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LECTURE 2.

RHIZOSPHERE
SYSTEM

Rhizosphere system

Root function/rhizosphere
Diversity and Ecology of Associated
Microbes
Rhizodeposition or Plant-Derived
compounds from plant roots
Root Colonization
Microbe-plant interactions in the
rhizosphere
Effect of biological activity in rhizosphere
on plant

PLANT ROOTS

Root Anatomy

Root Functions
Anchoring
Absorption of nutrients
storage

Root renewal

Perpetually changing
In young plantselongation and branching;

primary root a few cm to 9 cm per day


Secondary roots grow slower, Constantly renewed
Radicles die a few days after formation
In cotton 50% of root tissue disappeared in 100
days
Equilibrium between formation of new roots and
disappearance of old roots during last 2/3 of
growth.

Ripening stage, production of new roots stop,


followed by dimunition of root mass.

Importance of roots
Interaction between roots and
microbes
Dead root cells are precious nutritive
resources for soil organisms

Rhizosphere system
Zone or volume of soil under the
influence of roots
Extend > 5mm from roots, area of
increased microbial activities
Not uniform; consist of several
gradients that change as distance
from root increases
Approximately defined as the soil that
remains attached to roots when plant
is carefully uprooted, then gently
shaken.

Root anatomy

Epidermis
Cortex
Endodermis
Root hairs extension of epidermal cells,
important for ion uptake
Root cap- protect the meristem as root
moves through soil; produce mucilage and
substance that enhance plant cell growth
Rhizosphere boundaries: rhizosphere,
rhizoplane, endophytes

Rhizosphere - the area of soil directly influenced


by plant roots, variable. Soils that remains after
shaking. 1 - 4 mm or more from root surface

Rhizoplane - surface of plant root, root hairs


present large surface area (> 6 m2 for wheat
plant). Only 4-10 % of rhizoplane is in direct
contact with soil microbes. About 2 mm width.

Difficult to distinguish rhizosphere & rhizoplane


due to mucilage

Roots primary, root hairs- increase surface


area for absorption of ions
Soil-root interface has high microbial
community

Energy Flux of rhizosphere


Increased microbial populations due to abundance
energised substrates
A portion of compounds photosynthesised in aerial
parts is transported to roots through phloem
The carbon fixed; is transferred to roots; some
stored as reserves or incorporated into new
tissue; consumed for cell respiration (CO2); lost to
external environment (Rhizodeposition/root
exudates)

Rhizosphere Environment
Physical and chemical properties of
soil:Soil texture, structure, moisture,
temperature, pH, fertilization,
etc.

Diversity and Ecology of


Associated Microbes
Microbial Populations of Rhizosphere
Vast number of microbial species
Bacteria, fungi, algae, protozoa, nematode,
viruses
Variable nutrition patterns:- autotrophs,
chemoautotrophs, heterotrophs,
chemoheterotrops
Saprophytes, parasites, associative,
symbionts
Oxygen requirements, temperature,
physiology

Population decrease with increasing


distance
Rhizosphere/bulk soil (R/S) ratio
estimate how strongly the rhizosphere
affects organisms
determine the ability of organisms to
colonize rhizosphere (candidate as
inoculant)
R/S between 5 and 20, can also be >100

Bacteria
Most numerous typically 106 - 109 g -1
rhizosphere soil
Large R/S ratio
Abundance of non-sporulating rods
Pseudomonads and Gram - negative bacteria
are competitive and occupy large portion of
rhizosphere
Gram-positive rods & cocci is lower
Actinomycetes has smaller R/S ratio

Fungi
Considerable fungal growth along root
surfaces ; 12-14 mm hyphae mm -2 of root
surface
105 to 106 g 1 soil

Composition of microflora of
rhizosphere
Vary depending on the family of plants;
legumes have high R/S ration than
crucifers
Vary with plant age; high effect at
vegetative growth compared to
reproductive and maturation stage
Replacing plant chromosome will change the
microbial population
Dependent on root exudates

pH of Rhizosphere
Differs from pH of adjacent soil; about 2 pH
units
Depend on soil buffering capacity and plant
types
More acidic than soil; pH gradient along roots
Proton (H+) are excreted when cell elongate
Electric current circulating towards inside of
growing roots and towards outside of older
zones

Cont.

Absorption of mineral ions coupled to


hydrolysis of ATP by membrane ATPase;
transfer of proton from cytoplasm
towards outside cell creates a pH and
electric potential gradient
Concentration of H+ is not constant depend
on nature of ions taken up by roots; eg.
Fertilizer NH4+ (pH decrease) or NO3- (pH
increase)

Rhizodeposition or Plant-Derived
compounds from plant roots

a. Exudates- Low molecular wt. compounds


that leak from all cells into the intercellular
spaces then to the soil. Not metabolically
mediated
b. Secretions- Low and high mol. wt.
mucilages. Result of metabolic process
c. Mucilages
Originate in root cap
Hydrolysates of cell wall and sloughed root cap
cells
Secreted by epidermal cell
Produced by bacterial degradation of old, dead
epidermal cells

c. Mucigel- Gelatinous material at root


surface;
Natural and modified plant mucilages,
bacterial cells and metabolites, colloidal
mineral and organic matter.
Important in maintaining contact between root
and soil
Found on root/root hairs than root tip

d. Lysates compounds released by lysis of


epidermal cells.

