Phytomonas

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Phytomonas
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Phylum: Euglenozoa
Class: Kinetoplastea
Order: Trypanosomatida
Family: Trypanosomatidae
Genus: Phytomonas
Donovan, 1909 [1]
Species

See text.

Phytomonas is a genus of trypanosomatids that infect plant species. Initially described using existing genera in the family Trypanosomatidae, such as Trypanosoma or Leishmania, the nomenclature of Phytomonas was proposed in 1909 in light of their distinct hosts and morphology. [2] [3] When the term was originally coined, no strict criterion was followed, and the term was adopted by the scientific community to describe flagellate protozoa in plants as a matter of convenience. [3] Members of the taxon are globally distributed and have been discovered in members of over 24 plant families. [4] Of these 24, the two main families that are infected by Phytomonas are Euphorbiaceae and Asclepiadiacae. These protists have been found in hosts between 50° latitude North and South, and thus they can be found on all continents save for Antarctica. [3]

Contents

Phytomonas is believed to have arisen from a single monoxenous lineage of insect parasitizing trypanosomatids some 400 million years ago. [5] After this divergence, a heteroxenous lifestyle was developed, and most Phytomonas species are transferred between plant hosts by insect vectors in the Heteroptera suborder as a form of dixenous parasitism. [3] [5] Species with considerable economic impact include Phytomonas leptovasorum and Phytomonas staheli¸ the causative agents of phloem necrosis in coffee and wilt of coconut and oil palms.

Etymology

First proposed by Donovan in 1909, the term Phytomonas describes the relationship the genus has with plants compared to closely related organisms in Trypanosomatida. [2] Having been discovered as parasites of plants as opposed to mammals, the term phyto- was used to describe this relationship. In English, the term phyto- means plant, and comes from the Greek term for plant, phuton. The suffix -monas means 'unicellular organism' or 'single unit', and is now commonly used in the field of microbiology.

History of knowledge

The organisms now known as Phytomonas were first reported in 1909 by Dr. A. Lafont, after having discovered them in the latex of the spurge plant, Euphorbia pilulifera . [6] He named the organism Leptomonas davidi , after his lab technician David, the individual who first observed it. That same year, Donovan [2] confirmed this by also observing the organisms in the latex of Euphorbia piluifera in Mauritius and Chennai. Because these particular trypanosomatids differed distinctly from those that were known to be parasitic to animals, Donovan proposed that a new genus of trypanosomatids, Phytomonas, had to be created. [2]

Since 1909, trypanosomatids were reported to infect plants from around the planet, but initially, there was little scientific interest in the genus, aside from possible relations to disease-causing organisms in animals and humans. [7] [4] In 1931, Phytomonas were discovered in Coffea liberica plants, and was found to be the cause of coffee phloem necrosis. [6] [8] In following years, more parasites were observed in plants with stronger economic value, such as tomatoes ( Solanum lycopersicum ) [6] The infection of economically valuable crops attracted the attention of several research groups, but interest was again lost due to the inability to cultivate cultures in vitro. [7] [6] In the 1970s, scientific interest was revitalized due to increasingly devastating problems caused by the parasites in even more economically significant crops, namely coconut ( Cocos nucifera ) and oil palm ( Elaeis guineensis ). [7] [6] Finally, in 1982, Dollet successfully cultivated trypanosomatids in vitro, which allowed for the isolation of Phytomonas in 24 different plant families from across the globe. [7]

While there has been recent genetic work done on Phytomonas species, genome level analysis is limited compared to the genome data available for disease-causing trypanosomatids in animals and humans, especially Trypanosoma cruzi , Trypanosoma brucei , and Leishmania major . [9] In 2015, Phytomonas nordicus was found to be a part of the generally heteroxenous genus Phytomonas, despite it being a monoxenous parasite of the predatory bug Troilus luridus . [10] The species was described earlier in 1993. Having a monoxenous lifestyle, the species was not observed in plants, but was categorized as Phytomonas due to features that were similar to other phytomonads, namely long twisted promastigotes and flagellated stages in the salivary glands of bugs. Using molecular phylogenetic analyses, Frolov determined that the species was indeed a part of Phytomonas, despite not being a parasite of plants. [10] In most recent news, a new dixenous species, Phytomonas oxycareni n. sp. was discovered and described in 2017, after having been obtained from the salivary glands of the true bug Oxycarenus lavaterae . [5]

