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88-472: Microsporidia are a group of spore -forming unicellular parasites . These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or protists , but are now known to be fungi , or a sister group to true fungi . These fungal microbes are obligate eukaryotic parasites that use a unique mechanism to infect host cells. They have recently been discovered in

176-438: A holoparasite such as Cuscuta derives all of its nutrients from another plant. Parasitic plants make up about one per cent of angiosperms and are in almost every biome in the world. All these plants have modified roots, haustoria , which penetrate the host plants, connecting them to the conductive system—either the xylem , the phloem , or both. This provides them with the ability to extract water and nutrients from

264-457: A megasporangium that produces megaspores or a microsporangium that produces microspores. In flowering plants, these sporangia occur within the carpel and anthers, respectively. Fungi commonly produce spores during sexual and asexual reproduction. Spores are usually haploid and grow into mature haploid individuals through mitotic division of cells ( Urediniospores and Teliospores among rusts are dikaryotic). Dikaryotic cells result from

352-547: A zygote , which develops into a new sporophyte. This cycle is known as alternation of generations . The spores of seed plants are produced internally, and the megaspores (formed within the ovules) and the microspores are involved in the formation of more complex structures that form the dispersal units, the seeds and pollen grains. The term spore derives from the ancient Greek word σπορά spora , meaning " seed , sowing", related to σπόρος sporos , "sowing", and σπείρειν speirein , "to sow". In common parlance,

440-409: A 2017 Cornell study to infect Coleoptera on a large scale. So far, about 1500 of the probably more than one million species are named. Microsporidia are restricted to animal hosts , and all major groups of animals host microsporidia. Most infect insects , but they are also responsible for common diseases of crustaceans and fish . The named species of microsporidia usually infect one host species or

528-511: A cause of gastroenteritis , is spread by the fecal–oral route from animals, or by eating insufficiently cooked poultry , or by contaminated water. Haemophilus influenzae , an agent of bacterial meningitis and respiratory tract infections such as influenza and bronchitis , is transmitted by droplet contact. Treponema pallidum , the cause of syphilis , is spread by sexual activity . Viruses are obligate intracellular parasites, characterised by extremely limited biological function, to

616-437: A center pole. This shows that four spores shared a common origin and were initially in contact with each other forming a tetrahedron. A wider aperture in the shape of a groove may be termed a colpus . The number of colpi distinguishes major groups of plants. Eudicots have tricolpate spores (i.e. spores with three colpi). Envelope-enclosed spore tetrads are taken as the earliest evidence of plant life on land, dating from

704-559: A consequence, the genomes of microsporidia are much smaller than those of other eukaryotes. Currently known microsporidial genomes are 2.5 to 11.6 Mb in size, encoding from 1,848 to 3,266 proteins which is in the same range as many bacteria . Horizontal gene transfer (HGT) seems to have occurred many times in microsporidia. For instance, the genomes of Encephalitozoon romaleae and Trachipleistophora hominis contain genes that derive from animals and bacteria, and some even from fungi. The Rad9 - Rad1 -Hus1 protein complex (also known as

792-417: A digenean, the faustulid Bacciger israelensis , itself an intestinal parasite of a marine fish, the bogue Boops boops (Teleostei, Sparidae). Microsporidia have the smallest known (nuclear) eukaryotic genomes . The parasitic lifestyle of microsporidia has led to a loss of many mitochondrial and Golgi genes, and even their ribosomal RNAs are reduced in size compared with those of most eukaryotes . As

880-627: A faster rate than their hosts. Classic examples include interactions between vertebrate hosts and tapeworms , flukes , and those between the malaria-causing Plasmodium species, and fleas . Parasites reduce host fitness by general or specialised pathology , that ranges from parasitic castration to modification of host behaviour . Parasites increase their own fitness by exploiting hosts for resources necessary for their survival, in particular by feeding on them and by using intermediate (secondary) hosts to assist in their transmission from one definitive (primary) host to another. Although parasitism

968-485: A few are used in biological control of insect pests. Parasitic castration , gigantism, or change of host sex are all potential effects of microsporidian parasitism (in insects). In the most advanced cases of parasitism the microsporidium rules the host cell completely and controls its metabolism and reproduction, forming a xenoma . Replication takes place within the host's cells, which are infected by means of unicellular spores . These vary from 1–40 μm, making them some of

