Host (biology)

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The black rat is a reservoir host for bubonic plague: the oriental rat fleas that infest these rats are vectors for the disease. Roof rat-(rattus rattus).jpg
The black rat is a reservoir host for bubonic plague: the oriental rat fleas that infest these rats are vectors for the disease.

In biology and medicine, a host is a larger organism that harbours a smaller organism; [1] whether a parasitic, a mutualistic, or a commensalist guest (symbiont). The guest is typically provided with nourishment and shelter. Examples include animals playing host to parasitic worms (e.g. nematodes), cells harbouring pathogenic (disease-causing) viruses, a bean plant hosting mutualistic (helpful) nitrogen-fixing bacteria. More specifically in botany, a host plant supplies food resources to micropredators, which have an evolutionarily stable relationship with their hosts similar to ectoparasitism. The host range is the collection of hosts that an organism can use as a partner.

Contents

Symbiosis

Symbiosis spans a wide variety of possible relationships between organisms, differing in their permanence and their effects on the two parties. If one of the partners in an association is much larger than the other, it is generally known as the host. [1] In parasitism, the parasite benefits at the host's expense. [2] In commensalism, the two live together without harming each other, [3] while in mutualism, both parties benefit. [4]

Most parasites are only parasitic for part of their life cycle. By comparing parasites with their closest free-living relatives, parasitism has been shown to have evolved on at least 233 separate occasions. Some organisms live in close association with a host and only become parasitic when environmental conditions deteriorate. [5]

A parasite may have a long-term relationship with its host, as is the case with all endoparasites. The guest seeks out the host and obtains food or another service from it, but does not usually kill it. [6] In contrast, a parasitoid spends a large part of its life within or on a single host, ultimately causing the host's death, with some of the strategies involved verging on predation. Generally, the host is kept alive until the parasitoid is fully grown and ready to pass on to its next life stage. [7] A guest's relationship with its host may be intermittent or temporary, perhaps associated with multiple hosts, making the relationship equivalent to the herbivory of a wild-living animal. Another possibility is that the host–guest relationship may have no permanent physical contact, as in the brood parasitism of the cuckoo. [6]

Hosts to parasites

Micropredator, parasite, parasitoid, and predator strategies compared. Their interactions with their hosts form a continuum. Micropredation and parasitoidism are now considered to be evolutionary strategies within parasitism. Micropredator Parasite Parasitoid Predator strategies compared.svg
Micropredator, parasite, parasitoid, and predator strategies compared. Their interactions with their hosts form a continuum. Micropredation and parasitoidism are now considered to be evolutionary strategies within parasitism.

Parasites follow a wide variety of evolutionary strategies, placing their hosts in an equally wide range of relationships. [2] Parasitism implies host–parasite coevolution, including the maintenance of gene polymorphisms in the host, where there is a trade-off between the advantage of resistance to a parasite and a cost such as disease caused by the gene. [8]

Types of hosts

It is not always easy or even possible to identify which host is definitive and which secondary. As the life cycles of many parasites are not well understood, sometimes the subjectively more important organism is arbitrarily labelled as definitive, and this designation may continue even after it is found to be incorrect. For example, sludge worms are sometimes considered "intermediate hosts" for salmonid whirling disease, even though the myxosporean parasite reproduces sexually inside them. [9] In trichinosis, a disease caused by roundworms, the host has reproductive adults in its digestive tract and immature juveniles in its muscles, and is therefore both an intermediate and a definitive host. [10]

Plant hosts of micropredators

Buff ermine moth caterpillar, a polyphagous micropredator Spilarctia luteum larva.JPG
Buff ermine moth caterpillar, a polyphagous micropredator

Micropredation is an evolutionarily stable strategy within parasitism, in which a small predator lives parasitically on a much larger host plant, eating parts of it. [2]

The range of plants on which a herbivorous insect feeds is known as its host range. This can be wide or narrow, but it never includes all plants. A small number of insects are monophagous, feeding on a single plant. The silkworm larva is one of these, with mulberry leaves being the only food consumed. More often, an insect with a limited host range is oligophagous, being restricted to a few closely related species, usually in the same plant family. [15] The diamondback moth is an example of this, feeding exclusively on brassicas, [16] and the larva of the potato tuber moth feeds on potatoes, tomatoes and tobacco, all members of the same plant family, Solanaceae. [17] Herbivorous insects with a wide range of hosts in various different plant families are known as polyphagous. One example is the buff ermine moth whose larvae feed on alder, mint, plantain, oak, rhubarb, currant, blackberry, dock, ragwort, nettle and honeysuckle. [18]

Influenza virus can change by genetic reassortment as it travels between different hosts in its range. AntigenicShift HiRes.svg
Influenza virus can change by genetic reassortment as it travels between different hosts in its range.