Habitat and ecology

As the nomenclature Phytomonas suggests, this genus consists of trypanosomatids that infect plants. Though these organisms infect plant species from around the world, the two main plant families that contain the most Phytomonas hosts are Euphorbiaceae and Asclepiadiacae . [3] Within host plants, Phytomonas species have been observed in multiple tissue types, including phloem, seeds, fruits, flowers, and latex ducts. [4] Notable examples include Phytomonas serpens in tomato, P. staheli in coconut and oil palm, and P. leptovasorum in coffee. Most species do not appear to be pathogenic to their hosts or otherwise have any deleterious effects. [9] Because the term Phytomonas was not founded on any strict criterion other than being flagellated protists in plants, the nomenclature fails to reflect the wide range of lifestyles of the various species in the genus. [3] [9]

Overwhelmingly, the majority of trypanosomatid species are spread via insect vectors. [4] This is true for Phytomonas as well, and not very long after the categorization of the genus, it was found that the organisms could be transported from an infected plant to an uninfected plant using phytophagous hemipterans. [4] Subsequent research in following years showed that Phytomonas species could be spread between plant hosts by a broad range of insect species. However, this simple view of the relationship between parasite and hosts is complicated by nomenclature and limited research. [4] The current understanding is that the primary insect vectors are members of the Heteroptera suborder. [5] In fact, the two main taxa with species identified as vectors are the Heteropteran families of Lygaeidae and Coreidae . [3] In most cases, the organisms are transferred from the insect to the plant during feeding, as the parasites are present in the salivary glands. [3]

Description of the organism

Morphology and anatomy

As members of the family Trypanosomatidae, Phytomonas have structures that are characteristic of the family, including the flagellum-associated kinetoplast, subpellicular microtubules, the paraxial rod, and glycosomes. [6] Within a host plant, Phytomonas exhibits a fusiform structure twisted 2-5 times along the longitudinal axis. [3] Within the plant, the organisms can be in several flagellated stages: mostly promastigote with some paramastigotes in the phloem and lacticiferous tubes, and amastigote form in the latex. [6] However, most of the species are mainly observed in the promastigote form, with an elongated body and a single 10-15 μm long flagellum emerging from the anterior flagellar pocket. [3] [4] [6] This main form of their morphology is said to be consistent with insect parasites in their sister group Leishmania. [4] Species range from 10 to 20 μm in length and have widths close to 1.5 μm. [3] [6] It appears that reproduction of cells occurs only in the promastigote stage. [3] When viewed as live samples under the light microscope, Phytomonas can be seen to be incredibly active. [3] Different species that infect different host plants have differences in external morphology, for example, oil palm (Elaeis guineensis)-infecting Phytomonas exhibit fewer twists than those in coconut (Cocos nucifera). [6]

Like other trypanosomatids, the cell surface of Phytomonas species can be divided into the three regions of body surface, flagellar surface, and the flagellar pocket. [4] Also like other trypanosomatids, Phytomonas does not have a cell wall, but instead are protected from host responses and environmental conditions by membrane-anchored proteins and glycoinositol phospholipids. [4] The pellicular cell membrane is also lined with microtubules that run along the longitudinal axis of the organism, with a single row of four microtubules in the flagellar pocket. [3] A paraxial rod also runs parallel to the axoneme of the single flagellum on one side, giving the flagella increased thickness, robustness, and strength. [3]

As mentioned earlier, these organisms also have glycosomes, which are specialized peroxisomes. Depending on the species examined, these glycosomes may take the form of two rows separated by filamentous fibres. [6] Some isolates of the genus also have a contractile vacuole located at the anterior end, near the flagellar pocket. [6] This vacuole varies in size, but can be as large as 2 μm. It has been found that the endoplasmic reticulum (ER) of some species form subpellicular sheets that run parallel to the longitudinal axis. [6] Depending on the species and isolates in question, the endoplasmic reticulum (ER) may also have ribosomes in paracrystalline array and incredibly thin intracisternal space. [6] The cytoplasm is considered to be ribosome-rich. [3] In close association with the flagellum is the kinetoplast. The kinetoplast DNA networks of Phyotomonas species have been found to vary in isolates extracted from various insect and plant hosts, ranging from a loose appearance to compact networks reminiscent of those of other trypanosomatids. [6]

Life cycle

According to Dollet, reproduction occurs during the promastigote stage. [3] Elongation of the kinetoplast occurs first, and is followed by splitting of the anterior end of the cell. Because the single flagellum morphology of trypanosomes arose from the loss of one flagellum in the flagellar pocket, this splitting results in one piece of anterior cell having one flagellum and another that does not. [3] A new flagellum will later grow on the part that lacked a flagellum. [3] Further longitudinal division distributes the nuclei between the forming daughter cells.