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1056-419: A group of closely related taxa. Approximately 10 percent of the known species are parasites of vertebrates — several species, most of which are opportunistic, can infect humans, in whom they can cause microsporidiosis . After infection they influence their hosts in various ways and all organs and tissues are invaded, though generally by different species of specialised microsporidia. Some species are lethal, and

1144-595: A hormone or by diverting nutrients. For example, the trematode Zoogonus lasius , whose sporocysts lack mouths, castrates the intertidal marine snail Tritia obsoleta chemically, developing in its gonad and killing its reproductive cells. Directly transmitted parasites, not requiring a vector to reach their hosts, include such parasites of terrestrial vertebrates as lice and mites; marine parasites such as copepods and cyamid amphipods; monogeneans ; and many species of nematodes, fungi, protozoans, bacteria, and viruses. Whether endoparasites or ectoparasites, each has

1232-836: A parasite employs to identify and approach a potential host are known as "host cues". Such cues can include, for example, vibration, exhaled carbon dioxide , skin odours, visual and heat signatures, and moisture. Parasitic plants can use, for example, light, host physiochemistry, and volatiles to recognize potential hosts. There are six major parasitic strategies , namely parasitic castration ; directly transmitted parasitism; trophically -transmitted parasitism; vector -transmitted parasitism; parasitoidism ; and micropredation. These apply to parasites whose hosts are plants as well as animals. These strategies represent adaptive peaks ; intermediate strategies are possible, but organisms in many different groups have consistently converged on these six, which are evolutionarily stable. A perspective on

1320-618: A parasitic alien species. First used in English in 1539, the word parasite comes from the Medieval French parasite , from the Latinised form parasitus , from Ancient Greek παράσιτος (parasitos)  'one who eats at the table of another' in turn from παρά (para)  'beside, by' and σῖτος (sitos)  'wheat, food'. The related term parasitism appears in English from 1611. Parasitism

1408-432: A phenomenon termed the biotrophy-necrotrophy switch . Pathogenic fungi are well-known causative agents of diseases on animals as well as humans. Fungal infections ( mycosis ) are estimated to kill 1.6 million people each year. One example of a potent fungal animal pathogen are Microsporidia - obligate intracellular parasitic fungi that largely affect insects, but may also affect vertebrates including humans, causing

1496-533: A predator, the European sparrowhawk , giving her time to lay her eggs in the host's nest unobserved. Host species often combat parasitic egg mimicry through egg polymorphism , having two or more egg phenotypes within a single population of a species. Multiple phenotypes in host eggs decrease the probability of a parasitic species accurately "matching" their eggs to host eggs. In kleptoparasitism (from Greek κλέπτης ( kleptēs ), "thief"), parasites steal food gathered by

1584-595: A putrid odour, for dispersal of fungal spores is yet another strategy, most prominently used by the stinkhorns . In Common Smoothcap moss ( Atrichum undulatum ), the vibration of sporophyte has been shown to be an important mechanism for spore release. In the case of spore-shedding vascular plants such as ferns, wind distribution of very light spores provides great capacity for dispersal. Also, spores are less subject to animal predation than seeds because they contain almost no food reserve; however they are more subject to fungal and bacterial predation. Their chief advantage

1672-423: A simple asexual life cycle, while others have a complex life cycle involving multiple hosts and both asexual and sexual reproduction . Different types of spores may be produced at different stages, probably with different functions including autoinfection (transmission within a single host). In animals and humans, microsporidia often cause chronic, debilitating diseases rather than lethal infections. Effects on

1760-573: A single host-species. Within that species, most individuals are free or almost free of parasites, while a minority carry a large number of parasites; this is known as an aggregated distribution . Trophically -transmitted parasites are transmitted by being eaten by a host. They include trematodes (all except schistosomes ), cestodes , acanthocephalans , pentastomids , many roundworms , and many protozoa such as Toxoplasma . They have complex life cycles involving hosts of two or more species. In their juvenile stages they infect and often encyst in

1848-430: A suitable fungus soon after germinating. Parasitic fungi derive some or all of their nutritional requirements from plants, other fungi, or animals. Plant pathogenic fungi are classified into three categories depending on their mode of nutrition: biotrophs, hemibiotrophs and necrotrophs. Biotrophic fungi derive nutrients from living plant cells, and during the course of infection they colonise their plant host in such

