Plant–Microbe Communications for

Symbiosis
Definition :

Symbiosis is a biological phenomenon involving dynamic changes in the genome, metabolism and
signaling network, and a multidirectional comprehension of these interactions is required when
studying symbiotic organism.

Two symbiotic systems have been actively studied for many years.

One is arbuscular mycorrhizal (AM) symbiosis and the other is root nodule (RN) symbiosis.
Arbuscular mycorrhizal symbiosis :

AM symbiosis is probably the most widespread interaction between plants and microbes, in the
context of phylogeny and ecology (Kistner and Parniske 2002, Bonfante and Genre 2010). More than
80% of all land plant families are thought to have a symbiotic relationship with AM fungi that
belong to the Glomeromycota. The origin of AM symbiosis is thought to be in the early Devonian
period, approximately 400 million years ago. Thus, AM symbiosis is also called the mother of plant
root endosymbioses (Parniske 2008).
Root nodule symbiosis :

On the other hand, RN symbiosis involves morphogenesis and is formed by communication

Plant microbe interaction :

Plants are non-motile but they constantly encounter both the biotic and abiotic stress. There is a
constant war between the pathogenic microbes and the host plant – the outcome of which determines
resistance or disease.

Plants secrete various organic compounds resulting in a nutritionally enriched environment favorable
for microbial growth.Microbial communities influence plants in direct and indirect ways.

Plants and microbes can have variety of interactions including pathogenic, symbiotic and associative.

In plants, symbionts play significant roles in plant growth, development, and health by providing
nutrients and increasing tolerance to biotic and abiotic stress factors. For example, endophytic fungi
are very common in plants and many studies have shown that they increase growth and development
of colonized plants (Rodriguez, White, Arnold, & Redman, 2009). This is mainly achieved by promoting
root growth through the allocation of resources to the expanding root system before stem growth
(Rodriguez, Freeman, McArthur, Kim, & Redman, 2009). Indeed, evidence suggests that some of these
effects are due to epigenetic effects of plant symbiotic fungi or bacteria on their hosts. Conversely,
during the course of the establishment of endosymbiosis, genetic and/or epigenetic changes occur in
the interacting microorganisms with plant hosts.
Symbiotic relationships :

Symbiosis refers to relationships between organisms of different species that show an intimate
association with each other. Symbiotic relationships provide at least one of the participating species
with a nutritional advantage.

Three types of symbiosis have been recognized depending on the nature of relationship.

1. Mutualism
2. Commensalism
3. Parasitism

Mutualism :

Mutualism is a biological interaction between two species wherein both species benefit from each
other.

• This term was discovered by Pierre van Benden.

• They exchange food or provide shelter or protection, but may still be able to live independent life.

Mutualism describes the ecological interaction between two or more species where fitness is
increased through direct interaction. Mutualism is thought to be the most common type of ecological
interaction, and it is often dominant in most communities worldwide.

Mutualism plays a key part in ecology. For example, mutualistic interactions are vital for terrestrial
ecosystem function as more than 48% of land plants rely on mycorrhizal relationships with fungi to
provide them with inorganic compounds and trace elements. As another example, the estimate of
tropical forest trees with seed dispersal mutualisms with animals ranges from 70–90%. In addition,
mutualism is thought to have driven the evolution of much of the biological diversity we see, such as
flower forms (important for pollination mutualisms) and co-evolution between groups of species.
However mutualism has historically received less attention than other interactions such as predation
and parasitism.

Examples :

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.

There are many different examples of mutualistic relationship :

• Plants and Microbes : eg. Rhizobium in root nodules.

• Protists and Fungi : eg. Lichen

• Terrestrial plants and insects : eg. Pollination

Ocellaris clownfish and Ritter's sea anemones is a mutual service-service symbiosis, the fish driving off
butterflyfish and the anemone's tentacles protecting the fish from predators.
Commensalism :

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. 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 consonant with its habits, as in the remoras
that ride attached to sharks and other fishes. Both remora and 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.

Example :

Aspergillus and Staphylococcus:

Numerous genera of bacteria and fungi live on and in the human body as part of its natural flora.
Aspergillus is capable of living under considerable environmental stress, and thus is capable of
colonising the upper gastrointestinal tract where relatively few examples of the body's gut flora can
survive due to highly acidic or alkaline conditions produced by gastric acid and digestive juices. While
Aspergillus normally produces no symptoms, in individuals who are immunocompromised or suffering
from existing conditions such as tuberculosis, a condition called aspergillosis can occur, in which
populations of Aspergillus grow out of control.