Reproduction and multiplication of Phytomonas generally stops after entering the insect host. [3] Within the first week inside the host, the parasites elongate to form "giant" versions of themselves inside the pylorus of the host. After 12 days, cells migrate to the salivary glands via the haemolymph. [3] It is here, in the salivary glands, where multiplication resumes again, producing regular sized forms for infection of plant hosts. Depending on species and isolate, these promastigote forms may or may not be attached to the cell walls of the salivary glands with their flagella. [3] For example, in Phytomonas serpens, cells do attach to the interior of salivary glands, which indicates a true developmental stage within the insect host in some species. [4]

Agricultural importance

Compared to dangerous parasites of humans and animals, Phytomonas received little initial interest. Species of this genus have been found to infect many different host plants, and a majority of them do not seem to have negative effects on their hosts. However, several species are the cause of damaging diseases in many economically significant crop plants.

Phloem necrosis of coffee (Coffea liberica)

Described by Stahel in 1917, this disease is characterized by deposition of callose in the sieve tubes and necrosis. [3] Trypanosomatids were not discovered to be the cause until Stahel observed them in the sieve tubes of Coffea liberica plants in 1931. [6] It is now known to be caused by Phytomonas leptovasorum Stahel. [8] The insect vector of the parasite has not been confirmed, but Stahel mainly suspected Lincus spathuliger as a culprit. [3] Vermeulen also suspected a Hemipteran insect, due to the presence of flagellate protists in the midgut of certain bugs often found on coffee plant roots. [8] While in the plant host, these parasites inhabit only the phloem, a characteristic that it shares with P. staheli. [4] Despite being first characterized in C. liberica, the disease can also be caused by P. leptovasorum in C. arabica, C. excelsa, C. stenophylla, and C. abeocutae. [3] The disease caused by this phytomonad can be either acute or chronic, with the chronic form being much more frequent (95%). The two forms differ in the amount of foliage loss and killing time, but both cases involve the browning and death of the roots. [3]

Hartrot of coconut (Cocus nucifera)

Also known as fatal wilt, bronze leaf wilt, lethal yellowing, and Coronie wilt, this disease of coconut is caused by Phytomonas staheli. [3] [4] This is an acute wilt, and begins at the leaves of the tree, moving down the plant towards the root. [4] As such, the first symptoms to appear are yellowing of the leaf tips, followed by yellowing of new leaves. As unripe fruits begin to fall prematurely, rotting of the root begins. [3] After ten weeks, the plant dies, and a foul odor arises due to secondary infection by bacteria and fungi. [3] At first, it was believed that the disease was isolated to the east coast of South America, but in fact occurred in Colombia and Ecuador as well. [3]

Marchitez of oil palm (Elaeis guineensis)

Phytomonas staheli also causes disease in oil palm, namely marchitez sorpresiva (sudden wilt) and slow wilt. [4] While in the plant host, P. staheli remains in the phloem, a trait it shares with P. leptovasorum. [4] As in hartrot of coconut, disease symptoms first appear as yellowing of leaf tips. However, in oil palm, the lower leaves are targeted first, with upper leaves being invaded next. [3] Roots also deteriorate, starting from the growing tips and moving toward primary roots. As in coconut hartrot, fruits are also lost. Within 3–5 weeks, the plant dies and leaves become dried masses of grey. [3] P. staheli can spread between trees quickly, and can infect up to 30 trees within a few weeks. The disease has great impact across the north of Latin America, but can be partially controlled with insecticide application. [3] This is likely by virtue of killing insect vectors which are suspected to be known or new species in the genus Lincus. [3]