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1936-1047: A variety of methods to infect animal hosts, including physical contact, the fecal–oral route , free-living infectious stages, and vectors, suiting their differing hosts, life cycles, and ecological contexts. Examples to illustrate some of the many possible combinations are given in the table. social behaviour (grooming) Among the many variations on parasitic strategies are hyperparasitism, social parasitism, brood parasitism, kleptoparasitism, sexual parasitism, and adelphoparasitism. Hyperparasites feed on another parasite, as exemplified by protozoa living in helminth parasites, or facultative or obligate parasitoids whose hosts are either conventional parasites or parasitoids. Levels of parasitism beyond secondary also occur, especially among facultative parasitoids. In oak gall systems, there can be up to five levels of parasitism. Hyperparasites can control their hosts' populations, and are used for this purpose in agriculture and to some extent in medicine . The controlling effects can be seen in

2024-469: A variety of routes. To give a few examples, Bacillus anthracis , the cause of anthrax , is spread by contact with infected domestic animals ; its spores , which can survive for years outside the body, can enter a host through an abrasion or may be inhaled. Borrelia , the cause of Lyme disease and relapsing fever , is transmitted by vectors, ticks of the genus Ixodes , from the diseases' reservoirs in animals such as deer . Campylobacter jejuni ,

2112-474: A way as to keep it alive for a maximally long time. One well-known example of a biotrophic pathogen is Ustilago maydis , causative agent of the corn smut disease. Necrotrophic pathogens on the other hand, kill host cells and feed saprophytically , an example being the root-colonising honey fungi in the genus Armillaria . Hemibiotrophic pathogens begin their colonising their hosts as biotrophs, and subsequently killing off host cells and feeding as necrotrophs,

2200-563: Is a kind of symbiosis , a close and persistent long-term biological interaction between a parasite and its host. Unlike saprotrophs , parasites feed on living hosts, though some parasitic fungi, for instance, may continue to feed on hosts they have killed. Unlike commensalism and mutualism , the parasitic relationship harms the host, either feeding on it or, as in the case of intestinal parasites, consuming some of its food. Because parasites interact with other species, they can readily act as vectors of pathogens, causing disease . Predation

2288-675: Is aggregated. Coinfection by multiple parasites is common. Autoinfection , where (by exception) the whole of the parasite's life cycle takes place in a single primary host, can sometimes occur in helminths such as Strongyloides stercoralis . Vector-transmitted parasites rely on a third party, an intermediate host, where the parasite does not reproduce sexually, to carry them from one definitive host to another. These parasites are microorganisms, namely protozoa , bacteria , or viruses , often intracellular pathogens (disease-causers). Their vectors are mostly hematophagic arthropods such as fleas, lice, ticks, and mosquitoes. For example,

2376-443: Is by definition not a symbiosis, as the interaction is brief, but the entomologist E. O. Wilson has characterised parasites as "predators that eat prey in units of less than one". Within that scope are many possible strategies. Taxonomists classify parasites in a variety of overlapping schemes, based on their interactions with their hosts and on their life cycles , which can be complex. An obligate parasite depends completely on

2464-645: Is found in the ant Tetramorium inquilinum , an obligate parasite which lives exclusively on the backs of other Tetramorium ants. A mechanism for the evolution of social parasitism was first proposed by Carlo Emery in 1909. Now known as " Emery's rule ", it states that social parasites tend to be closely related to their hosts, often being in the same genus. Intraspecific social parasitism occurs in parasitic nursing, where some individual young take milk from unrelated females. In wedge-capped capuchins , higher ranking females sometimes take milk from low ranking females without any reciprocation. In brood parasitism ,

2552-585: Is often unambiguous, it is part of a spectrum of interactions between species , grading via parasitoidism into predation, through evolution into mutualism , and in some fungi, shading into being saprophytic . Human knowledge of parasites such as roundworms and tapeworms dates back to ancient Egypt , Greece , and Rome . In early modern times, Antonie van Leeuwenhoek observed Giardia lamblia with his microscope in 1681, while Francesco Redi described internal and external parasites including sheep liver fluke and ticks . Modern parasitology developed in

2640-419: Is seen in some species of anglerfish , such as Ceratias holboelli , where the males are reduced to tiny sexual parasites , wholly dependent on females of their own species for survival, permanently attached below the female's body, and unable to fend for themselves. The female nourishes the male and protects him from predators, while the male gives nothing back except the sperm that the female needs to produce

2728-478: Is that spores were an adaptation of early land plant species, such as embryophytes , that allowed for plants to easily disperse while adapting to their non-aquatic environment. This is particularly supported by the observation of a thick spore wall in cryptospores . These spore walls would have protected potential offspring from novel weather elements. The second more recent hypothesis is that spores were an early predecessor of land plants and formed during errors in

