Oral Ecology
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Oral ecology is the
microbial ecology Microbial ecology (or environmental microbiology) is the ecology of microorganisms: their relationship with one another and with their environment. It concerns the three major domains of life—Eukaryota, Archaea, and Bacteria—as well as viru ...
of the
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s found in
mouth In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
s. Oral ecology, like all forms of
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
, involves the study of the living things found in oral cavities as well as their interactions with each other and with their environment. Oral ecology is frequently investigated from the perspective of oral disease prevention, often focusing on conditions such as dental
caries Tooth decay, also known as cavities or caries, is the breakdown of teeth due to acids produced by bacteria. The cavities may be a number of different colors from yellow to black. Symptoms may include pain and difficulty with eating. Complicatio ...
(or "cavities"),
candidiasis Candidiasis is a fungal infection due to any type of '' Candida'' (a type of yeast). When it affects the mouth, in some countries it is commonly called thrush. Signs and symptoms include white patches on the tongue or other areas of the mouth ...
("thrush"),
gingivitis Gingivitis is a non-destructive disease that causes inflammation of the gums. The most common form of gingivitis, and the most common form of periodontal disease overall, is in response to bacterial biofilms (also called plaque) that is attached ...
,
periodontal disease Periodontal disease, also known as gum disease, is a set of inflammatory conditions affecting the tissues surrounding the teeth. In its early stage, called gingivitis, the gums become swollen and red and may bleed. It is considered the main caus ...
, and others. However, many of the interactions between the microbiota and oral environment protect from disease and support a healthy oral cavity. Interactions between microbes and their environment can result in the stabilization or destabilization of the oral
microbiome A microbiome () is the community of microorganisms that can usually be found living together in any given habitat. It was defined more precisely in 1988 by Whipps ''et al.'' as "a characteristic microbial community occupying a reasonably well ...
, with destabilization believed to result in disease states. Destabilization of the microbiome can be influenced by several factors, including diet changes, drugs or
immune system disorders An immune disorder is a dysfunction of the immune system. These disorders can be characterized in several different ways: * By the component(s) of the immune system affected * By whether the immune system is overactive or underactive * By whether t ...
.


History

Bacteria were first detected under the microscope of Dutch scientist
Anton van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the F ...
in the late 17th century from his own healthy human oral sample. After using this technology on a healthy sample, Leeuwenhoek applied his tool to the decayed tooth matter of his wife, where he noted that the organisms present were highly similar to those found in cheese. These are believed to likely have been lactic acid bacteria, however the link between bacterial acid production and tooth decay was not further uncovered until much later. After this discovery and the further development of microscopy, bacteria was found within tooth cavities by multiple scientists throughout the 19th century. Willoughby Miller was the first recorded oral microbiologist, and he performed much of his foundational microbiology research in the laboratory of famed microbiologist
Robert Koch Heinrich Hermann Robert Koch ( , ; 11 December 1843 – 27 May 1910) was a German physician and microbiologist. As the discoverer of the specific causative agents of deadly infectious diseases including tuberculosis, cholera (though the Vibrio ...
. In this time, Miller generated the chemo-parasitic (also referred to as "acidogenic") theory of caries, which proposed that tooth decay is initiated by bacterial acid production on the surface of teeth. This theory is considered to be foundational to the field of dentistry as well as oral ecology, by drawing connections between the activities of microbial entities and its effects on their non-living microscopic environment. In ecological terms, early work in oral microbiology largely falls into a category of microbial research now described as "reductionist", generally meaning it focused heavily on the isolation of individual microbes before observation or testing. It wasn't until the late 20th century that "holistic" approaches to oral microbiology were coming into the mainstream, and thus microbial ecology was intentionally studied. Holistic microbiology considers not only an organism of interest but also the biological and abiotic context in which the organism naturally is found. Scientist Philip Marsh is credited with developing the ecological plaque hypothesis in 1994, in which he ideated that
dental plaque Dental plaque is a biofilm of microorganisms (mostly bacteria, but also fungi) that grows on surfaces within the mouth. It is a sticky colorless deposit at first, but when it forms tartar, it is often brown or pale yellow. It is commonly found bet ...
can be both normal and healthy as well as "cariogenic" (creates cavities), depending on the microbial community (or "
consortia A consortium (plural: consortia) is an association of two or more individuals, companies, organizations or governments (or any combination of these entities) with the objective of participating in a common activity or pooling their resources for ...
") present in the biofilm and the community's stability. Furthermore, in his theory, Marsh links the exposure of nonliving environmental influences on the microbial community to the selection and change in microbial constituents that can cause cariogenic conditions.


Oral environment

Teeth, saliva, and oral tissues are the major components of the oral environment in which the oral microbiome resides. Much like most environments, some of the oral environment, such as teeth and
saliva Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can be ...
, are
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
(non-living), and some are living, such as the host immune system or host mouth mucosal tissues- including gums, cheek ("buccal") and tongue (when present).


