) surrounded by the fine stands of the Reticulopodial Network (R)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Phylum: Retaria
Subphylum: Foraminifera
Class: Monothalamea
Order: Allogromiida
Family: Reticulomyxidae
Genus: Reticulomyxa
Species:
R. filosa
Binomial name
Reticulomyxa filosa
Nauss 1949

Reticulomyxa is a monospecific genus of freshwater foraminiferans. [1] The type species is the unicellular Reticulomyxa filosa. It is found in freshwater environments as well as moist environments, like decomposing matter and damp soils. The heterotrophic naked foraminiferan can feed on microbes as well has larger organisms and is able to be sustained in culture by supplemented nutrients such as wheat germ and oats. [2] [3] [4] The large, multinucleate foraminferan is characteristic for its lack of test and named for the network of connecting pseudopodia surrounding its central body mass. [2] The organism has unique bidirectional cytoplasmic streaming throughout the anastomosing pseudopodia that is some of the fastest reported organelle transport observed. [5] Reticulomyxa was first described in 1949 and is commonly used as a model organism for the unique transport of organelles throughout the cytoplasm of pseudopodia by cytoskeletal mechanisms. Only asexual reproduction of this genus has been observed in culture, but the genome possesses genes related to meiosis suggesting it is capable of sexually reproductive life stages. [6]

Contents

History and Etymology

In 1937 Reticulomyxa was isolated from a slime mold culture and observed in culture for twelve years before being officially described. It was continued to be found in damp environments in that area. This organism was initially cultured in semi aquatic conditions on damp blotting paper and in dishes of freshwater. [2] From this original isolation, samples were cultivated and observed before R.filosa was first described by Ruth Nauss in 1949 and named for its network of filose pseudopodia. Since this initial description, Reticulomyxa has been isolated from various freshwater environments as well as terrestrially. [2] [3]

The lack of test seen in Reticulomyxa is important for the history of foraminferans. Fossils of tests have been recorded from marine environments but any potentially unknown naked foraminferans would not be identified this way. The existence of Reticulomyxa is important because it suggests that there are potentially unstudied naked foraminiferans in marine ecosystems that have not been identified due to their lack of fossils. [1] The organism has been used as a model system in studies of cytoskeletal transport due to the unique fast bidirectional cytoplasmic streaming in network of pseudopodia and on the cell body surface. [5] [7]

Ecology

Reticulomyxa is found in aquatic and semi aquatic terrestrial environments. R. filosa has been isolated from lakes, moist soil, decomposing organic matter, and artificial bodies of water. [2] [3] [4] Under different conditions the plasmodium will take on various morphologies. When transferred to an aqueous environment, the plasmodia will dissociate into smaller cell bodies that reform pseudopodia. [2]

Reticulomyxa is a heterotroph that can feed on prey of a range of sizes. Previous studies have observed the ingestion of bacteria and other protists, as well as large aquatic zooplankton. [2] [4] The vegetative plasmodium will stay in one location while eating until surrounding food sources have been depleted. Once devoid of food, the cell will excrete waste from the protoplasm and move to a new location. [2] Feeding occurs through the pseudopodia of the cell, filose pseudopodia engulf and phagocytize prey. [5] Prey is encapsulated in food vacuoles and are then transported back to the central body via the cytoskeletal machinery. [2]

Description

Morphology

Reticulomyxa has a plasmodial morphology, including a central body surrounded by filose pseudopodia, which have a diameter of approximately 50 μm. [2] [5] This organism lacks a test that is a defining characteristic of foraminiferans. This test is suspected to have been lost throughout its evolution due to the adaptation to freshwater environments. [8] When in the vegetative stage, the central body of Reticulomyxa is round and has many pseudopodia extending outwards forming an array or network-like plasmodia which facilitate locomotion and feeding. These pseudopodia consistently change shape and length due to changing environmental and nutrient conditions. In aquatic conditions the pseudopodia branch out in all directions whereas when in moist condition the pseudopodia contract into the central body. [2] The cytoplasm of this organism is thick and streams bidirectionally through the veins. [2] [3] Cytoplasmic streaming allowing for the movement of organelles and vacuoles throughout the network can reach speeds of 25 μm/s. [9] Organelle movement is facilitated through the cytoskeleton, which consists of actin and microtubules. Organelles are attached to the microtubules of the cytoskeleton and movement is due to the sliding of microtubules along each other in both directions. [10] This giant freshwater foraminferan contains many haploid nuclei, approximately 5 μm in diameter. [6]