Staphylococcus aureus, a common bacterial species, is known best for its numerous pathogenic
strains that can cause numerous illnesses and conditions. However, many strains of S. aureus are
metabiotic commensals, and are present on roughly 20 to 30% of the human population as part of the
skin flora. S. aureus also benefits from the variable ambient conditions created by the body's mucous
membranes, and as such can be found in the oral and nasal cavities, as well as inside the ear canal.
Other Staphylococcus species including S. warneri, S. lugdunensis and S. epidermidis, will also engage
in commensalism for similar purposes .

Tillandsia bourgaei growing on an oak tree in Mexico.

Ammensalism :

Ammensalism is the ecological interaction in which an individual species harms another without
obtaining benefit.
• This type of symbiotic relationship is common, but not considered as an important process
structuring communities because they are accidental and do not benefit the species doing the harm.

Ammensalism examples :

Algal blooms can lead to the death of many species of fish and other animals, however the algae do
not benefit from the deaths of these individuals.

• Black walnut trees secretes a chemical from its roots which prevents the growth of neighboring
trees.

• Elephants stepping on ants or leveling brush does not benefit the elephant but harms the ants and
brush.

Algal bloom

Black Walnut trees secrete a substance into the soil called juglone that inhibits the growth of
neighboring plants.
African Elephant With Bird On Back Walking On Savanna By Mountain Kenya showing the example of
Ammensalism.

Parasitism :

In evolutionary biology, parasitism is a relationship between species, where one organism, the
parasite, lives on or in another organism, the host, causing it some harm, and is adapted structurally
to this way of life. Parasites are therefore chemoautotrophs. This relationship is detrimental to the
host, however a true parasite does normally not kill its host.

Examples :

A hemiparasite or partial parasite, such as mistletoe derives some of its nutrients from another living
plant, and a holoparasite such as dodder derives all of its nutrients from another plant. Parasitic plants
make up about one per cent of angiosperms and are in almost every biome in the world. All these
plants have modified roots, haustoria, which penetrate the host plants, connecting them to the
conductive system either the xylem, the phloem, or both. This provides them with the ability to extract
water and nutrients from the host.

Classification based on host :

A parasitic plant is classified depending on where it latches onto the host, either the stem or the root,
and the amount of nutrients it requires. Since holoparasites have no chlorophyll and therefore cannot
make food for themselves by photosynthesis, they are always obligate parasites, deriving all their food
from their hosts.( Heide-Jørgensen, Henning S. (2008).

Releasing of chemicals :

Some parasitic plants can locate their host plants by detecting chemicals in the air or soil given off by
host shoots or roots, respectively. About 4,500 species of parasitic plant in approximately 20 families
of flowering plants are known. Nickrent, D. L.; Musselman, L. J. (2004). ,Heide-Jørgensen, Henning S.
(2008).

Interaction of soil with parasitic members of family Orobanchaceae :

Species within Orobanchaceae (broomrapes) are some of the most economically destructive of all
plants. Species of Striga (witchweeds) are estimated to cost billions of dollars a year in crop yield loss,
infesting over 50 million hectares of cultivated land within Sub-Saharan Africa alone. Striga infects
both grasses and grains, including corn, rice and sorghum, undoubtedly some of the most important
food crops. Orobanche also threatens a wide range of other important crops, including peas,
chickpeas, tomatoes, carrots, and varieties of cabbage. Yield loss from Orobanche can be total; despite
extensive research, no method of control has been entirely successful.

Parasitic fungi :

Parasitic fungi derive some or all of their nutritional requirements from plants, other fungi, or animals.
Unlike mycorrhizal fungi which have a mutualistic relationship with their host plants, they are
pathogenic.

For example,

1. the honey fungi in the genus Armillaria grow in the roots of a wide variety of trees, and
eventually kill them. They then continue to live in the dead wood, feeding saprophytically.
2. Fungal infection (mycosis) is widespread in animals including humans; it kills some 1.6 million
people each year.
3. Microsporidia are obligate intracellular parasitic fungi that can also be hyperparasites. They
largely affect insects, but some affect vertebrates including humans, where they can cause
the intestinal infection microsporidiosis.

