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.
It is advantageous for the parasite to preserve the health of its host when this is compatible with its nutritional and reproductive requirements, except when the death of the host is necessary for transmission. [1]
Obligate parasitism is exhibited in a range of organisms, with examples in viruses, bacteria, fungi, plants, and animals. [2] They are unable to complete their development without passing through at least one parasitic stage which is necessary to their life-cycle.
Whether one regards viruses as living organisms or not, they cannot reproduce except by means of resources within living cells. Accordingly, it is convenient and customary to regard them as obligate intracellular parasites.
Among the Vespidae family, Vespula austriaca is an example of an obligate reproductive parasite; its common host is Vespula acadica . [3] In the genus Bombus , B. bohemicus is an obligate parasite of B. locurum, B. cryptarum, and B. terrestris. [4]
Parasitic life cycles involve the exploitation of at least one host. Parasites that infect a single species are said to have direct life-cycles. [5] For example, the hookworm species Necator americanus . Parasites that infect more than one host are said to have a complex or indirect life-cycle. [5] For example, the malaria plasmodium.
An intermediate or secondary host is exploited by the parasite only for a short transition period. A final or primary host is exploited by the parasite and is the only location in which the parasite is able to reach maturity and if possible, reproduce sexually. For example, Ribeiroia ondatrae uses ramshorn snails as its first intermediate host, amphibians and fish as second intermediate hosts and birds as definitive hosts. [6]
Obligate parasites may not necessarily spend all of their time behaving as parasites. When a parasite is permanent, a number of generations occur in or on the host of an infested individual. Head lice are an example of this. Temporary parasites are organisms whose parasitic mode of life is limited to a few or even one stage of development. [2] An example of this is the larval stage of harvest mites, while the adult stage is non-parasitic.
The parasite may live outside of the host ectoparasite ; for example, a tick. Alternatively, the parasite may live within the host endoparasite ; for example, the fluke. An obligate parasite that does not live directly in or on the host, but rather acts at a distance – for example, a cuckoo which hatches and is raised by non-relatives – is known as a brood parasite.
In order to establish infestation in a susceptible host, obligate parasites must evade defences before, during and after entry into the host. [7] Due to the wide range of obligate parasite types, it is impossible to identify a general invasion strategy. Intracellular parasites use various strategies to invade cells and subvert cellular signalling pathways. Most bacteria and viruses undergo passive uptake, where they rely on the host cell for uptake. However, apicomplexans engage in active entry. [8] One obligate wasp parasite, Polistes atrimandibularis , infiltrates its hosts' colony by modifying its chemical signature to match that of the hosts. [9] This tricks the host wasps into thinking the parasite is one of their own.
A number of obligate intracellular parasites have evolved mechanisms for evading their hosts' cellular defences, including the ability to survive in distinct cellular compartments. [10] One of the mechanisms that hosts employ in their attempt to reduce the replication and spread of pathogens is apoptosis (programmed cell death). Some obligate parasites have developed ways to suppress this phenomenon, for example Toxoplasma gondii , although the mechanism is not yet fully understood. [11]
Changes in a host’s behaviour following infection with obligate parasites are extremely common. [12] Unusual behaviour observed in infected individuals is noted, and if its complexity suggests that this behaviour will benefit the transmission of the parasite, then this is said to be an example of adaptive manipulation. [13] However, there is a difficulty in demonstrating changes in behaviour are the result of a selective process favouring transmission of the parasite. [14] It has been suggested that these changes may merely be a side-effect of infection. [15] Most behaviour changes have not been demonstrated to lead to fitness gains in either the host or the parasite. [16] An example of this behaviour is the attraction of rats to cat urine after infection with Toxoplasma gondii . [17] However, the "scientific metaphors, including anthropomorphisms" sometimes used in "popular media and the scientific literature" to describe the manipulation of host behavior have been described as "catchy, yet misleading". [18]
In some cases the behaviour we observe in an organism is not due to the expression of its genes, but rather to the genes of parasites infecting it. This behaviour is an extended phenotype. [13]
Three main evolutionary routes have been suggested for the appearance of host behaviour manipulation by parasites. The first is a parasite driven scenario of manipulation, while the second and third are host driven scenarios of manipulation.