Medical importance

Aside from the harmful impact these organisms can have, Phytomonas species can also be useful as parallel models for the study of dangerous diseases caused by organisms in other infections trypanosomatid genera. Trypanosoma cruzi is the causative agent of Chagas' disease, and as a member of the family Trypanosomatidae, is related to organisms of the genus Phytomonas. [11] In a 2015 study, Phytomonas Jma was tested as a model for the expression of heterologous proteins in the dangerous T. cruzi. [12] It was found that Phytomonas was able to express GFP levels similar to that of T. cruzi, and it was concluded that organisms in the genus could be used as human-safe models for functional expression of trypanosomatid proteins. [12]

Research was also done on the practical uses of antigen similarities between P. serpens and trypanosomatid pathogens. It was found that T. cruzi shares some antigens with P. serpens, and that these antigens could be recognized by rabbit and mouse sera. [11] Immunization of mice with P. serpens induced a partial immune response against T. cruzi infection. [13] Though more research is required, the phytomonads could potentially be used as vaccine agents to prime defense responses to T. cruzi antigens in order to interfere with the development of Chagas' disease in humans. [11] [13]

List of better-known described species

Related Research Articles

<span class="mw-page-title-main">Chagas disease</span> Mammal parasitic disease

Chagas disease, also known as American trypanosomiasis, is a tropical parasitic disease caused by Trypanosoma cruzi. It is spread mostly by insects in the subfamily Triatominae, known as "kissing bugs". The symptoms change over the course of the infection. In the early stage, symptoms are typically either not present or mild, and may include fever, swollen lymph nodes, headaches, or swelling at the site of the bite. After four to eight weeks, untreated individuals enter the chronic phase of disease, which in most cases does not result in further symptoms. Up to 45% of people with chronic infections develop heart disease 10–30 years after the initial illness, which can lead to heart failure. Digestive complications, including an enlarged esophagus or an enlarged colon, may also occur in up to 21% of people, and up to 10% of people may experience nerve damage.

<i>Leishmania</i> Genus of parasitic flagellate protist

Leishmania is a parasitic protozoan, a single-celled organism of the genus Leishmania that is responsible for the disease leishmaniasis. They are spread by sandflies of the genus Phlebotomus in the Old World, and of the genus Lutzomyia in the New World. At least 93 sandfly species are proven or probable vectors worldwide. Their primary hosts are vertebrates; Leishmania commonly infects hyraxes, canids, rodents, and humans.

<span class="mw-page-title-main">Trypanosomatida</span> Flagellate kinetoplastid excavate order

Trypanosomatida is a group of kinetoplastid unicellular organisms distinguished by having only a single flagellum. The name is derived from the Greek trypano (borer) and soma (body) because of the corkscrew-like motion of some trypanosomatid species. All members are exclusively parasitic, found primarily in insects. A few genera have life-cycles involving a secondary host, which may be a vertebrate, invertebrate or plant. These include several species that cause major diseases in humans. Some trypanosomatida are intracellular parasites, with the important exception of Trypanosoma brucei.

<span class="mw-page-title-main">Kinetoplastida</span> Flagellated protists belonging to the phylum Euglenozoa

Kinetoplastida is a group of flagellated protists belonging to the phylum Euglenozoa, and characterised by the presence of a distinctive organelle called the kinetoplast, a granule containing a large mass of DNA. The group includes a number of parasites responsible for serious diseases in humans and other animals, as well as various forms found in soil and aquatic environments. The organisms are commonly referred to as "kinetoplastids" or "kinetoplasts".

<i>Trypanosoma</i> Genus of parasitic flagellate protist in the Kinetoplastea class

Trypanosoma is a genus of kinetoplastids, a monophyletic group of unicellular parasitic flagellate protozoa. Trypanosoma is part of the phylum Sarcomastigophora. The name is derived from the Greek trypano- (borer) and soma (body) because of their corkscrew-like motion. Most trypanosomes are heteroxenous and most are transmitted via a vector. The majority of species are transmitted by blood-feeding invertebrates, but there are different mechanisms among the varying species. Trypanosoma equiperdum is spread between horses and other equine species by sexual contact. They are generally found in the intestine of their invertebrate host, but normally occupy the bloodstream or an intracellular environment in the vertebrate host.