Exudate composition
(water soluble, insoluble, volatile)
Sugars, amino acids, vitamins, tannin,
Alkaloid, phosphatides, growth factors,
flourescent substance, stimulants,
inhibitors.
Composition varies with plant genome and
stage of development and environmental
factors (light, temperature, water, redox
potential, minerals, phytotoxicity,
reduction in leaf surface (by insect or
diseases)

Functions of root exudate:


Provide carbon , energy and nutrients
Sugars- sources of carbon for microbial growth
Amino acids- source of N
Organic acids- metal chelation, affects pH,
absorption and translocation of nutrients
Vitamins- influence bacterial population,
species.

Exudation Rate and factors affecting


a. Gradients of release; greatest amount of
exudate from cells involved in cell elongation
and lateral root formation. Concentration
declines exponentially with increase in distance
from roots.
b. Can be increased by extreme temperature,
water stress, phosphorus deficiency, increase
light intensity, increase microbial population,
herbicides, pathogens, foliar treatments,
symbiotic associations.
c. Can be reduced by N deficiency, decrease light
intensity, decrease microbial population

Root Colonization
Factors influencing rhizosphere colonization
Microbial characteristics:- nutrient
requirements, early growth rate, cellulase
production, antibiotic tolerance and
production, siderophore production,
tolerance to fungicides or chemicals.
Plant characteristics :- plant species, plant
age, plant genetics, foliar treatments
Environment:- soil type, texture, moisture,
atmosphere, temperature, fertility,
pesticides applied

Only 7 -15 % root surface occupied by


microbes
Rhizoplane bacteria scattered in small
colonies in holes, tears, crevices,
junctions of epidermal cells in roots
Recognition signals

Colonization on rhizoplane
Mostly by epiphytes and saprophytes
Not distributed uniformly; only 10% of surface of
young roots; ~ 37% in older roots
Colonies extend very little compared to root
elongation
Ability of bacteria to disperse along root is limited
Implanted at the junction of epidermal cells and at
base of absorbent hairs where exudation is most
intense and protective mucilage is thickest
Live on senescent cells of cortical cells, epidermal
cells

Endophytes
Microbes can penetrate into central
cylinder of roots
Entrance through rapture of endodermis
during emergence of secondary roots
Minimal colonization by microbes except
when attacked by pathogens
Bacteria can move from roots to stems and
leaves

Microbe-plant interactions in
the rhizosphere
N2-fixation

Mycorrhizal associations
Plant growth promotion
Decomposition
nutrient cycling

Rhizospheric affinity/signals
Microbes receive signals from plants that triggers
chemotaxis process directed towards roots
Signals consist of mainly sugars and amino acids
that diffuse in soil solution
Rapid respond resulting in successful colonization
Lectins in legumes root hair tips important in early
stages of nodulation, it may bind the Rhizobium to
the root tips
N-acyl homoserine lactones in root exudate
regulate expression of bacterial genes for quarum
sensing in plant-microbe interaction

Rhizocompetence
The ability of microbes to grow,
function and maintain itself in
rhizosphere;
through production of antibiotic
Resistant to foreign antibiotics
Optimal utilization of available molecules

Microbiostasis and effect of


exudates
Microbiostasis or dormancy of microbes
Root exudate affect germination of
clamydospores
- stimulate parasites and symbionts
- eg. Root exudate of Pinus sp. trigger
germination of ectomycorrhizae spore of
Suilus sp. but not other fungal species
- hatching of parasitic nematode eggs

Microbial Movement towards


host
Attraction of microbes to roots is
associated with amino acids, fatty
acids, aldehydes, alcohols
Eg. Luteoline (alfalfa seeds) attract
Sinorhizobium
Daidzein, genistein (soybean roots)
attract Phytopthora sojae

Attachment/fixation on host
Secretion of adhesive materials
attract cells to plant
Mass bacterial cells affix themselves
by forming extracellular fibrils of
cellulose material

Effect of biological activity in


rhizosphere on plant
i. Recycling of mineral elements
Root exudate increase microbial
population; increase predators; increase
enzymes, decomposition and NH4+
release

ii. Decrease Redox Potential


Respiration of roots and microbes
consume O2; reduce redox potential;
increase solubility of certain cations
(Fe and Mn)
Increase toxic compounds

iii. Synthesis of growth factors


Biosynthesis of hormones; giberrelin, auxin,
cytokinin, ethylene
iv. Chelating agents
Siderophores that has high affinity for
Fe3+
organic acids( oxalic or ketogluconic acid)
involve in P solubilization
v. Production of toxic compounds
antibiotics; rhizobitoxin produce by certain
Bradyrhizobium cause chlorosis in soybean
vi. Detoxification of toxic compounds