Mistletoe : a parasitic plant

Parasitic plant
The honey fungus, Armillaria mellea, is a parasite of trees, and a saprophyte feeding on the trees it
has killed.

Plant -symbionts interaction examples :

Symbionts have been demonstrated to contribute significantly to plant tolerance to stress and this
could be regarded as an epigenetic plasticity trait.

Example 1:

The endophytic fungus Piriformospora indica confers drought tolerance in plants by colonizing their
roots. In Arabidopsis, it was shown that in P. indica-colonized plants several genes were upregulated
earlier and in higher quantities compared to uncolonized plants (Sherameti, Tripathi, Varma, &
Oelmüller, 2008). Among the upregulated genes was histone acetyltransferase, which is responsible
for histone acetylation. Higher histone acetylation is associated with more active transcription. It is
likely that by induction of this gene early, P. indica regulates expression of a variety of genes involved
in stress response, which leads to priming the aerial parts of the plant to be more prepared for water
stress.

Example 2 :

Induced Resistance by Trichoderma spp. :

Trichoderma spp. facilitate root colonization of their hosts by the production and regulation of
hormonal signals. For example, Trichoderma strains that promote plant growth are found to produce
the plant hormone auxin (Gravel et al., 2007; Contreras-Cornejo et al., 2009; Hoyos-Carvajal et al.,
2009; Zhang et al., 2012) that promotes root growth. Auxin-induced modifications in root architecture
(e.g. increased number of root hairs etc.), increases total absorptive surface of the roots, thereby
facilitating nutrient uptake resulting to increased plant growth (Contreras-Cornejo et al., 2009;
Samolski et al., 2012). Trichoderma spp. equally utilize a vast array of proteins to facilitate root
colonization.

Once inside the roots, Trichoderma spp. must suppress or tolerate plant defense mechanisms in order
to facilitate root invasion. Trichoderma spp. are found to tolerate toxicants like antibiotics, plant
antimicrobial compounds, and synthetic chemicals or contaminants.

Example 3:

Role of parasitic fungi :

In the terrestrial environment, fungi are of fundamental importance as decomposers, plant pathogens
and symbionts (mycorrhizas), playing important roles in carbon, nitrogen and other biogeochemical
cycles (Wainwright, 1988). They are often dominant in acidic conditions and, in soil, can comprise the
largest pool of biomass (including other microorganisms and invertebrates) (Metting, 1992). This,
combined with their branching filamentous explorative growth habit and high surface area to mass
ratio ensures that fungal–metal interactions are an integral component of major environmental
cycling processes. Metals, both essential and inessential, and their derivatives can interact with fungi
in various ways depending on the metal species, organism and environment, while fungal metabolic
activities can also influence speciation and mobility (Gadd, 1993; Wainwright and Gadd, 1997). Certain
mechanisms may mobilize metals into forms available for cellular uptake and leaching from the
system, e.g. complexation with organic acids, other metabolites and siderophores (Francis, 1994),
while immobilization may result from sorption on to cell components and exopolymers, transport and
intracellular and extracellular sequestration or precipitation (Morley and Gadd, 1995; Gadd, 1996;
Sayer and Gadd, 1997; White et al., 1997). Such apparently opposing processes of solubilization and
immobilization are important for biogeochemical cycles for indigenous or introduced metals, and
fundamental determinants of fungal growth, morphogenesis and physiology (Morley et al., 1996;
Ramsay et al., 1999). Furthermore, several processes are relevant to environmental bioremediation
(White et al., 1997; Sayer etal., 1998).

Example 4:

Mycorrhizas:

Fungi are another crucial, but often under-appreciated part of all forest ecosystems. Mycorrhizas are
symbiotic relationships between certain fungi and the roots of plants. The fine fungal threads (called
hyphae) either ensheath or penetrate the host plant's roots. The fungus helps the plant to extract
nutrients and water from the soil, and also protects its host against harmful organisms. In return, and
in common with lichens, the fungus receives sugars via the plant's photosynthesis.

