Topic 3: The root-soil interface/Rhizosphere

#Discussion points
o Ion concentration, rhizosphere pH, root exudates.

o Microbial activity in the rhizosphere

o Mycorrhizae (ecto and endo)

i) Infection and energetics
ii) Phosphorous uptake, other effects
iii) Mycorrhizal dependency and utility
#What is a rhizosphere
Rhizo – roots Sphere – field of influence

o The rhizosphere is the narrow region of soil that is directly influenced by root
secretions and associated soil microorganisms.

o Soil which is not part of the rhizosphere is known as bulk soil.

o The rhizosphere is a micro-ecological zone in direct proximity of plant roots.

o It is functionally defined as the particulate matter and microorganisms that

cling to roots after being gently shaken in water.

o The theoretical extent of the rhizosphere is dependent on the zone of influence

of the plant roots and associated microorganisms.
 #Rhizosphere

A=Amoeba consuming bacteria BL=Energy limited bacteria BU=Non-energy limited

bacteria RC=Root derived carbon SR=Sloughed root hair cells F=Fungal hyphae
N=Nematode worm
o The rhizosphere is a metabolically busier, faster moving, more competitive

environment than the surrounding soil.

o The rhizosphere contains many bacteria that feed on sloughed-off plant cells,

termed rhizodeposition, and the proteins and sugars released by roots.

o Protozoa and nematodes that graze on bacteria are also more abundant in the
rhizosphere.

o Microbes are most abundant where the integrity of the root is compromised.

o For this reason, rhizoplane microorganisms tend to be found on older rather than
younger roots.
#Root exudates - secretions
o Plants secrete many compounds into the rhizosphere which serve different functions.

o Theses can be primary or secondary metabolites

o Primary metabolites include; amino acid, soluble sugars, lipids, organic acids,
vitamins, enzymes, plant DNA, etc.

o These are low molecular weight compounds, most off-which act as messengers.

o Secondary metabolites include; mucilage, enzymes, etc

o Most secondary metabolite are produced in the Golgi apparatus in root cap cell, but
mucilage is produced by bacterial degradation of root epidermal cells
#Functions of root secretions

1. Defence of rhizosphere against pathogenenic microbes

2. Attract higher numbers of beneficial organisms

3. Helps to keep the soil moist

4. To obtain nutrients – enzymes

5. Changes soil chemical properties

6. Inhibits the growth of other competing spp.

#Examples:

o Symbiotic nitrogen fixing bacteria, such as the Rhizobium species, detect compounds
secreted by the roots of leguminous plants which initiate rhizoidal infection.

o Strigolactones are detected by mycorrhizal fungi, stimulate germination of spores and

initiate changes in the mycorrhizae that allow it to colonize the root.

o The parasitic plant, Striga also detects the presence of strigolactones and will germinate
when it detects them.

o Some plants secrete allelochemicals which inhibit the growth of other organisms, for
example, garlic mustard produces a chemical which is believed to prevent mutualisms
forming between the trees and mycorrhizae in North American temperate forests.
#Enzymes in the rhizosphere

o Are produced by both plants and microbes:

1. Oxidoreductases- transfer electrons from one molecule to another.

2. Hydrolases- nutrient mineralisation, protein breakdown.

3. Lyases- degrade cell walls.

4. Transferases- transfer the functional group from one molecule to

another.
## Physical Environment

1. Water status – water potential

o The plant roots which the rhizosphere is associated with can effect the
physical environment of the rhizosphere.

o As plants transpire water with more force during the day than during the
night, they change the soil water potential immediately near their roots and
so the rhizosphere undergoes fluctuations that the bulk soil avoids.
2. Soil Texture

o Movement of organic matter away from the root as well as bacteria

colonizing new locations occurs more readily in sandy soils than clay
soils.

o Sand has larger pores between each granule which allow free
movement of microorganisms and exudates, therefore, the larger the
granule size, the further the rhizosphere and microorganisms associated
with it will extend into the surrounding soil.

o Plant roots compact the soil on the short term as they grow, and once
they die and decay, can actually aid the soil to be less porous.
3. Soil pH

o Several factors can lower the pH in the rhizosphere, for instance, respiration leads to carbon

dioxide, and eventually to bicarbonate/carbonic acid generation.

o In addition to respiration of the roots themselves, the rhizosphere is very rich in carbon,

resulting in other organisms (e.g., prokaryotes, eukaryotes & other small animals living

organisms) respiring in the rhizosphere more than in the bulk soil.

o The available habitat that microbes can inhabit are limited in part by pH of the soil.

o Fungi prefer more acidic soils, while bacteria have a very broad pH spectrum where they can
survive.

o The influencing effects of pH in the rhizosphere is critical in supporting a biologically diverse

microbial community.
 #Organisms in the rhizosphere
1. Bacteria
o Bacteria are the most ubiquitous organisms in the soil, averaging between
10^6 to 10^9 organisms per gram of rhizosphere soil.
o Due to their small mass, they only account for a small amount of the biomass
of soil.
o Non-sporulating rods and pseudomonads are the most common bacteria in the
soil.