Small section of the Reticulopodial Network of Reticulomyxa filosa: Fine reticulopods alter with small Lamellipodial sections. (Phase contrast micrograph; bar = 50 micrometers) Reticulomyxa filosa - Reticulopodial network.jpg
Small section of the Reticulopodial Network of Reticulomyxa filosa: Fine reticulopods alter with small Lamellipodial sections. (Phase contrast micrograph; bar = 50 micrometers)

Cysts produced by Reticulomyxa can be covered or uncovered. Covered cysts are enclosed by two layers, with the outer mucus layer being much thicker. Uncovered cysts do not have the thick outer layer as seen in covered cysts and have irregular sizes. [4]

Life cycle

Plasmodia of this genera have only been seen to undergo mitosis in culture conditions. Asexual reproduction through multiple fission, where the nuclei will undergo many rounds of mitotic division before the cytoplasm separates. [2] In Reticulomyxa, mitosis is closed, the nuclear membrane remains intact during spindle formation and chromosome separation. [6] Under culture conditions, division was initiated following the movement to a new location. Division starts at the anterior end of the cell and produces three daughter cells. [2]

The Reticulomyxa plasmodium may encyst for dispersal or in response to adverse environmental conditions. [4] Cysts with and without coverings have been produced by Reticulomyxa. Cysts covered with a thick envelope can withstand harsher conditions and disperse by wind as well as water. Covered cells were only observed to form during summer months in culture conditions. [4] [11] Uncovered cysts are smooth and have irregular sizes. These uncovered resting cysts were produced in culture conditions in response unfavorable environmental conditions like low temperatures or when nutrient availability was low. [2] [4] [11]

Genetics

The genome of Reticulomyxa is repetitive and approximately 320 Mbp in size. The genome contains genes for flagellar components, despite no flagellated form observed. Also, genes coding for proteins associated with meiosis are present in the Reticulomyxa genome but are not actively transcribed. The presence of flagella and meiosis related genes suggests that there is a possibility of sexual reproduction and gamete production in this genus. [6]

Related Research Articles

<span class="mw-page-title-main">Actinophryid</span> Order of heliozoan protists

The actinophryids are an order of heliozoa, a polyphyletic array of stramenopiles, having a close relationship with pedinellids and Ciliophrys. They are common in fresh water and occasionally found in marine and soil habitats. Actinophryids are unicellular and roughly spherical in shape, with many axopodia that radiate outward from the cell body. Axopodia are a type of pseudopodia that are supported by hundreds of microtubules arranged in interlocking spirals and forming a needle-like internal structure or axoneme. Small granules, extrusomes, that lie under the membrane of the body and axopodia capture flagellates, ciliates and small metazoa that make contact with the arms.

<span class="mw-page-title-main">Apicomplexa</span> Phylum of parasitic alveolates

The Apicomplexa are a large phylum of mainly parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. The organelle is an adaptation that the apicomplexan applies in penetration of a host cell.

<span class="mw-page-title-main">Cell (biology)</span> Basic unit of many life forms

The cell is the basic structural and functional unit of all forms of life. Every cell consists of cytoplasm enclosed within a membrane, and contains many macromolecules such as proteins, DNA and RNA, as well as many small molecules of nutrients and metabolites. The term comes from the Latin word cellula meaning 'small room'.