As with most mutualistic relationships, each partner grows better in association with the other than it
would individually. Birch (Betula spp.) has a number of these partnerships, the most familiar being
with the red and white fly agaric (Amanita muscaria), as well as with the chanterelle (Cantharellus
cibarius). Scots pine has mycorrhizal associations with over 200 species of fungi in Scotland, including
another kind of chanterelle (Cantharellus lutescens). In fact, the majority of plants in the Caledonian
Forest benefit from mycorhizzal relationships, and it is thought that mycorrhizas helped plants to
colonise the land, millions of years ago.

Example 5 :

Symbiotic bacteria:

Symbiosis works on many different scales, as is clearly illustrated by the relationship between alder
(Alnus glutinosa) and a bacterium (Frankia alni). In this case, Frankia lives within special nodules on
the roots of the alder (another example of endosymbiosis), and absorbs nitrogen from the
atmosphere, 'fixing' it in the soil. This benefits the alder, which via photosynthesis provides the
bacteria with sugars. The soil becomes enriched as a result of this process, and alder has been used in
ecological restoration projects in various parts of the world, to restore depleted soils.

Example 6 :

Wood Ants

Wood ants (Formica spp.) have symbiotic relationships with a number of other organisms in the forest.
Some species of flowering plant in the forest depend on ants for their dispersal. Cow-wheat
(Melampyrum pratense) seeds have a fatty attachment on them called an elaiosome. The ants take
the seeds to their nests, and feed the elaiosome to their larvae, thus helping to disperse the plant. In
areas that have been deforested, such plants cannot return easily without the aid of ants, and this
understanding is crucial when attempting to restore woodland ground flora to areas that have been
deforested for a long time.

Example 7 :

Pollination:

Many of these relationships are difficult to see, but pollination is a form of symbiosis that can be
observed quite easily. There are many flowering plants in the Caledonian Forest. Flowers act as
powerful advertisements to insects, offering energy-rich nectar. The visiting insect, having fed upon
the sugary liquid, then goes on to carry pollen to fertilise other flowers, benefiting the overall
population of that particular plant species.

Some insects are fairly specific in their choice of plant. Certain bee species have a longer 'tongue' than
others, and this affects their choice of flower. The three banded white-tail bumblebee (Bombus
hortorum) (a species found in Glen Affric) for example, chooses deeper flowers such as foxglove
(Digitalis purpurea). The shorter-tongued bees can only drink nectar from flowers that are not as deep,
such as raspberry (Rubus idaeus) and goat willow (Salix caprea).

Butterflies are also very visible in their role as pollinators. For example, the pearl-bordered fritillary
(Boloria euphrosyne) can be found in open deciduous woodland where it visits spring flowers such as
bugle (Ajuga reptans), birds-foot trefoil (Lotus corniculatus) or dandelion (Taraxacum officinale), for
their nectar.

Pollination probably evolved in response to early insects eating the pollen itself. Plants that offered
organic matter (ie nectar) as an alternative to pollen for insects to feed on would increase their
reproductive success - the pollen was not only spared, but carried from plant to plant. Insects could
still feed, and flowering plants evolved and thrived.

References :

Miller, Allie. "Intricate Relationship Allows the Other to Flourish: the Sea Anemone and the Clownfish".
AskNature. The Biomimicry Institute. Retrieved 15 February 2015.

Haskell, E. F. (1949). A clarification of social science. Main Currents in Modern Thought 7: 45–51.

Burkholder, P. R. (1952) Cooperation and Conflict among Primitive Organisms. American Scientist, 40,
601-631. link.

Bronstein, J. L. (2015). The study of mutualism. In: Bronstein, J. L. (ed.). Mutualism. Oxford University
Press, Oxford.

Haskell, E. F. (1949). A clarification of social science. Main Currents in Modern Thought 7: 45–51.

Burkholder, P. R. (1952) Cooperation and Conflict among Primitive Organisms. American Scientist, 40,
601-631. link.

Bronstein, J. L. (2015). The study of mutualism. In: Bronstein, J. L. (ed.). Mutualism. Oxford ULatorre,
A.; Durban, A.; Moya, A.; Pereto, J. (2011). The role of symbiosis in eukaryotic evolution. Origins and
evolution of life – An astrobiological perspective. pp. 326–339.niversity Press, Oxford.

"Symbiosis". Bloomsbury Guide to Human Thought. London: Bloomsbury Publishing Ltd, 1993. Credo
Reference. Web. 17 September 2012.