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2816-451: Is that, of all forms of progeny, spores require the least energy and materials to produce. In the spikemoss Selaginella lepidophylla , dispersal is achieved in part by an unusual type of diaspore , a tumbleweed . Spores have been found in microfossils dating back to the mid-late Ordovician period. Two hypothesized initial functions of spores relate to whether they appeared before or after land plants. The heavily studied hypothesis

2904-469: Is then carried to a nest, sometimes alongside other prey if it is not large enough to support a parasitoid throughout its development. An egg is laid on top of the prey and the nest is then sealed. The parasitoid develops rapidly through its larval and pupal stages, feeding on the provisions left for it. Koinobiont parasitoids, which include flies as well as wasps, lay their eggs inside young hosts, usually larvae. These are allowed to go on growing, so

2992-417: Is useful in distinguishing between different species. Spores of most species are oval or pyriform, but rod-shaped or spherical spores are not unusual. A few genera produce spores of unique shape for the genus. The spore is protected by a wall, consisting of three layers: In most cases there are two closely associated nuclei , forming a diplokaryon , but sometimes there is only one. The anterior half of

3080-428: The host , causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria , sleeping sickness , and amoebic dysentery ; animals such as hookworms , lice , mosquitoes , and vampire bats ; fungi such as honey fungus and

3168-491: The meiosis of algae , a hypothesized early ancestor of land plants. Whether spores arose before or after land plants, their contributions to topics in fields like paleontology and plant phylogenetics have been useful. The spores found in microfossils, also known as cryptospores, are well preserved due to the fixed material they are in as well as how abundant and widespread they were during their respective time periods. These microfossils are especially helpful when studying

3256-505: The snubnosed eel is probably a facultative endoparasite (i.e., it is semiparasitic) that opportunistically burrows into and eats sick and dying fish. Plant-eating insects such as scale insects , aphids , and caterpillars closely resemble ectoparasites, attacking much larger plants; they serve as vectors of bacteria, fungi and viruses which cause plant diseases . As female scale insects cannot move, they are obligate parasites, permanently attached to their hosts. The sensory inputs that

3344-504: The 19th century. In human culture, parasitism has negative connotations. These were exploited to satirical effect in Jonathan Swift 's 1733 poem "On Poetry: A Rhapsody", comparing poets to hyperparasitical "vermin". In fiction, Bram Stoker 's 1897 Gothic horror novel Dracula and its many later adaptations featured a blood-drinking parasite. Ridley Scott 's 1979 film Alien was one of many works of science fiction to feature

3432-712: The 9-1-1 complex) in eukaryotes is recruited to sites of DNA damage where it is considered to help trigger the checkpoint-signaling cascade. Genes that code for heterotrimeric 9-1-1 are present in microsporidia. In addition to the 9-1-1 complex, other components of the DNA repair machinery are also present indicting that repair of DNA damage likely occurs in microsporidia. Phylogeny of Rozellomycota Rozellales Mitosporidiaceae Morellosporaceae Paramicrosporidiaceae Nucleophagaceae Chytridiopsida Metchnikovellida Neopereziida Ovavesiculida Amblyosporida Glugeida Nosematida The first described microsporidian genus, Nosema ,

3520-458: The agents of ringworm ; and plants such as mistletoe , dodder , and the broomrapes . There are six major parasitic strategies of exploitation of animal hosts, namely parasitic castration , directly transmitted parasitism (by contact), trophically-transmitted parasitism (by being eaten), vector-transmitted parasitism, parasitoidism , and micropredation. One major axis of classification concerns invasiveness: an endoparasite lives inside

3608-696: The air or soil given off by host shoots or roots , respectively. About 4,500 species of parasitic plant in approximately 20 families of flowering plants are known. Species within the Orobanchaceae (broomrapes) are among the most economically destructive of all plants. Species of Striga (witchweeds) are estimated to cost billions of dollars a year in crop yield loss, infesting over 50 million hectares of cultivated land within Sub-Saharan Africa alone. Striga infects both grasses and grains, including corn , rice , and sorghum , which are among

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3696-442: The amoebula. In plants, spores are usually haploid and unicellular and are produced by meiosis in the sporangium of a diploid sporophyte . In some rare cases, diploid spore is also produced in some algae, or fungi. Under favourable conditions, the spore can develop into a new organism using mitotic division, producing a multicellular gametophyte , which eventually goes on to produce gametes. Two gametes fuse to form