2. Fungi
o Both pathogenic and symbiotic fungi associate with the rhizosphere.
o They average between 10^5 and 10^6 organisms per gram of rhizosphere soil.
o Zygomycetes and Hyphomycetes establish the most readily in the rhizosphere
because they metabolize simple sugars.
#Biotic Interactions in the Rhizosphere
o A range of interactions are present in the rhizosphere from beneficial symbiotic relationships to detrimental
pathogenic interactions:
#General Impacts of Microbes on Rhizosphere
o Chemical changes occur as a result of humification of organic matter.
o The resultant mineralization of various organic compounds (e.g., phosphorous, sulfur &
nitrogen) provides plants with forms of nutrition that are readily available for uptake.
o The turnover of microbial populations also results in the release of nutrients.
o The fixation of atmospheric dinitrogen by both asymbiotic and symbiotic bacteria results in
increases in the available nitrogen pool that can be accessed by plants in and near the
rhizosphere.
o Symbiotic mycorrhizae cause an increase in the effective rooting area of plants, thereby
providing added nutrient mining capabilities to the plant.
o Rhizosphere microbes can also release plant growth regulators.
o Physical changes occur primarily through the production of extracellular polymeric
substances such as polysaccharides which improve soil aggregation and soil texture.
o The presence of mucigel in the rhizoplane is crucial to the water relations of plants,
providing a bridge that prevents desiccation by maintaining the water column during water
stress events.
o Plant roots cause chemical and physical changes to the soil they inhabit and these changes will
affect the microbial diversity in and around the rhizosphere.

o Root exudates will select for/against certain populations of microorganisms.

o Many plants exhibit genetic resistance/tolerance to rhizosphere microorganisms; the genetic

make-up of the plant will determine, in part, the community makeup of the microorganisms in
the rhizospere.

o The ability of a plant to form symbiotic relationships with soil microbes will also determine
rhizosphere microbial populations.

o The age and health of the plants present will also play a role in the microbial community
dynamics of the rhizosphere.

o In some cases, plants can also compete with rhizosphere microorganisms for resources like water
and nutrients.
#Mycorrhizae

Gk. mykós, "fungus" riza, "roots

o Is a symbiotic association between a fungus and the roots of a vascular

plant

o Mycorrhizae are mutualistic symbiotic associations established between

the roots of about 90% of the land plants and a great number of soil fungi.
#Symbiotic or Mutualistic Relationships

o In such a relationship, the plant usually provides a source of carbon and the bacteria or

fungus fills some other more specialized function.

o In many cases the association is not absolutely necessary for the survival of both members,

but provides significant benefit to both.

o The microbial partner can also help the plant to survive in an otherwise inhospitable

environment.

o For example, specific endophytes have been shown to confer heat resistance to grasses that

grow in a thermal area, or salt tolerance to costal grasses.

#Plant infection by fungus

o 1st contact between partners

o 2nd changes of the cell surface

o 3rd development of interfacial compartments in order to increase the exchange area

o 4th modification of protein expression

#Types of mycorrhizae
o Mycorrhizae (fungal infections) are commonly divided into
Ectomycorrhizae and Endomycorrhizae.

o The two types are differentiated by the fact that the hyphae of
Ectomycorrhizae fungi (during infection) do not penetrate
individual cells within the root.

o While the hyphae of endomycorrhizae fungi (during infection)

penetrates the cell wall and invaginate the cell membrane.
#Ectomycorrhizae
o Ectomycorrhizae interact with the plant root by producing a
net-like structure called a Hartig net that weaves between the
root cortical cells.

o A sheath or mantle of fungal tissue covering part or all of the

root is responsible for most of the increase in surface area
caused by mycorrhizae.

o Some fungi, such as Boletus betulicola, have only a narrow

range of plants they can associate with.