It has been suggested that extended phenotype behaviours are not adaptive, but are Exaptative. [21] While they may have a benefit for the parasitic organism, they did not arise with the intention of this benefit. [20]
The cowbird and cuckoo require the nests and parental care of other passerines in order for their young to fledge. These are known as brood parasites. The parasitic bird species mimics egg patterns and colours of the host species, which reduces egg rejection. [22] The chicks of some species are able to manipulate host behaviour by making rapid calls that mimic the sound made by up to four of the host chicks. [23] Mimicry of the host species also occurs in the paper wasp species Polistes semenowi and Polistes sulcifer and the bumblebee species Bombus bohemicus , with the parasite changing its proportions of cuticular hydrocarbons, species- and colony-specific identifying chemicals, to match that of the usurped host species. [4] [24] [25]
Several butterfly species will also exhibit brood parasitic behavior. An example is Niphanda fusca, a butterfly that will release cuticular hydrocarbons (CHCs) to trick the host ant, C. japonicus, into adopting the larva as their own in their own nest. The ant will then raise the larva of the butterfly, feeding it directly from mouth-to-mouth, until it pupates. [26]
It is proposed that this mimicry has evolved through two processes: either as coevolutionary responses to host defences against brood parasites or modifying pre-existing host provisioning strategies. [27] Competition between the parasite and host young for parental resources might lead to exaggeration of the aspects of the signal that most effectively exploit host parents. [28] The parasitic young are likely to experience stronger selection for exaggerated signals than host young, because they are unrelated to the other chicks in the nest and therefore under selection to behave more selfishly. [29]
Current theory in evolutionary biology indicates that host-parasite relationships may evolve towards equilibrial states of severe disease. [30] This differs from the conventional belief that commensalism is the ideal equilibrium for both the host and parasite. [1]
Parasitism is a close 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 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.
Toxoplasmosis is a parasitic disease caused by Toxoplasma gondii, an apicomplexan. Infections with toxoplasmosis are associated with a variety of neuropsychiatric and behavioral conditions. Occasionally, people may have a few weeks or months of mild, flu-like illness such as muscle aches and tender lymph nodes. In a small number of people, eye problems may develop. In those with a weak immune system, severe symptoms such as seizures and poor coordination may occur. If a person becomes infected during pregnancy, a condition known as congenital toxoplasmosis may affect the child.
The Extended Phenotype is a 1982 book by the evolutionary biologist Richard Dawkins, in which the author introduced a biological concept of the same name. The book’s main idea is that phenotype should not be limited to biological processes such as protein biosynthesis or tissue growth, but extended to include all effects that a gene has on its environment, inside or outside the body of the individual organism.
Toxoplasma gondii is a parasitic protozoan that causes toxoplasmosis. Found worldwide, T. gondii is capable of infecting virtually all warm-blooded animals, but felids are the only known definitive hosts in which the parasite may undergo sexual reproduction.
In biology and medicine, a host is a larger organism that harbours a smaller organism; 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, cells harbouring pathogenic (disease-causing) viruses, or 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.
Kleptoparasitism is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, such as when food is scarce or when victims are abundant. Many kleptoparasites are arthropods, especially bees and wasps, but including some true flies, dung beetles, bugs, and spiders. Cuckoo bees are specialized kleptoparasites which lay their eggs either on the pollen masses made by other bees, or on the insect hosts of parasitoid wasps. They are an instance of Emery's rule, which states that insect social parasites tend to be closely related to their hosts. The behavior occurs, too, in vertebrates including birds such as skuas, which persistently chase other seabirds until they disgorge their food, and carnivorous mammals such as spotted hyenas and lions. Other species opportunistically indulge in kleptoparasitism.
A supernormal stimulus or superstimulus is an exaggerated version of a stimulus to which there is an existing response tendency, or any stimulus that elicits a response more strongly than the stimulus for which it evolved.
Brood parasitism is a subclass of parasitism and phenomenon and behavioural pattern of animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own, usually using egg mimicry, with eggs that resemble the host's. The strategy involves a form of aggressive mimicry called Kirbyan mimicry.
Coccidia (Coccidiasina) are a subclass of microscopic, spore-forming, single-celled obligate intracellular parasites belonging to the apicomplexan class Conoidasida. As obligate intracellular parasites, they must live and reproduce within an animal cell. Coccidian parasites infect the intestinal tracts of animals, and are the largest group of apicomplexan protozoa.