Plants often produce toxic or unpalatable secondary metabolites to deter herbivores from feeding on them. Monophagous insects have developed specific adaptations to overcome those in their specialist hosts, giving them an advantage over polyphagous species. However, this puts them at greater risk of extinction if their chosen hosts suffer setbacks. Monophagous species are able to feed on the tender young foliage with high concentrations of damaging chemicals on which polyphagous species cannot feed, having to make do with older leaves. There is a trade off between offspring quality and quantity; the specialist maximises the chances of its young thriving by paying great attention to the choice of host, while the generalist produces larger numbers of eggs in sub-optimal conditions. [19]

Some insect micropredators migrate regularly from one host to another. The hawthorn-carrot aphid overwinters on its primary host, a hawthorn tree, and migrates during the summer to its secondary host, a plant in the carrot family. [20]

Host range

The host range is the set of hosts that a parasite can use as a partner. In the case of human parasites, the host range influences the epidemiology of the parasitism or disease. For instance, the production of antigenic shifts in Influenza A virus can result from pigs being infected with the virus from several different hosts (such as human and bird). This co-infection provides an opportunity for mixing of the viral genes between existing strains, thereby producing a new viral strain. An influenza vaccine produced against an existing viral strain might not be effective against this new strain, which then requires a new influenza vaccine to be prepared for the protection of the human population. [21]

Non-parasitic associations

Mutualistic hosts

Mycorrhiza, a mutualistic interaction between a plant's roots and a fungus Mycorrhizal root tips (amanita).jpg
Mycorrhiza, a mutualistic interaction between a plant's roots and a fungus

Some hosts participate in fully mutualistic interactions with both organisms being completely dependent on the other. For example, termites are hosts to the protozoa that live in their gut and which digest cellulose, [22] and the human gut flora is essential for efficient digestion. [23] Many corals and other marine invertebrates house zooxanthellae, single-celled algae, in their tissues. The host provides a protected environment in a well-lit position for the algae, while benefiting itself from the nutrients produced by photosynthesis which supplement its diet. [24] Lamellibrachia luymesi , a deep sea giant tubeworm, has an obligate mutualistic association with internal, sulfide-oxidizing, bacterial symbionts. The tubeworm extracts the chemicals that the bacteria need from the sediment, and the bacteria supply the tubeworm, which has no mouth, with nutrients. [25] Some hermit crabs place pieces of sponge on the shell in which they are living. These grow over and eventually dissolve away the mollusc shell; the crab may not ever need to replace its abode again and is well-camouflaged by the overgrowth of sponge. [26]

An important hosting relationship is mycorrhiza, a symbiotic association between a fungus and the roots of a vascular host plant. The fungus receives carbohydrates, the products of photosynthesis, while the plant receives phosphates and nitrogenous compounds acquired by the fungus from the soil. Over 95% of plant families have been shown to have mycorrhizal associations. [27] Another such relationship is between leguminous plants and certain nitrogen-fixing bacteria called rhizobia that form nodules on the roots of the plant. The host supplies the bacteria with the energy needed for nitrogen fixation and the bacteria provide much of the nitrogen needed by the host. Such crops as beans, peas, chickpeas and alfalfa are able to fix nitrogen in this way, [28] and mixing clover with grasses increases the yield of pastures. [29]

Neurotransmitter tyramine produced by commensal Providencia bacteria, which colonize the gut of the nematode Caenorhabditis elegans , bypasses the requirement for its host to biosynthesise tyramine. This product is then probably converted to octopamine by the host enzyme tyramine β-hydroxylase and manipulates a host sensory decision. [30]

Cleaning symbiosis: a Hawaiian cleaner wrasse with its client, a yellowtail wrasse Coris gaimard and Labroides phthirophagus.JPG
Cleaning symbiosis: a Hawaiian cleaner wrasse with its client, a yellowtail wrasse