3784-475: The apex. The posterior vacuole swells, forcing the polar filament to rapidly eject the infectious content into the cytoplasm of the potential host. Simultaneously the material of the filament is rearranged to form a tube which functions as a hypodermic needle and penetrates the gut epithelium. Once inside the host cell, a sporoplasm grows, dividing or forming a multinucleate plasmodium , before producing new spores. The life cycle varies considerably. Some have

3872-864: The deer tick Ixodes scapularis acts as a vector for diseases including Lyme disease , babesiosis , and anaplasmosis . Protozoan endoparasites, such as the malarial parasites in the genus Plasmodium and sleeping-sickness parasites in the genus Trypanosoma , have infective stages in the host's blood which are transported to new hosts by biting insects. Parasitoids are insects which sooner or later kill their hosts, placing their relationship close to predation. Most parasitoids are parasitoid wasps or other hymenopterans ; others include dipterans such as phorid flies . They can be divided into two groups, idiobionts and koinobionts, differing in their treatment of their hosts. Idiobiont parasitoids sting their often-large prey on capture, either killing them outright or paralysing them immediately. The immobilised prey

3960-426: The difference between a "spore" and a " gamete " is that a spore will germinate and develop into a sporeling , while a gamete needs to combine with another gamete to form a zygote before developing further. The main difference between spores and seeds as dispersal units is that spores are unicellular, the first cell of a gametophyte, while seeds contain within them a developing embryo (the multicellular sporophyte of

4048-438: The early periods of earth as macrofossils such as plants are not common nor well preserved. Both cryptospores and modern spores have diverse morphology that indicate possible environmental conditions of earlier periods of Earth and evolutionary relationships of plant species. Autoinfection Parasitism is a close relationship between species , where one organism, the parasite, lives on or inside another organism,

4136-409: The energy that would have gone into reproduction into host and parasite growth, sometimes causing gigantism in the host. The host's other systems remain intact, allowing it to survive and to sustain the parasite. Parasitic crustaceans such as those in the specialised barnacle genus Sacculina specifically cause damage to the gonads of their many species of host crabs . In the case of Sacculina ,

4224-491: The evolutionary options can be gained by considering four key questions: the effect on the fitness of a parasite's hosts; the number of hosts they have per life stage; whether the host is prevented from reproducing; and whether the effect depends on intensity (number of parasites per host). From this analysis, the major evolutionary strategies of parasitism emerge, alongside predation. Parasitic castrators partly or completely destroy their host's ability to reproduce, diverting

4312-444: The family Cuculidae , over 40% of cuckoo species are obligate brood parasites, while others are either facultative brood parasites or provide parental care. The eggs of some brood parasites mimic those of their hosts, while some cowbird eggs have tough shells, making them hard for the hosts to kill by piercing, both mechanisms implying selection by the hosts against parasitic eggs. The adult female European cuckoo further mimics

4400-438: The fitness of host mosquitoes, and was transmitted vertically (through inheritance). Microsporidian infections of humans sometimes cause a disease called microsporidiosis . At least 14 microsporidian species, spread across eight genera, have been recognized as human pathogens . These include Trachipleistophora hominis . Microsporidia can infect a variety of hosts, including hosts which are themselves parasites. In that case,

4488-569: The fluids of the ascus that lead to explosive discharge of the ascospores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water ( Buller's drop ), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g . Other fungi rely on alternative mechanisms for spore release, such as external mechanical forces, exemplified by puffballs . Attracting insects, such as flies, to fruiting structures, by virtue of their having lively colours and

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4576-735: The fusion of two haploid gamete cells. Among sporogenic dikaryotic cells, karyogamy (the fusion of the two haploid nuclei) occurs to produce a diploid cell. Diploid cells undergo meiosis to produce haploid spores. Spores can be classified in several ways such as by their spore producing structure, function, origin during life cycle, and mobility. Below is a table listing the mode of classification, name, identifying characteristic, examples, and images of different spore species. Under high magnification , spores often have complex patterns or ornamentation on their exterior surfaces. A specialized terminology has been developed to describe features of such patterns. Some markings represent apertures, places where

4664-467: The habitat: Aquasporidia, Marinosporidia and Terresporidia. A second classification by Cavalier-Smith 1993: Spore In biology , a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants , algae , fungi and protozoa . They were thought to have appeared as early as