A rhoptry is a specialized secretory organelle. They are club-shaped organelles connected by thin necks to the extreme apical pole of the parasite. These organelles, like micronemes, are characteristic of the motile stages of Apicomplexa protozoans. They can vary in number and shape and contain numerous enzymes that are released during the process of host penetration. The proteins they contain are important in the interaction between the host and the parasite, including the formation of the parasitophorous vacuole (PV).
Intracellular parasites are microparasites that are capable of growing and reproducing inside the cells of a host. They are also called intracellular pathogens.
Bombus bohemicus, also known as the gypsy's cuckoo bumblebee, is a species of socially parasitic cuckoo bumblebee found in most of Europe with the exception of the southern Iberian Peninsula and Iceland. B. bohemicus practices inquilinism, or brood parasitism, of other bumblebee species. B. bohemicus is a generalist parasite, successfully invading several species from genus Bombus. The invading queen mimics the host nest's chemical signals, allowing her to assume a reproductively dominant role as well as manipulation of host worker fertility and behavior.
Neospora hughesi is an obligate protozoan apicomplexan parasite that causes myelitis and equine protozoal myeloencephalitis (EPM) in horses, and has only been documented in North America. EPM is a neurological disease from lesions in the spinal cord, brain stem, or brain from parasites such as N. hughesi or Sarcocystis neurona. Signs that a horse may have EPM include ataxia, muscle atrophy, difficulty swallowing, and head tilt. There are antiprotozoal drugs, such as the 28-day course of ponazuril, to treat the disease, as well as anti-inflammatories to alleviate neurologic symptoms
Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism is typified by a cellular variety with a distinct morphology and biochemistry.
Behavior-altering parasites are parasites capable of causing changes in the behavior of their hosts species to enhance their transmission, sometimes directly affecting the hosts' decision-making and behavior control mechanisms. By way of example, a parasite that reproduces in an intermediate host may require, as part of their life cycle, that the intermediate host be eaten by a predator at a higher trophic level, and some parasites are capable of altering the behavior of the intermediate host to make such predation more likely; a mechanism that has been called parasite increased trophic facilitation or parasite increased trophic transmission. Examples can be found in bacteria, protozoa, viruses, and animals. Parasites may also alter the host behavior to increase protection of the parasites or their offspring; the term bodyguard manipulation is used for such mechanisms.
Hammondia hammondi is a species of obligate heteroxenous parasitic alveolates of domestic cats. Intracellular cysts develop mainly in striated muscle. After the ingestion of cysts by cats, a multiplicative cycle precedes the development of gametocytes in the epithelium of the small intestine. Oocyst shedding persists for 10 to 28 days followed by immunity. Cysts in skeletal muscle measure between 100 and 340 μm in length and 40 and 95 μm in width. Some of the intermediate hosts develop low levels of antibody and some cross-immunity against Toxoplasma.
Polistes austroccidentalis is a kleptoparasitic paper wasp that is found in several regions of high altitude in Europe, and until 2017 was universally mistakenly referred to as Polistes semenowi, which is instead the correct name of the species formerly known as "Polistes sulcifer". As one of only four obligate parasites in the subgenus Polistes, it uses the nests of other paper wasps to rear its young. To evade detection by the host nest, P. austroccidentalis employs mimicry by adjusting its cuticular hydrocarbons to match those of the host. Once the host nest has been infiltrated, the parasitic female physically attacks the host queen to subdue her and become the colony's new queen. P. austroccidentalis displays several morphological adaptations for parasitism such as increased mandible size and an enlarged Van der Vecht's organ. This species is unusual because it does not have the ability to produce workers and is only able to produce individuals who have the capacity to reproduce.
Polistes semenowi is a species of paper wasp in the genus Polistes that is found in southeastern and southern central Europe, as well as central Asia, and was until 2017 erroneously known by the name Polistes sulcifer, while a different species was incorrectly believed to represent P. semenowi. It is one of only four known Polistes obligate social parasites, sometimes referred to as "cuckoo paper wasps", and its host is the congeneric species Polistes dominula. As an obligate social parasite, this species has lost the ability to build nests, and relies on the host workers to raise its brood. P. semenowi females use brute force, followed by chemical mimicry in order to successfully usurp a host nest and take over as the queen.
The name cuckoo paper wasp refers to a monophyletic species group of brood-parasitic paper wasps in the genus Polistes. This species group contains only four species; Polistes atrimandibularis, P. austroccidentalis, P. maroccanus, and P. semenowi, all of them obligate social parasites of other Polistes species.
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.