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A phytobiome consists of a
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclu ...
(phyto) situated in its specific ecological area (
biome A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
), including its environment and the associated communities of
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
s which inhabit it. These organisms include all macro- and
micro-organism 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 living in, on, or around the plant including
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
, archaea,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
, protists,
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
,
animals Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
, and other
plants Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
. The environment includes the
soil Soil, also commonly referred to as earth or dirt Dirt is an unclean matter, especially when in contact with a person's clothes, skin, or possessions. In such cases, they are said to become dirty. Common types of dirt include: * Debri ...
,
air The atmosphere of Earth is the layer of gases, known collectively as air, retained by Earth's gravity that surrounds the planet and forms its planetary atmosphere. The atmosphere of Earth protects life on Earth by creating pressure allowing f ...
, and
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
. Examples of ecological areas are fields,
rangeland Rangelands are grasslands, shrublands, woodlands, wetlands, and deserts that are grazed by domestic livestock or wild animals. Types of rangelands include tallgrass and shortgrass prairies, desert grasslands and shrublands, woodlands, sava ...
s,
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s. Knowledge of the interactions within a phytobiome can be used to create tools for agriculture, crop management, increased health, preservation,
productivity Productivity is the efficiency of production of goods or services expressed by some measure. Measurements of productivity are often expressed as a ratio of an aggregate output to a single input or an aggregate input used in a production proces ...
, and sustainability of cropping and forest systems.


Signaling


Diversity

The microbial community within the phytobiome is perhaps one of the most rich and diverse microbiomes on Earth. Plants form associations with billions of organisms across every kingdom of life. Recent metagenomic and metatranscriptomic approaches have allowed scientists to discover novel taxonomic species that are not easily cultured in a laboratory.


Bacteria

Recent research has shown that inter-kingdom communication between organisms is essential for proper phytobiome function. There are numerous physical and chemical signals such as secreted lipids, peptides and polysaccharides that allow organisms to recognize and interact within the phytobiome. Bacteria are known to produce
quorum sensing In biology, quorum sensing or quorum signalling (QS) is the ability to detect and respond to cell population density by gene regulation. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at ...
molecules such as homoserine lactones, lipid-like diffusable factors and signaling peptides that mediate plant-bacteria interactions such as colonization. Homoserine lactones have been are reportedly produced by a large number of bacteria found in the
rhizosphere The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microo ...
. Plant growth promoting bacteria (PGPBs) often produce Nod factors (nodulation factors) that initiate nodule formation in plants. In addition to plant-bacterial interactions, bacteria often secrete bactericidal or fungicidal compounds into the phytobiome to reduce local competition for niches and resources. Additionally, organisms that feed on bacteria such as some species of algae and protists are attracted to these small signaling molecules.


Phages

Bacteriophages also play a critical role in the phytobiome through
predator-prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
interactions. Bacteriophages use a signaling peptides such as
arbitrium Arbitrium is a viral peptide produced by bacteriophages to communicate with each other. It is six amino acids long, and is produced when a phage infects a bacterial host. It signals to other phages that a host has been infected. Discovery Arbitr ...
to mediate the initiation of cell lysis and lysogeny in the host cell.


Fungi

Fungi communicate in the phytobiome through chemical signaling to aid in sexual reproduction,
sporulation In biology, a spore is a unit of sexual 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, ...
, cell-to-cell recognition and antibiosis; however, only a fraction of these chemicals have been studied for their function.
Mycorrhizal   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the pla ...
fungi establish symbiotic relationships with plants through the production of Myc factors, or chitooligosaccharides that are recognized by receptors in the plant. Nematode-trapping fungi often utilize fungal signaling molecules to initiate morphogenesis towards prey. Other organisms can interfere with fungal signaling, such as plant-produced
oxylipins Oxylipins constitute a family of oxygenated natural products which are formed from fatty acids by pathways involving at least one step of dioxygen-dependent oxidation. Oxylipins are derived from polyunsaturated fatty acids (PUFAs) by COX enzymes ( ...
that mimic fungal signaling molecules and can regulate fungal development or reduce
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 ...
. Multiple species of bacteria, insects and nematodes have all been reported to respond to fungal signaling compounds.


Nematodes

Very little is known about nematode communication within the phytobiome. Plant-pathogenic nematodes often communicate through production of pheromones. Plants can detect these compounds and induce defense pathways. Nematodes also produce plant hormones such as
cytokinins Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and le ...
that aid in the establishment of association with plants.


Protists

Perhaps even less is known about the ecological role of protists and viruses within the phytobiome. Some
amoebae An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopo ...
species use
cyclic nucleotides A cyclic nucleotide (cNMP) is a single-phosphate nucleotide with a cyclic bond arrangement between the sugar and phosphate groups. Like other nucleotides, cyclic nucleotides are composed of three functional groups: a sugar, a nitrogenous base, ...
or peptide signals to adapt social behavior.
Phytohormones Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
produced by algae-associated bacteria can greatly impact microalgae populations in the soil. The presence of amoeba can also trigger the bacterium '' P. fluorescens'' to produce anti-amoebal toxins.