Hosts in cleaning symbiosis

Hosts of many species are involved in cleaning symbiosis, both in the sea and on land, making use of smaller animals to clean them of parasites. Cleaners include fish, shrimps and birds; hosts or clients include a much wider range of fish, marine reptiles including turtles and iguanas, octopus, whales, and terrestrial mammals. [4] The host appears to benefit from the interaction, but biologists have disputed whether this is a truly mutualistic relationship or something closer to parasitism by the cleaner. [31] [32]

Nurse shark playing host to commensal remoras, which gain a free ride and which may serve as cleaners Nurse shark with remoras (cropped).jpg
Nurse shark playing host to commensal remoras, which gain a free ride and which may serve as cleaners

Commensal hosts

Remoras (also called suckerfish) can swim freely but have evolved suckers that enable them to adhere to smooth surfaces, gaining a free ride (phoresis), and they spend most of their lives clinging to a host animal such as a whale, turtle or shark. [3] However, the relationship may be mutualistic, as remoras, though not generally considered to be cleaner fish, often consume parasitic copepods: for example, these are found in the stomach contents of 70% of the common remora. [33] Many molluscs, barnacles and polychaete worms attach themselves to the carapace of the Atlantic horseshoe crab; for some this is a convenient arrangement, but for others it is an obligate form of commensalism and they live nowhere else. [22]

History

The first host to be noticed in ancient times was human: human parasites such as hookworm are recorded from ancient Egypt from 3000 BC onwards, while in ancient Greece, the Hippocratic Corpus describes human bladder worm. [34] The medieval Persian physician Avicenna recorded human and animal parasites including roundworms, threadworms, the Guinea worm and tapeworms. [34] In Early Modern times, Francesco Redi recorded animal parasites, while the microscopist Antonie van Leeuwenhoek observed and illustrated the protozoan Giardia lamblia from "his own loose stools". [34]

Hosts to mutualistic symbionts were recognised more recently, when in 1877 Albert Bernhard Frank described the mutualistic relationship between a fungus and an alga in lichens. [35]

See also

Related Research Articles

Endosymbiont Organism that lives within the body or cells of another organism

An endosymbiont or endobiont is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living".) Examples are nitrogen-fixing bacteria, which live in the root nodules of legumes; single-cell algae inside reef-building corals, and bacterial endosymbionts that provide essential nutrients to about 10–15% of insects.

A human pathogen is a pathogen that causes disease in humans.

Symbiosis Type of a close and long-term biological interaction between two different biological organisms

Symbiosis is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasitic. The organisms, each termed a symbiont, must be of different species. In 1879, Heinrich Anton de Bary defined it as "the living together of unlike organisms". The term was subject to a century-long debate about whether it should specifically denote mutualism, as in lichens. Biologists have now abandoned that restriction.

Parasitism relationship between species where one organism lives on or in another organism, causing it harm

Parasitism is a symbiotic relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has 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 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, trophically transmitted parasitism, vector-transmitted parasitism, parasitoidism, and micropredation.

Mutualism (biology) A relationship between organisms of different species in which each individual benefits from the activity of the other

Mutualism describes the ecological interaction between two or more species where each species has a net benefit. Mutualism is a common type of ecological interaction. Prominent examples include most vascular plants engaged in mutualistic interactions with mycorrhizae, flowering plants being pollinated by animals, vascular plants being dispersed by animals, and corals with zooxanthellae, among many others. Mutualism can be contrasted with interspecific competition, in which each species experiences reduced fitness, and exploitation, or parasitism, in which one species benefits at the "expense" of the other.

Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit from each other; amensalism, where one is harmed while the other is unaffected; parasitism, where one is harmed and the other benefits, and parasitoidism, which is similar to parasitism but the parasitoid has a free-living state and instead of just harming its host it eventually ends up killing it. The commensal may obtain nutrients, shelter, support, or locomotion from the host species, which is substantially unaffected. The commensal relation is often between a larger host and a smaller commensal; the host organism is unmodified, whereas the commensal species may show great structural adaptation consistent with its habits, as in the remoras that ride attached to sharks and other fishes. Remoras feed on their hosts' fecal matter, while pilot fish feed on the leftovers of their hosts' meals. Numerous birds perch on bodies of large mammal herbivores or feed on the insects turned up by grazing mammals.