Abiotic

Saliva is holds multiple roles in oral ecology; for one example, it creates physical disturbance to microbes through a washing action. Increase in saliva flow via stimulation (i.e. chewing gum) has been shown to diminish cariogenic plaque formation. Saliva is also largely responsible for environmental pH, water content, nutrients, and host-produced immune cells and
antimicrobial An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
s. One major antimicrobial found in saliva (as well as mucus) is
lysozyme Lysozyme (EC 3.2.1.17, muramidase, ''N''-acetylmuramide glycanhydrolase; systematic name peptidoglycan ''N''-acetylmuramoylhydrolase) is an antimicrobial enzyme produced by animals that forms part of the innate immune system. It is a glycoside ...
, an enzyme that shears bacterial cells. Another critical role saliva plays in the microscopic environment is supplying the
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glycos ...
s bacteria use to cling to the surface of teeth. Teeth are another example of the abiotic environmental factors involved in oral ecology. Bacteria settle on the tooth surface as a solid substrate on which to grow. Compared to floating in saliva, bacteria on teeth gain environmental stability in that they will experience a consistent environment of temperature, relative oxygen exposure, nutrient density, physical disturbances, etc. While teeth provide stability to the microbial community, the overgrowth of bacteria is known to result in tooth decay primarily due to acid production from sugar-consuming fermentative metabolisms. Some organisms associated with this condition are ''
lactobacilli The ''Lactobacillaceae'' are a family of lactic acid bacteria. It is the only family in the lactic acid bacteria which includes homofermentative and heterofermentative organisms; in the ''Lactobacillaceae,'' the pathway used for hexose fermentati ...
'', which produce the lactic acid that breaks down tooth enamel. As a result, host diet also influences the ecology of the mouth by altering saliva pH and nutrient content, and as a result the microbial life that interacts with the oral environment. Oxygen content is a major variable that can influence the type of microbial flora present in the oral cavity. This variable is slightly unique to the oral cavity due to its exposure to the outside of the host body. In ecology, niches are a set of conditions that can be associated with the presence of a certain organism. Thus, oxygen concentration variation throughout the mouth can be a factor in niche differentiation within this environment. At the microscopic scale, oxygen concentration can dictate where in the mouth
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cellu ...
,
anaerobic Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to: * Anaerobic adhesive, a bonding a ...
,
facultative anaerobic A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococc ...
,
aerotolerant Aerotolerant anaerobes use fermentation to produce ATP. They do not use oxygen, but they can protect themselves from reactive oxygen molecules. In contrast, obligate anaerobes can be harmed by reactive oxygen molecules. There are three categorie ...
, or
microaerophilic A microaerophile is a microorganism that requires environments containing lower levels of dioxygen than that are present in the atmosphere (i.e. < 21% O2; typically 2–10% O2) for optimal growth. A more re ...
microbes grow or form biofilm. Biofilms themselves can also help regulate oxygen exposure and keep anaerobic organisms at the interior, adding to the complexity of the niches within the oral cavity. Another abiotic environmental influence on oral ecology includes the use of drugs, especially
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention of ...
s and orally-administered antibiotics. Antibiotics can kill oral bacteria as well as cause secondary environmental effects such as a decrease in saliva, leading to further changes in the abiotic microenvironment. The destabilization of the bacteria in a microbiome which results in disease is known as bacterial dysbiosis. For example, the destabilization of the bacterial community in the mouth can lead to a bloom in fungal communities, resulting in diseases such as thrush. Furthermore, the development of antibiotic-resistant populations in response to the treatment can result in an overpopulation of the resistant bacteria after treatment is completed, disturbing the relative abundances found pre-treatment.  


Biotic (non-bacterial)

The host of the oral cavity in which the oral ecology is studied is also of importance. This is an example of a biotic, or living, environmental factor. General host health and immune system function is critical to oral microflora, as it determines which microbes are able to survive in the mouth. The
innate immune system The innate, or nonspecific, immune system is one of the two main immunity strategies (the other being the adaptive immune system) in vertebrates. The innate immune system is an older evolutionary defense strategy, relatively speaking, and is the ...
, which operates in animals continuously regardless of the presence of disease, is most relevant due to its constant role in oral ecology both in healthy and unhealthy hosts. This includes the production of free-floating
antibodies An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of the ...
, macrophages, and other immune cells present in saliva. At a healthy, stable state, the host immune system permits the colonization of certain microbes by not targeting them. This can be described as "immune equilibrium", or the conditions where the host and the microbiota in the oral microbiome symbiose.