o Other fungi have a much larger range of host plant, for

example, Pisolithus tinctorius associates with 46 tree species.
#Arbuscular mycorrhizae
o Endomycorrhizae are variable and have been further classified as
arbuscular, ericoid, arbutoid, monotropoid and orchid mycorrhizae.

o Arbuscular mycorrhizae (AM) were formerly known as vesicular-

arbuscular mycorrhizae (VAM).

o Hyphae enter into the plant cells, producing structures that are either
balloon-like (vesicles) or dichotomously-branching invaginations
(arbuscules).

o The fungal hyphae do not in fact penetrate the protoplast (i.e. the interior
of the cell), but invaginate the cell membrane.

o The structure of the arbuscules greatly increases the contact surface area
between the hyphae and the cell cytoplasm to facilitate the transfer of
nutrients between them.
o Direct connection between the plant and fungal cytoplasm
allows the transfer of nutrients from the fungi to the plant and
carbon from the plant to the fungi to be more efficient.

o Arbuscular mycorrhizae are formed only by fungi in the

division Glomeromycota.

o The hyphae of AM fungi produce the glycoprotein glomalin,

which may be one of the major stores of carbon in the soil.

o Arbuscular mycorrhizal fungi have (possibly) been asexual for

many millions of years and, unusually, individuals can contain
many genetically different nuclei (a phenomenon called
heterokaryosis).
#Mycorrhizae Energetics (Mutualistic benefits)
o This mutualistic association provides the fungus with relatively constant
and direct access to carbohydrates, such as glucose and sucrose which are
energy sources.

o The carbohydrates are translocated from their source (usually leaves) to

root tissues and on to the plant's fungal partners.

o In return, the plant gains the benefits of the mycelium's higher absorptive
capacity for water and mineral nutrients

o The comparatively large surface area of mycelium –to- root ratio improves
the plant's mineral absorption capabilities.
#Phosphorous uptake
o This element, like nitrogen gas, can not be absorbed by the plant in its
elemental form.

o First, it must be combined to form the orthophosphate ion (H2PO4 -)

before uptake.

o Increasing pH may limit the availability of phosphorus to the plant.

o Plant roots alone may be incapable of taking up phosphate ions that are
de-mineralized in soils with a slightly basic pH.

o The mycelium of the mycorrhizal fungus can, however, access these

phosphorus sources and make them available to the plants they colonize.

o The mechanisms of increased absorption are both physical and chemical.

o Mycorrhizal mycelia are much smaller in diameter than the
smallest root, and thus can explore a greater volume of soil,
providing a larger surface area for absorption.

o Also, the cell membrane chemistry of fungi is different from

that of plants (includes organic acid excretion which aids in
ion displacement)

o Mycorrhizae are therefore especially beneficial for the plant

partner in nutrient-poor soils
#Dependency and utility
oMycorrhizal dependency can be defined as the degree to which a
plant is dependent on Mycorrhizal conditions to produce its
maximum yield.

oTo determine the MD of a particular spp, there is need to compare

the growth of the mycorrhizal and non-mycorrhizal spp.

oPlant utility denotes the various benefits to the plant of having

mycorrhizal associations and these include:

1. Disease and drought resistance

oMycorrhizal plants are often more resistant to diseases, such as
those caused by microbial soil-borne pathogens and are also more
resistant to the effects of drought.
2. Colonization of barren soil
o Plants grown in sterile soils and growth media often perform
poorly without the addition of spores or hyphae of mycorrhizal
fungi to colonize the plant roots and aid in the uptake of soil
mineral nutrients.

o The absence of mycorrhizal fungi can also slow plant growth

in early succession or on degraded landscapes.

o The introduction of alien mycorrhizal plants to nutrient-

deficient ecosystems puts indigenous non-mycorrhizal plants
at a competitive disadvantage.
#Resistance to toxicity
o Fungi have been found to have a protective role for plants rooted in soils
with high metal concentrations such as acidic and contaminated soils.

o Pine trees inoculated with Pisolithus tinctorius planted in several

contaminated sites displayed high tolerance to the prevailing
contaminant, survivorship and growth.

o One study discovered the existence of Suillus luteus strains with varying
tolerance of zinc.

o This was probably due to binding of the metal to the extrametrical

mycelium of the fungus without affecting the exchange of beneficial
substances.
#Recap
o The plant genotype shape the structure of the rhizosphere microbial community, but
edaphic factors and biogeography also have a great influence
***END***