4752-424: The host and parasitoid develop together for an extended period, ending when the parasitoids emerge as adults, leaving the prey dead, eaten from inside. Some koinobionts regulate their host's development, for example preventing it from pupating or making it moult whenever the parasitoid is ready to moult. They may do this by producing hormones that mimic the host's moulting hormones ( ecdysteroids ), or by regulating

4840-418: The host include reduced longevity, fertility, weight, and general vigor. Vertical transmission of microsporidia is frequently reported. In the case of insect hosts, vertical transmission often occurs as transovarial transmission, where the microsporidian parasites pass from the ovaries of the female host into eggs and eventually multiply in the infected larvae. Amblyospora salinaria n. sp. which infects

4928-442: The host to complete its life cycle, while a facultative parasite does not. Parasite life cycles involving only one host are called "direct"; those with a definitive host (where the parasite reproduces sexually) and at least one intermediate host are called "indirect". An endoparasite lives inside the host's body; an ectoparasite lives outside, on the host's surface. Mesoparasites—like some copepods , for example—enter an opening in

5016-422: The host's body and remain partly embedded there. Some parasites can be generalists, feeding on a wide range of hosts, but many parasites, and the majority of protozoans and helminths that parasitise animals, are specialists and extremely host-specific. An early basic, functional division of parasites distinguished microparasites and macroparasites. These each had a mathematical model assigned in order to analyse

5104-453: The host's body; an ectoparasite lives outside, on the host's surface. Like predation, parasitism is a type of consumer–resource interaction , but unlike predators , parasites, with the exception of parasitoids, are much smaller than their hosts, do not kill them, and often live in or on their hosts for an extended period. Parasites of animals are highly specialised , each parasite species living on one given animal species, and reproduce at

5192-656: The host's endocrine system. A micropredator attacks more than one host, reducing each host's fitness by at least a small amount, and is only in contact with any one host intermittently. This behavior makes micropredators suitable as vectors, as they can pass smaller parasites from one host to another. Most micropredators are hematophagic , feeding on blood. They include annelids such as leeches , crustaceans such as branchiurans and gnathiid isopods, various dipterans such as mosquitoes and tsetse flies , other arthropods such as fleas and ticks, vertebrates such as lampreys , and mammals such as vampire bats . Parasites use

5280-425: The host. A parasitic plant is classified depending on where it latches onto the host, either the stem or the root, and the amount of nutrients it requires. Since holoparasites have no chlorophyll and therefore cannot make food for themselves by photosynthesis , they are always obligate parasites, deriving all their food from their hosts. Some parasitic plants can locate their host plants by detecting chemicals in

5368-482: The host. Furthermore, spore-forming organisms in general do have a complex system of reproduction, both sexual and asexual, which look far from primitive. Since the mid-2000s microsporidia are placed within the Fungi or as a sister-group of the Fungi with a common ancestor. Work to identify clades is largely based on habitat and host. Three classes of Microsporidia are proposed by Vossbrinck and Debrunner-Vossbrinck, based on

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5456-477: The host. The parasitism is often on close relatives, whether within the same species or between species in the same genus or family. For instance, the many lineages of cuckoo bees lay their eggs in the nest cells of other bees in the same family. Kleptoparasitism is uncommon generally but conspicuous in birds; some such as skuas are specialised in pirating food from other seabirds, relentlessly chasing them down until they disgorge their catch. A unique approach

5544-423: The hosts suffer increased parental investment and energy expenditure to feed parasitic young, which are commonly larger than host young. The growth rate of host nestlings is slowed, reducing the host's fitness. Brood parasites include birds in different families such as cowbirds , whydahs , cuckoos , and black-headed ducks . These do not build nests of their own, but leave their eggs in nests of other species . In

5632-472: The intermediate host. When the intermediate-host animal is eaten by a predator, the definitive host, the parasite survives the digestion process and matures into an adult; some live as intestinal parasites . Many trophically transmitted parasites modify the behaviour of their intermediate hosts, increasing their chances of being eaten by a predator. As with directly transmitted parasites, the distribution of trophically transmitted parasites among host individuals

5720-471: The intestinal infection microsporidiosis . Protozoa such as Plasmodium , Trypanosoma , and Entamoeba are endoparasitic. They cause serious diseases in vertebrates including humans—in these examples, malaria, sleeping sickness, and amoebic dysentery —and have complex life cycles. Many bacteria are parasitic, though they are more generally thought of as pathogens causing disease. Parasitic bacteria are extremely diverse, and infect their hosts by