Insects

Insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
communicate to transfer information regarding external threats, social status, food availability and mating through the production of volatile
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
, also known as semiochemicals. This has made pheromones a subject of research since the 1950s for various applications in
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people t ...
and insect-vectored diseases such as
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...
. Plants can have profound impacts on insect pheromone production. Rattlebox plants produce various
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
compounds that insects use as a precursor for sex pheromone synthesis. Many plant species have evolved production of volatile chemicals that interfere with pheromone signaling, often through inhibition of proper olfactory neuron function. Bacteria and fungi can also produce volatile chemicals that affect insect behavior.


Plants

The presence of plants and their communication with other community members fundamentally shapes the phytobiome. Root exudates contain numerous sugars, amino acids, polysaccharides and
secondary metabolites Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the nor ...
. The production of these exudates is heavily influenced by environmental factors and plant physiology and can alter the community composition of the rhizosphere and rhizoplane. The secretion of
flavonoids Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
helps to recruit '' Rhizobia'' bacteria that form a mutualistic symbiosis with numerous plant species. ''Rhizobia'' can also recognize other plant compounds such as betaines, aldonic acids and jasmonic acid. These signal molecules can have multiple or even counteracting effects. For example, plant cutins trigger
arbuscular mycorrhizal An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscules. ( ...
colonization and symbiosis but can also be recognized by plant-pathogenic
oomycetes Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the result ...
and trigger
pathogenesis Pathogenesis is the process by which a disease or disorder develops. It can include factors which contribute not only to the onset of the disease or disorder, but also to its progression and maintenance. The word comes from Greek πάθος ''pat ...
. Plant volatile chemicals also attract herbivores, pollinators and seed carriers. When plants recognize the presence of microbes, they often activate the production of phytohormone signals that are transported throughout the plant. Plants respond to pathogens and herbivores through production of hormones including salicylic acid, jasmonic acid and ethylene. In addition, many phytohormones that function in abiotic stress tolerance or plant growth also trigger responses with the microbial community. The production of salicylic acid in '' Arabidopsis'' was shown to influence the
root microbiome The root microbiome (also called rhizosphere microbiome) is the dynamic community of microorganisms associated with plant roots. Because they are rich in a variety of carbon compounds, plant roots provide unique environments for a diverse assemblag ...
composition by acting as a signal or carbon source. Secretion of
strigolactone Strigolactones are a group of chemical compounds produced by a plant's roots. Due to their mechanism of action, these molecules have been classified as plant hormones or phytohormones. So far, strigolactones have been identified to be responsible f ...
is known to stimulate spore germination and Myc factor production in arbuscular mycorrhizal fungi. The microbial community can also manipulate phytohormone function or the production of specific phytohormones in plants.


Research

In 2015 the
American Phytopathological Society The American Phytopathological Society (APS) is an international scientific organization devoted to the study of plant diseases (phytopathology). APS promotes the advancement of modern concepts in the science of plant pathology and in plant healt ...
(APS) launched a research framework, the Phytobiomes Initiative, to facilitate the organization of research into phytobiome. As part of this effort, in 2016 it launched '' Phytobiomes Journal'', an
open-access journal Open access (OA) is a set of principles and a range of practices through which research outputs are distributed online, free of access charges or other barriers. With open access strictly defined (according to the 2001 definition), or Gratis v ...
. The journal focuses on transdisciplinary research that impacts the entire plant ecosystem. An overall research strategy has been published in the Phytobiomes Roadmap, a document developed by a group of scientific societies, companies, research institutes, and governmental agencies. It is intended to present a strategic plan to study phytobiomes and propose an action plan to apply phytobiome studies. The connected Phytobiomes Alliance is an international, nonprofit consortium of academic institutions, large and small companies, and governmental agencies coordinating public-private research projects on various aspects of agriculturally relevant phytobiomes.


See also

*
Plant microbiome The plant microbiome, also known as the phytomicrobiome, plays roles in plant health and productivity and has received significant attention in recent years.. Material was copied from this source, which is available under Creative Commons Attri ...
*
Soil biology Soil biology is the study of microbial and faunal activity and ecology in soil. Soil life, soil biota, soil fauna, or edaphon is a collective term that encompasses all organisms that spend a significant portion of their life cycle within a soil ...


References


External links

* {{cite web , title=Phytobiomes , website=phytobiomes.org , date=2018-06-21 , url=http://www.phytobiomes.org/Pages/default.aspx , ref={{sfnref , Phytobiomes , 2018 , access-date=2018-06-21 Plant ecology Biomes Agriculture Plant products