<span class="mw-page-title-main">Pseudopodia</span> False leg found on slime molds, archaea, protozoans, leukocytes and certain bacteria

A pseudopod or pseudopodium is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filaments and may also contain microtubules and intermediate filaments. Pseudopods are used for motility and ingestion. They are often found in amoebas.

<span class="mw-page-title-main">Cytoskeleton</span> Network of filamentous proteins that forms the internal framework of cells

The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is composed of similar proteins in the various organisms. It is composed of three main components:microfilaments, intermediate filaments, and microtubules, and these are all capable of rapid growth or disassembly depending on the cell's requirements.

<span class="mw-page-title-main">Heliomonadida</span> Order of single-celled organisms

The Heliomonadida are a small group of heliozoan amoeboids that are unusual in possessing flagella throughout their life cycle.

<span class="mw-page-title-main">Foraminifera</span> Phylum of amoeboid protists

Foraminifera are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly an external shell of diverse forms and materials. Tests of chitin are believed to be the most primitive type. Most foraminifera are marine, the majority of which live on or within the seafloor sediment, while a smaller number float in the water column at various depths, which belong to the suborder Globigerinina. Fewer are known from freshwater or brackish conditions, and some very few (nonaquatic) soil species have been identified through molecular analysis of small subunit ribosomal DNA.

<i>Naegleria</i> Genus of protists

Naegleria is a free living amoebae protist genus consisting of 47 described species often found in warm aquatic environments as well as soil habitats worldwide. It has three life cycle forms: the amoeboid stage, the cyst stage, and the flagellated stage, and has been routinely studied for its ease in change from amoeboid to flagellated stages. The Naegleria genera became famous when Naegleria fowleri, a human pathogenic strain and the causative agent of primary amoebic meningoencephalitis (PAM), was discovered in 1965. Most species in the genus, however, are nonpathogenic, meaning they do not cause disease.

<span class="mw-page-title-main">Axoneme</span>

An axoneme, also called an axial filament is the microtubule-based cytoskeletal structure that forms the core of a cilium or flagellum. Cilia and flagella are found on many cells, organisms, and microorganisms, to provide motility. The axoneme serves as the "skeleton" of these organelles, both giving support to the structure and, in some cases, the ability to bend. Though distinctions of function and length may be made between cilia and flagella, the internal structure of the axoneme is common to both.

<span class="mw-page-title-main">Molecular motor</span> Biological molecular machines

Molecular motors are natural (biological) or artificial molecular machines that are the essential agents of movement in living organisms. In general terms, a motor is a device that consumes energy in one form and converts it into motion or mechanical work; for example, many protein-based molecular motors harness the chemical free energy released by the hydrolysis of ATP in order to perform mechanical work. In terms of energetic efficiency, this type of motor can be superior to currently available man-made motors. One important difference between molecular motors and macroscopic motors is that molecular motors operate in the thermal bath, an environment in which the fluctuations due to thermal noise are significant.

<span class="mw-page-title-main">Growth cone</span> Large actin extension of a developing neurite seeking its synaptic target

A growth cone is a large actin-supported extension of a developing or regenerating neurite seeking its synaptic target. It is the growth cone that drives axon growth. Their existence was originally proposed by Spanish histologist Santiago Ramón y Cajal based upon stationary images he observed under the microscope. He first described the growth cone based on fixed cells as "a concentration of protoplasm of conical form, endowed with amoeboid movements". Growth cones are situated on the tips of neurites, either dendrites or axons, of the nerve cell. The sensory, motor, integrative, and adaptive functions of growing axons and dendrites are all contained within this specialized structure.

<i>Physarum polycephalum</i> Species of slime mold, model organism

Physarum polycephalum, an acellular slime mold or myxomycete popularly known as "the blob", is a protist with diverse cellular forms and broad geographic distribution. The “acellular” moniker derives from the plasmodial stage of the life cycle: the plasmodium is a bright yellow macroscopic multinucleate coenocyte shaped in a network of interlaced tubes. This stage of the life cycle, along with its preference for damp shady habitats, likely contributed to the original mischaracterization of the organism as a fungus. P. polycephalum is used as a model organism for research into motility, cellular differentiation, chemotaxis, cellular compatibility, and the cell cycle.