<span class="mw-page-title-main">Carlos Chagas</span> Brazilian doctor and scientist (1879–1934)

Carlos Justiniano Ribeiro Chagas, or Carlos Chagas, was a Brazilian sanitary physician, scientist, and microbiologist who worked as a clinician and researcher. Most well known for the discovery of an eponymous protozoal infection called Chagas disease, also called American trypanosomiasis, he also discovered the causative fungi of the pneumocystis pneumonia. He described the two pathogens in 1909, while he was working at the Oswaldo Cruz Institute in Rio de Janeiro, and named the former Trypanosoma cruzi to honour his friend Oswaldo Cruz.

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

A cytostome or cell mouth is a part of a cell specialized for phagocytosis, usually in the form of a microtubule-supported funnel or groove. Food is directed into the cytostome, and sealed into vacuoles. Only certain groups of protozoa, such as the Ciliophora and Excavata, have cytostomes. An example is Balantidium coli, a ciliate. In other protozoa, and in cells from multicellular organisms, phagocytosis takes place at any point on the cell or feeding takes place by absorption.

<i>Trypanosoma brucei</i> Species of protozoan parasite

Trypanosoma brucei is a species of parasitic kinetoplastid belonging to the genus Trypanosoma that is present in sub-Saharan Africa. Unlike other protozoan parasites that normally infect blood and tissue cells, it is exclusively extracellular and inhabits the blood plasma and body fluids. It causes deadly vector-borne diseases: African trypanosomiasis or sleeping sickness in humans, and animal trypanosomiasis or nagana in cattle and horses. It is a species complex grouped into three subspecies: T. b. brucei, T. b. gambiense and T. b. rhodesiense. The first is a parasite of non-human mammals and causes nagana, while the latter two are zoonotic infecting both humans and animals and cause African trypanosomiasis.

<i>Retortamonas</i> Unicellular organism

Retortamonas is a genus of flagellated excavates. It is one of only two genera belonging to the family Retortamonadidae along with the genus Chilomastix. The genus parasitizes a large range of hosts including humans. Species within this genus are considered harmless commensals which reside in the intestine of their host. The wide host diversity is a useful factor given that species are distinguished based on their host rather than morphology. This is because all species share similar morphology, which would present challenges when trying to make classifications based on structural anatomy. Although Retortamonas currently includes over 25 known species, it is possible that some defined species are synonymous, given that such overlapping species have been discovered in the past. Further efforts into learning about this genus must be done such as cross-transmission testing as well as biochemical and genetic studies. One of the most well-known species within this genus is Retortamonas intestinalis, a human parasite that lives in the large intestine of humans.

<i>Crithidia</i> Genus of parasitic flagellate protist in the Kinetoplastea class

Crithidia is a genus of trypanosomatid Euglenozoa. They are parasites that exclusively parasitise arthropods, mainly insects. They pass from host to host as cysts in infective faeces and typically, the parasites develop in the digestive tracts of insects and interact with the intestinal epithelium using their flagellum. They display very low host-specificity and a single parasite can infect a large range of invertebrate hosts. At different points in its life-cycle, it passes through amastigote, promastigote, and epimastigote phases; the last is particularly characteristic, and similar stages in other trypanosomes are often called crithidial.

Paratransgenesis is a technique that attempts to eliminate a pathogen from vector populations through transgenesis of a symbiont of the vector. The goal of this technique is to control vector-borne diseases. The first step is to identify proteins that prevent the vector species from transmitting the pathogen. The genes coding for these proteins are then introduced into the symbiont, so that they can be expressed in the vector. The final step in the strategy is to introduce these transgenic symbionts into vector populations in the wild. One use of this technique is to prevent mortality for humans from insect-borne diseases. Preventive methods and current controls against vector-borne diseases depend on insecticides, even though some mosquito breeds may be resistant to them. There are other ways to fully eliminate them. “Paratransgenesis focuses on utilizing genetically modified insect symbionts to express molecules within the vector that are deleterious to pathogens they transmit.” The acidic bacteria Asaia symbionts are beneficial in the normal development of mosquito larvae; however, it is unknown what Asais symbionts do to adult mosquitoes.