5808-441: The large blue butterfly, Phengaris arion , its larvae employing ant mimicry to parasitise certain ants, Bombus bohemicus , a bumblebee which invades the hives of other bees and takes over reproduction while their young are raised by host workers, and Melipona scutellaris , a eusocial bee whose virgin queens escape killer workers and invade another colony without a queen. An extreme example of interspecific social parasitism

5896-567: The largest group is the parasitoid wasps in the Hymenoptera. The phyla and classes with the largest numbers of parasitic species are listed in the table. Numbers are conservative minimum estimates. The columns for Endo- and Ecto-parasitism refer to the definitive host, as documented in the Vertebrate and Invertebrate columns. A hemiparasite or partial parasite such as mistletoe derives some of its nutrients from another living plant, whereas

5984-537: The marine worm Bonellia viridis has a similar reproductive strategy, although the larvae are planktonic. Examples of the major variant strategies are illustrated. Parasitism has an extremely wide taxonomic range, including animals, plants, fungi, protozoans, bacteria, and viruses. Parasitism is widespread in the animal kingdom, and has evolved independently from free-living forms hundreds of times. Many types of helminth including flukes and cestodes have complete life cycles involving two or more hosts. By far

6072-400: The microsporidian species is a hyperparasite , i.e. a parasite of a parasite. As an example, more than eighteen species are known which parasitize digeneans (parasitic flatworms ). These digeneans are themselves parasites in various vertebrates and molluscs . Eight of these species belong to the genus Nosema . Similarly, the microsporidian species Toguebayea baccigeri is a parasite of

6160-527: The mid-Ordovician (early Llanvirn, ~ 470  million years ago ), a period from which no macrofossils have yet been recovered. Individual trilete spores resembling those of modern cryptogamic plants first appeared in the fossil record at the end of the Ordovician period. In fungi, both asexual and sexual spores or sporangiospores of many fungal species are actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of

6248-425: The mid-late Ordovician period as an adaptation of early land plants. Bacterial spores are not part of a sexual cycle, but are resistant structures used for survival under unfavourable conditions. Myxozoan spores release amoeboid infectious germs ("amoebulae") into their hosts for parasitic infection, but also reproduce within the hosts through the pairing of two nuclei within the plasmodium, which develops from

6336-498: The mosquito Culex salinarius Coquillett, and Amblyospora californica which infects the mosquito Culex tarsalis Coquillett, provide typical examples of transovarial transmission of microsporidia. Microsporidia, specifically the mosquito-infecting Vavraia culicis , are being explored as a possible 'evolution-proof' malaria-control method. Microsporidian infection of Anopheles gambiae (the principal vector of Plasmodium falciparum malaria) reduces malarial infection within

6424-479: The mosquito, and shortens the mosquito lifespan. As the majority of malaria-infected mosquitoes naturally die before the malaria parasite is mature enough to transmit, any increase in mosquito mortality through microsporidian-infection may reduce malaria transmission to humans. In May 2020, researchers reported that Microsporidia MB , a symbiont in the midgut and ovaries of An. arabiensis , significantly impaired transmission of P. falciparum , had "no overt effect" on

6512-443: The next generation), produced by the fusion of the male gamete of the pollen tube with the female gamete formed by the megagametophyte within the ovule. Spores germinate to give rise to haploid gametophytes, while seeds germinate to give rise to diploid sporophytes. Vascular plant spores are always haploid . Vascular plants are either homosporous (or isosporous) or heterosporous . Plants that are homosporous produce spores of

6600-407: The next generation. Adelphoparasitism, (from Greek ἀδελφός ( adelphós ), brother ), also known as sibling-parasitism, occurs where the host species is closely related to the parasite, often in the same family or genus. In the citrus blackfly parasitoid, Encarsia perplexa , unmated females may lay haploid eggs in the fully developed larvae of their own species, producing male offspring, while

6688-417: The point where, while they are evidently able to infect all other organisms from bacteria and archaea to animals, plants and fungi, it is unclear whether they can themselves be described as living. They can be either RNA or DNA viruses consisting of a single or double strand of genetic material ( RNA or DNA , respectively), covered in a protein coat and sometimes a lipid envelope. They thus lack all

6776-535: The population movements of the host–parasite groupings. The microorganisms and viruses that can reproduce and complete their life cycle within the host are known as microparasites. Macroparasites are the multicellular organisms that reproduce and complete their life cycle outside of the host or on the host's body. Much of the thinking on types of parasitism has focused on terrestrial animal parasites of animals, such as helminths. Those in other environments and with other hosts often have analogous strategies. For example,