<i>Amoeba</i> (genus) Genus of Protozoa

Amoeba is a genus of single-celled amoeboids in the family Amoebidae. The type species of the genus is Amoeba proteus, a common freshwater organism, widely studied in classrooms and laboratories.

<i>Phacus</i> Genus of algae

Phacus is a genus of unicellular excavates, of the phylum Euglenozoa, characterized by its flat, leaf-shaped structure, and rigid cytoskeleton known as a pellicle. These eukaryotes are mostly green in colour, and have a single flagellum that extends the length of their body. They are morphologically very flat, rigid, leaf-shaped, and contain many small discoid chloroplasts.

<span class="mw-page-title-main">Vampyrellida</span> Order of single-celled organisms

The vampyrellids, colloquially known as vampire amoebae, are a group of free-living predatory amoebae classified as part of the lineage Endomyxa. They are distinguished from other groups of amoebae by their irregular cell shape with propensity to fuse and split like plasmodial organisms, and their life cycle with a digestive cyst stage that digests the gathered food. They appear worldwide in marine, brackish, freshwater and soil habitats. They are important predators of an enormous variety of microscopic organisms, from algae to fungi and animals. They are also known as aconchulinid amoebae.

<span class="mw-page-title-main">Intracellular transport</span> Directed movement of vesicles and substances within a cell

Intracellular transport is the movement of vesicles and substances within a cell. Intracellular transport is required for maintaining homeostasis within the cell by responding to physiological signals. Proteins synthesized in the cytosol are distributed to their respective organelles, according to their specific amino acid’s sorting sequence. Eukaryotic cells transport packets of components to particular intracellular locations by attaching them to molecular motors that haul them along microtubules and actin filaments. Since intracellular transport heavily relies on microtubules for movement, the components of the cytoskeleton play a vital role in trafficking vesicles between organelles and the plasma membrane by providing mechanical support. Through this pathway, it is possible to facilitate the movement of essential molecules such as membrane‐bounded vesicles and organelles, mRNA, and chromosomes.

<i>Vampyrella</i> Genus of single-celled organisms

Vampyrella is a genus of amoebae belonging to the vampyrellid cercozoans usually ranging from 30-60 µm. Members of the genus alternate between two life stages: a free-living trophozoite stage and a cyst stage in which mitosis occurs. This taxon has received a great deal of attention due to their peculiar feeding behaviour of perforating the cell wall of algal cells and drawing out the contents for nourishment.

Neocercomonas is a protist genus of the order Cercomonadida. It consists of single-celled bacteriophagous organisms that usually live on or nearby terrestrial plants, both above and belowground. Species are biflagellate and may grow up to 60 micrometers long, with a trailing tail-like mass of protoplasm at their posterior end and a pair of roots connecting their posterior flagellum to the cytoskeleton.

<i>Vampyrella lateritia</i> Species of protist

Vampyrella lateritia is a freshwater species of predatory amoebae that feeds on species of algae and is known for its specialized feeding strategy of removing, digesting, and ingesting the cellular contents of its prey. It is the type species of the genus Vampyrella and has been identified in numerous locations around the world including Brazil, Germany, and the eastern United States. Along with Vampyrella pendula, its genome was sequenced in 2012.

<i>Aphelidium tribonemae</i>

Aphelidium tribonemae is a species within the Aphelid group. Their classification in the kingdom Fungi is a subject of controversy. Some argue for the classification of aphelids as ‘fungal animals', and for a period of time in the 1950s, aphids were classified as protists due to their amoeboid stage. Recently, molecular phylogenetics placed the aphelids within Opisthosporidia, a super phylum within Opisthokonta. Aphelids have posterior uniflagellate zoospores which place them as Opisthokonts. They are an early diverging lineage in Kingdom Fungi. While the aphelid group only contains three genera, it spans many both freshwater and marine ecosystems.