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

A kinetoplast is a network of circular DNA inside a mitochondrion that contains many copies of the mitochondrial genome. The most common kinetoplast structure is a disk, but they have been observed in other arrangements. Kinetoplasts are only found in Excavata of the class Kinetoplastida. The variation in the structures of kinetoplasts may reflect phylogenic relationships between kinetoplastids. A kinetoplast is usually adjacent to the organism's flagellar basal body, suggesting that it is bound to some components of the cytoskeleton. In Trypanosoma brucei this cytoskeletal connection is called the tripartite attachment complex and includes the protein p166.

<span class="mw-page-title-main">Beet leafhopper</span> Species of insect

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Crithidia fasciculata is a species of parasitic excavates. C. fasciculata, like other species of Crithidia have a single host life cycle with insect host, in the case of C. fasciculata this is the mosquito. C. fasciculata have low host species specificity and can infect many species of mosquito.

<span class="mw-page-title-main">Disease vector</span> Agent that carries and transmits an infectious pathogen into another living organism

In epidemiology, a disease vector is any living agent that carries and transmits an infectious pathogen to another living organism; agents regarded as vectors are organisms, such as parasites or microbes. The first major discovery of a disease vector came from Ronald Ross in 1897, who discovered the malaria pathogen when he dissected a mosquito.

<span class="mw-page-title-main">Protozoan infection</span> Parasitic disease caused by a protozoan

Protozoan infections are parasitic diseases caused by organisms formerly classified in the kingdom Protozoa. They are usually contracted by either an insect vector or by contact with an infected substance or surface and include organisms that are now classified in the supergroups Excavata, Amoebozoa, SAR, and Archaeplastida.

Trypanosoma rangeli is a species of hemoflagellate excavate parasites of the genus Trypanosoma. Although infecting a variety of mammalian species in a wide geographical area in Central and South America, this parasite is considered non-pathogenic to these hosts. T. rangeli is transmitted by bite of infected triatomine bugs of the Reduviidae family, commonly known as barbeiro, winchuka(vinchuca), chinche, pito ou chupão.

<i>Leishmania mexicana</i> Species of parasitic protist

Leishmania mexicana is a species of obligate intracellular parasites of the protozoan genus Leishmania. In Mexico and Central America, this parasite is the primary cause of cutaneous leishmaniasis.

<i>Strigomonas culicis</i> Species of parasitic flagellate protist in the Kinetoplastea class

Strigomonas culicis is a protist and member of flagellated trypanosomatids. It is an obligate parasite in the gastrointestinal tract of mosquito, and is in turn a host to symbiotic bacteria. It maintains strict mutualistic relationship with the bacteria as a sort of cell organelle (endosymbiont) so that it cannot lead an independent life without the bacteria. Along with Angomonas deanei, S. culicis is researched as model organism for the evolution of symbiotic relationsships with intracellular bacteria.

Novymonas esmeraldas is a protist and member of flagellated trypanosomatids. It is an obligate parasite in the gastrointestinal tract of a bug, and is in turn a host to symbiotic bacteria. It maintains strict mutualistic relationship with the bacteria as a sort of cell organelle (endosymbiont) so that it cannot lead an independent life without the bacteria. Its discovery in 2016 suggests that it is a good model in the evolution of prokaryotes into eukaryotes by symbiogenesis. The endosymbiotic bacterium was identified as member of the genus Pandoraea.

References

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  13. 1 2 3 Santos, André L.S.; d'Avila-Levy, Claudia M.; Elias, Camila G.R.; Vermelho, Alane B.; Branquinha, Marta H. (2007). "Phytomonas serpens: immunological similarities with the human trypanosomatid pathogens". Microbes and Infection. 9 (8): 915–921. doi:10.1016/j.micinf.2007.03.018. ISSN   1286-4579. PMID   17556002.
  14. ↑ Brazil, Reginaldo Peçanha; Fiorini, João Evangelista; Silva, Paulo Márcio Faria e; Brazil, Reginaldo Peçanha; Fiorini, João Evangelista; Silva, Paulo Márcio Faria e (June 1990). "Phytomonas sp., a trypanosomatid parasite of tomato, isolated from salivary glands of Phthia picta (Hemiptera: Coreidae) in southeast Brazil". Memórias do Instituto Oswaldo Cruz. 85 (2): 239–240. doi: 10.1590/S0074-02761990000200016 . ISSN   0074-0276.
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