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Horizontal transmission is the transmission of organisms between biotic and/or abiotic members of an ecosystem that are not in a parent-progeny relationship. This concept has been generalized to include transmissions of infectious agents, symbionts, and cultural traits between humans. Because the evolutionary fate of the agent is not tied to reproductive success of the host, horizontal transmission tends to evolve
virulence Virulence is a pathogen's or microorganism's ability to cause damage to a host. In most, especially in animal systems, virulence refers to the degree of damage caused by a microbe to its host. The pathogenicity of an organism—its ability to c ...
. It is therefore a critical concept for
evolutionary medicine Evolutionary medicine or Darwinian medicine is the application of modern evolutionary theory to understanding health and disease. Modern biomedical research and practice have focused on the molecular and physiological mechanisms underlying heal ...
.


Biological


Pathogen transmission

In biological, but not cultural, transmissions the carriers (also known as vectors) may include other species. The two main biological modes of transmission are ''anterior station'' and ''posterior station''. In anterior station, transmission occurs via the bite of an infected organism (the vector), like in
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects humans and other animals. Malaria causes Signs and symptoms, symptoms that typically include fever, fatigue (medical), tiredness, vomiting, and headaches. In se ...
,
dengue fever Dengue fever is a mosquito-borne tropical disease caused by the dengue virus. Symptoms typically begin three to fourteen days after infection. These may include a high fever, headache, vomiting, muscle and joint pains, and a characteristic ...
, and
bubonic plague Bubonic plague is one of three types of plague caused by the plague bacterium (''Yersinia pestis''). One to seven days after exposure to the bacteria, flu-like symptoms develop. These symptoms include fever, headaches, and vomiting, as well ...
. Posterior station is transmission via contact with infected feces. Examples are rickettsiae driven diseases (like
typhus Typhus, also known as typhus fever, is a group of infectious diseases that include epidemic typhus, scrub typhus, and murine typhus. Common symptoms include fever, headache, and a rash. Typically these begin one to two weeks after exposure. ...
), which are contracted by a body louse's fecal material being scratched into the bloodstream. The vector is not necessarily another species, however. For example, a dog infected with
Rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
may infect another dog via anterior station transmission. Moreover, there are other modes of biological transmission, such as generalized bleeding in
ebola Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after becom ...
.


Symbiont transmission

Symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
describes a relationship in which at least two organisms are in an intimately integrated state, such that one organism acts a
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ...
and the other as the symbiont. There are obligate, those that require the host for survival, and facultative symbionts, those that can survive independently of the host. Symbionts can follow
vertical Vertical is a geometric term of location which may refer to: * Vertical direction, the direction aligned with the direction of the force of gravity, up or down * Vertical (angles), a pair of angles opposite each other, formed by two intersecting s ...
, horizontal, or a mixed mode of
transmission Transmission may refer to: Medicine, science and technology * Power transmission ** Electric power transmission ** Propulsion transmission, technology allowing controlled application of power *** Automatic transmission *** Manual transmission ** ...
to their host. Horizontal, or lateral, transmission describes the acquisition of a facultative symbiont from the environment or from a nearby host. The life cycle of the host includes both symbiotic and aposymbiotic phases. The aposymbiotic phase generally begins in the
germ cell Germ or germs may refer to: Science * Germ (microorganism), an informal word for a pathogen * Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually * Germ layer, a primary layer of cells that forms during embr ...
s, and during development, the host organism acquires the symbiont and translocates it to a symbiont-housing organ. The host will release the symbiont before reproduction. Determining the transmission mode of a symbiont requires identifying the key aspects that define transmission. For horizontal transmission one would need to check the host at each life stage for the presence of the symbiont and determine whether the symbiont is released from the host into the environment. In hydrothermal vent tubeworms, release of a symbiont upon host death was demonstrated using a symbiont recruitment plates and fluorescence ''in situ'' hybridization (FISH). Each life cycle stage of the tubeworm was sampled and examined also using FISH to determine the aposymbiotic and acquisition life cycle stages as larvae and settled larvae >250μm in length. Implications of horizontal transmission include the critical need for specificity in recognition and acquisition methods and the larger genetic diversity maintained by the symbiont species. Recognition specificity can be achieved through complex signaling through the release of hormones, such as with Rhizobia species and legumes. The release of the symbiont allows it to exchange genetic material with external microbes as well as between the genetically diverse individuals within the host. This maintaining of genetic exchange allows for the symbiont's host range to be extended and the selection for new functionality or adaptations of hosts, symbionts, and
holobiont A holobiont is an assemblage of a host and the many other species living in or around it, which together form a discrete ecological unit through symbiosis, though there is controversy over this discreteness. The components of a holobiont are i ...
. An example of a horizontally transmitted symbiont with a high specificity recognition and acquisition method is the
Rhizobia Rhizobia are diazotrophic bacteria that fix nitrogen after becoming established inside the root nodules of legumes (Fabaceae). To express genes for nitrogen fixation, rhizobia require a plant host; they cannot independently fix nitrogen. In ...
-
Legume A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock for ...
symbiosis. The establishment of the symbiosis begins with the aposymbiotic plant releasing flavinoids that are detected by a specific Rhizobium species and triggers the induction of ''nod'' genes in the bacterium. Nod factors, which are
lipooligosaccharide Lipopolysaccharides (LPS) are large molecules consisting of a lipid and a polysaccharide that are bacterial toxins. They are composed of an O-antigen, an outer core, and an inner core all joined by a covalent bond, and are found in the outer me ...
signals, are released as a result of the expression of the ''nod'' genes and when detected by the host plant initiate
root nodule Root nodules are found on the roots of plants, primarily legumes, that form a symbiosis with nitrogen-fixing bacteria. Under nitrogen-limiting conditions, capable plants form a symbiotic relationship with a host-specific strain of bacteria known a ...
formation which eventually trap the Rhizobia. The established symbiosis can be further contained in a
symbiosome A symbiosome is a specialised compartment in a host cell that houses an endosymbiont in a symbiotic relationship. The term was first used in 1983 to describe the vacuole structure in the symbiosis between the animal host the ''Hydra'', and the end ...
in which the Rhizobia symbionts reside and carry out the nitrogen-fixation.


Anthropological

Cultural transmission may also be horizontal which is explicitly reified in
Dual Inheritance Theory Dual inheritance theory (DIT), also known as gene–culture coevolution or biocultural evolution, was developed in the 1960s through early 1980s to explain how human behavior is a product of two different and interacting evolutionary processes: ge ...
. Horizontal transmission is implicit in the meme theory of cultural evolution, where the "meme" has been characterized by
Richard Dawkins Richard Dawkins (born 26 March 1941) is a British evolutionary biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An ath ...
as a " Virus of the Mind".


See also

* Horizontal disease transmission * Horizontal gene transmission *
Vertical transmission Vertical transmission of symbionts is the transfer of a microbial symbiont from the parent directly to the offspring.  Many metazoan species carry symbiotic bacteria which play a mutualistic, commensal, or parasitic role.  A symbiont is acquire ...


References

{{reflist, 30em Infectious diseases