References

  1. 1 2 PAWLOWSKI, JAN; BOLIVAR, IGNACIO; FAHRNI, JOSE F.; VARGAS, COLOMBAN DE; BOWSER, SAMUEL S. (1999-11-01). "Molecular Evidence That Reticulomyxa Filosa Is A Freshwater Naked Foraminifer". The Journal of Eukaryotic Microbiology. 46 (6): 612–617. doi:10.1111/j.1550-7408.1999.tb05137.x. ISSN   1550-7408. PMID   10568034. S2CID   36497475.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Nauss, Ruth N. (1949). "Reticulomyxa filosa Gen. Et Sp. Nov., A New Primitive Plasmodium". Bulletin of the Torrey Botanical Club. 76 (3): 161–173. doi:10.2307/2482203. JSTOR   2482203.
  3. 1 2 3 4 Koonce, M. P.; Euteneuer, U.; Schliwa, M. (1986). "Reticulomyxa: a new model system of intracellular transport". Journal of Cell Science. Supplement. 5: 145–159. doi: 10.1242/jcs.1986.supplement_5.9 . ISSN   0269-3518. PMID   3477550.
  4. 1 2 3 4 5 6 7 Gothe, G; Bohm, K. J.; Unger, E (1997). "Different resting stages of the plasmodial rhizopod Reticulomyxa filosa". Acta Protozoologica. 36: 23–29.
  5. 1 2 3 4 Euteneur, Ursula; McDonald, Kent L; Koonce, Michael P; Schliwa, Manfred (1986). "Intracellular-transport in Reticulomyxa". Annals of the New York Academy of Sciences. 466 (1): 936–939. Bibcode:1986NYASA.466..936E. doi:10.1111/j.1749-6632.1986.tb38479.x. S2CID   84102411.
  6. 1 2 3 4 Glöckner, Gernot; Hülsmann, Norbert; Schleicher, Michael; Noegel, Angelika A.; Eichinger, Ludwig; Gallinger, Christoph; Pawlowski, Jan; Sierra, Roberto; Euteneuer, Ursula (2014). "The Genome of the Foraminiferan Reticulomyxa filosa". Current Biology. 24 (1): 11–18. doi: 10.1016/j.cub.2013.11.027 . PMID   24332546.
  7. Orokos, D. D.; Bowser, S. S.; Travis, J. L. (1997). "Reactivation of cell surface transport in Reticulomyxa". Cell Motility and the Cytoskeleton. 37 (2): 139–148. doi:10.1002/(SICI)1097-0169(1997)37:2<139::AID-CM6>3.0.CO;2-3. ISSN   0886-1544. PMID   9186011.
  8. Pawlowski, Jan; Holzmann, Maria (2002). "Molecular phylogeny of Foraminifera a review". European Journal of Protistology. 38 (1): 1–10. doi:10.1078/0932-4739-00857.
  9. Kube-Granderath, Eckhard; Schliwa, Manfred (1998). "Unusual Tubulins for Unusual Cells". Protist. 149 (2): 123–126. doi: 10.1016/s1434-4610(98)70016-1 . PMID   23196161.
  10. Orokos, D. D.; Cole, R. W.; Travis, J. L. (December 2000). "Organelles are transported on sliding microtubules in Reticulomyxa". Cell Motility and the Cytoskeleton. 47 (4): 296–306. doi:10.1002/1097-0169(200012)47:4<296::AID-CM4>3.0.CO;2-4. ISSN   0886-1544. PMID   11093250.
  11. 1 2 Ross, Benjamin J.; Hallock, Pamela (2016-10-01). "Dormancy in the Foraminifera: A Review". Journal of Foraminiferal Research. 46 (4): 358–368. doi:10.2113/gsjfr.46.4.358. ISSN   0096-1191.

Further reading