Human


Bacterial

In microbial ecology, the principle of
priority effect In ecology, a priority effect is an impact that a particular species can have on community development due to prior arrival at a site. There are two basic types: An ''inhibitory priority effect'' which occurs when a species that arrives first at ...
refers to the competitive advantage some microorganisms gain by colonizing a surface first. It is generally believed that primary colonization occurs by transmission from the mother or their breastmilk (
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 ...
), as well as the environment of the newborn (
horizontal transmission 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, symbiont ...
). It has been found that at different locations in the oral cavity, different microbes are early colonizers. The very initial colonizers of teeth are considered to be ''
Streptococcus ''Streptococcus'' is a genus of gram-positive ' (plural ) or spherical bacteria that belongs to the family Streptococcaceae, within the order Lactobacillales (lactic acid bacteria), in the phylum Bacillota. Cell division in streptococci occurs ...
,'' a genus of bacteria that are usually
facultative anaerobes A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...
that can grow in both aerobic and anaerobic conditions. This is advantageous in an environment that is variably exposed to oxygen throughout the day as well as throughout the oral cavity. Despite over 700 unique species of bacteria being associated with the human mouth, in tooth plaque only between 7-9 "major players" have been repeatedly identified as early colonizers, including ''
Actinomyces ''Actinomyces'' is a genus of the Actinomycetia class of bacteria. They all are gram-positive. ''Actinomyces'' species are facultatively anaerobic and they grow best under anaerobic conditions. ''Actinomyces'' species may form endospores, an ...
'', ''
Streptococcus ''Streptococcus'' is a genus of gram-positive ' (plural ) or spherical bacteria that belongs to the family Streptococcaceae, within the order Lactobacillales (lactic acid bacteria), in the phylum Bacillota. Cell division in streptococci occurs ...
'', ''
Neisseria ''Neisseria'' is a large genus of bacteria that colonize the mucosal surfaces of many animals. Of the 11 species that colonize humans, only two are pathogens, '' N. meningitidis'' and ''N. gonorrhoeae''. ''Neisseria'' species are Gram-negativ ...
'', and ''
Veillonella ''Veillonella'' are Gram-negative bacteria (Gram stain pink) anaerobic cocci, unlike most Bacillota, which are Gram-positive bacteria. This bacterium is well known for its lactate fermenting abilities. It is a normal bacterium in the intestine ...
'' species. It is believed that the colonization of these specific genera of bacteria influence the stability and
homeostasis In biology, homeostasis (British English, British also homoeostasis) Help:IPA/English, (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physics, physical, and chemistry, chemical conditions maintained by organism, living systems. Thi ...
of the resulting oral microflora. This colonization occurs by the construction of and adhesion to a pellicle made of glycoproteins from host saliva. Upon adhesion to the pellicle, early colonizing bacteria begin to produce the biofilm intended to anchor the colony to the tooth. As is common in microbiomes, this biofilm does not remain a single genera or species. In fact, the vast majority of relevant microbes perform co-aggregation within a biofilm. However, it is understood that not all microbes will co-aggregate together, and ammensal activity does occur between specific species, such as ''S. mutans'' and ''
P. gingivalis ''Porphyromonas gingivalis'' belongs to the phylum ''Bacteroidota'' and is a nonmotile, Gram-negative, rod-shaped, anaerobic, pathogenic bacterium. It forms black colonies on blood agar. It is found in the oral cavity, where it is implicated i ...
''. The interbacterial interactions as well as the interactions with the host teeth, oxygen conditions, and saliva are what compose bacterial oral ecology.  


Nonbacterial

Bacteria, while being the most abundant, are not the only kind of microbiota present in the oral cavity. Fungal/yeast cells are also present, particularly including the genus '' Candida''. The yeast species ''C. albicans'' and '' C. tropicalis'' are known as commensals in the human mouth, which means that they are a part of normal flora that engages in a mutually-beneficial relationship with its host. They are the most abundant non-bacterial microbes isolated from the human mouth. As described in the above section, co-aggregation within a biofilm is not uncommon, including the cohabitation of yeasts with bacteria. ''Candida albicans'' is known to selectively participate in "dual-species" biofilms with certain species of ''Streptococcus'' bacteria through the actual attachment of the yeast to the bacterial cell surface. This allows the yeast to be anchored to the tooth surface indirectly to gain stability. Some other, but significantly less abundant, non-bacterial microbes in the human mouth include the fungi genera ''
Cryptococcus ''Cryptococcus'' is a genus of fungi in the family Cryptococcaceae that includes both yeasts and filamentous species. The filamentous, sexual forms or teleomorphs were formerly classified in the genus ''Filobasidiella'', while ''Cryptococcus'' ...
,
Aspergillus ' () is a genus consisting of several hundred mold species found in various climates worldwide. ''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Miche ...
,'' and ''
Fusarium ''Fusarium'' is a large genus of filamentous fungi, part of a group often referred to as hyphomycetes, widely distributed in soil and associated with plants. Most species are harmless saprobes, and are relatively abundant members of the soil mi ...
''.


References


Further reading

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