6864-399: The same size and type. Heterosporous plants, such as seed plants , spikemosses , quillworts , and ferns of the order Salviniales produce spores of two different sizes: the larger spore (megaspore) in effect functioning as a "female" spore and the smaller (microspore) functioning as a "male". Such plants typically give rise to the two kind of spores from within separate sporangia, either

6952-617: The smallest eukaryotes . Microsporidia that infect mammals are 1.0–4.0 μm. They also have the smallest eukaryotic genomes . The terms "microsporidium" (pl. "microsporidia") and "microsporidian" are used as vernacular names for members of the group. The name Microsporidium Balbiani, 1884 is also used as a catchall genus for incertae sedis members. Microsporidia lack mitochondria , instead possessing mitosomes . They also lack motile structures, such as flagella . Microsporidia produce highly resistant spores, capable of surviving outside their host for up to several years. Spore morphology

7040-430: The spore contains a harpoon-like apparatus with a long, thread-like polar filament , which is coiled up in the posterior half of the spore. The anterior part of the polar filament is surrounded by a polaroplast , a lamella of membranes. Behind the polar filament, there is a posterior vacuole . In the gut of the host the spore germinates; it builds up osmotic pressure until its rigid wall ruptures at its thinnest point at

7128-407: The spores from the reproductive structures as well as travelling through the air over long distances. Many fungi thereby possess specialized mechanical and physiological mechanisms as well as spore-surface structures, such as hydrophobins , for spore ejection. These mechanisms include, for example, forcible discharge of ascospores enabled by the structure of the ascus and accumulation of osmolytes in

7216-466: The testes of over two-thirds of their crab hosts degenerate sufficiently for these male crabs to develop female secondary sex characteristics such as broader abdomens, smaller claws and egg-grasping appendages. Various species of helminth castrate their hosts (such as insects and snails). This may happen directly, whether mechanically by feeding on their gonads, or by secreting a chemical that destroys reproductive cells; or indirectly, whether by secreting

7304-418: The tough outer coat of the spore can be penetrated when germination occurs. Spores can be categorized based on the position and number of these markings and apertures. Alete spores show no lines. In monolete spores , there is a single narrow line (laesura) on the spore. Indicating the prior contact of two spores that eventually separated. In trilete spores , each spore shows three narrow lines radiating from

7392-403: The tropics, however effectively cheat by taking carbon from a fungus rather than exchanging it for minerals. They have much reduced roots, as they do not need to absorb water from the soil; their stems are slender with few vascular bundles , and their leaves are reduced to small scales, as they do not photosynthesize. Their seeds are small and numerous, so they appear to rely on being infected by

7480-506: The usual machinery of the cell such as enzymes , relying entirely on the host cell's ability to replicate DNA and synthesise proteins. Most viruses are bacteriophages , infecting bacteria. Parasitism is a major aspect of evolutionary ecology; for example, almost all free-living animals are host to at least one species of parasite. Vertebrates, the best-studied group, are hosts to between 75,000 and 300,000 species of helminths and an uncounted number of parasitic microorganisms. On average,

7568-585: The way that the CHV1 virus helps to control the damage that chestnut blight , Cryphonectria parasitica , does to American chestnut trees, and in the way that bacteriophages can limit bacterial infections. It is likely, though little researched, that most pathogenic microparasites have hyperparasites which may prove widely useful in both agriculture and medicine. Social parasites take advantage of interspecific interactions between members of eusocial animals such as ants , termites , and bumblebees . Examples include

7656-471: The world's most important food crops. Orobanche also threatens a wide range of other important crops, including peas , chickpeas , tomatoes , carrots , and varieties of cabbage . Yield loss from Orobanche can be total; despite extensive research, no method of control has been entirely successful. Many plants and fungi exchange carbon and nutrients in mutualistic mycorrhizal relationships. Some 400 species of myco-heterotrophic plants, mostly in

7744-680: Was initially put by Nägeli in the fungal group Schizomycetes together with some bacteria and yeasts . For some time microsporidia were considered as very primitive eukaryotes, placed in the protozoan group Cnidospora . Later, especially because of the lack of mitochondria, they were placed along with the other Protozoa such as diplomonads , parabasalids and archamoebae in the protozoan -group Archezoa . More recent research has falsified this theory of early origin (for all of these). Instead, microsporidia are proposed to be highly developed and specialized organisms, which just dispensed functions that are needed no longer, because they are supplied by

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