Ectosymbiosis Symbiosis in which the symbiont lives on the body surface of the host

Ectosymbiosis is a form of symbiotic behavior in which a parasite lives on the body surface of the host, including internal surfaces such as the lining of the digestive tube and the ducts of glands. The parasitic species is generally an immobile, or sessile, organism existing off of biotic substrate through mutualism, commensalism, or parasitism. Ectosymbiosis is found throughout a diverse array of environments and in many different species.

Parasitoid Organism that lives with host and kills it

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

Biological interaction Any process in which an organism has an effect on another organism

In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species, or of different species. These effects may be short-term, like pollination and predation, or long-term; both often strongly influence the evolution of the species involved. A long-term interaction is called a symbiosis. Symbioses range from mutualism, beneficial to both partners, to competition, harmful to both partners. Interactions can be indirect, through intermediaries such as shared resources or common enemies. This type of relationship can be shown by net effect based on individual effects on both organisms arising out of relationship.

In medicine, public health, and biology, transmission is the passing of a pathogen causing communicable disease from an infected host individual or group to a particular individual or group, regardless of whether the other individual was previously infected.

An obligate parasite or holoparasite is a parasitic organism that cannot complete its life-cycle without exploiting a suitable host. If an obligate parasite cannot obtain a host it will fail to reproduce. This is opposed to a facultative parasite, which can act as a parasite but does not rely on its host to continue its life-cycle. Obligate parasites have evolved a variety of parasitic strategies to exploit their hosts. Holoparasites and some hemiparasites are obligate.

Symbiotic bacteria are bacteria living in symbiosis with another organism or each other. For example, Zoamastogopera, found in the stomach of termites, enable them to digest cellulose.

Vector (epidemiology) Agent that carries and transmits an infectious pathogen into another living organism

In epidemiology, a disease vector is any agent which carries and transmits an infectious pathogen into another living organism; agents regarded as vectors are organisms, such as intermediate 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.

Introduction to viruses Non-technical introduction to viruses

A virus is a tiny infectious agent that reproduces inside the cells of living hosts. When infected, the host cell is forced to rapidly produce thousands of identical copies of the original virus. Unlike most living things, viruses do not have cells that divide; new viruses assemble in the infected host cell. But unlike simpler infectious agents like prions, they contain genes, which allow them to mutate and evolve. Over 4,800 species of viruses have been described in detail out of the millions in the environment. Their origin is unclear: some may have evolved from plasmids—pieces of DNA that can move between cells—while others may have evolved from bacteria.

Insect ecology The study of how insects interact with the surrounding environment

Insect ecology is the scientific study of how insects, individually or as a community, interact with the surrounding environment or ecosystem.

Paleoparasitology is the study of parasites from the past, and their interactions with hosts and vectors; it is a subfield of Paleontology, the study of living organisms from the past. Some authors define this term more narrowly, as "Paleoparasitology is the study of parasites in archaeological material." (p. 103) K.J. Reinhard suggests that the term "archaeoparasitology" be applied to "... all parasitological remains excavated from archaeological contexts ... derived from human activity" and that "the term 'paleoparasitology' be applied to studies of nonhuman, paleontological material." (p. 233) This article follows Reinhard's suggestion and discusses the protozoan and animal parasites of non-human animals and plants from the past, while those from humans and our hominid ancestors are covered in archaeoparasitology.

Phoresis

Phoresis or phoresy is a non-permanent, commensalistic interaction in which one organism attaches itself to another solely for the purpose of travel. Phoresis has been observed directly in ticks and mites since the 1700s and indirectly in fossils 320 million years old, but is not restricted to arthropods or animals. Plants with seeds that disperse by attaching themselves to animals are also considered to be phoretic.

In biology, a pathogen in the oldest and broadest sense, is any organism that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ.

Parasites in fiction

Parasites appear frequently in biology-inspired fiction from ancient times onwards, with a flowering in the nineteenth century. These include intentionally disgusting alien monsters in science fiction films, often with analogues in nature. Authors and scriptwriters have to some extent exploited parasite biology: lifestyles including parasitoid, behaviour-altering parasite, brood parasite, parasitic castrator, and many forms of vampire are found in books and films. Some fictional parasites, like Count Dracula and Alien's Xenomorphs, have become well known in their own right.

Symbiosis (mutualism) appears in fiction, especially science fiction, as a plot device. It is distinguished from parasitism in fiction, a similar theme, by the mutual benefit to the organisms involved, whereas the parasite inflicts harm on its host.

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