Alkaline soils

Lecture 14
Prof Sergey Shabala, 2017
Geographical distribution
Alkaline soils
Boron toxicity
Iron deficiency
(lime –induced chlorosis)
-Decrease in soluble
inorganic Fe
-Impairment of H+-Pumping
-Decrease in release of
phenolics
-Decrease in Fe(III)
reduction
-Indirect effects

Zinc deficiency
-Cereals; flooded rice
 Boron availability in the soil
• Soil concentrations > 12 mg B/kg
considered as toxic
• Present in the soil solution in
either B(OH)3 or B(OH)4- form
• Readily leached from the soil
• Strong pH dependence
 Boron as essential micronutrient
B Sugar transport; cell wall synthesis &
structure; lignification; pollen
germination; respiration; metabolism of
IAA, RNA and carbohydrates

• Critical
deficiency:
5-10 mg kg-
1 DW in
cereals
• Critical
deficiency:
20-70 mg in
dicotes
Boron as essential micronutrient
Boron toxicity
 Basis for boron toxicity
• Reduced glutathione and
tocopherol content
• Reducing root cell division
• Reduced cell wall expansion
• Decreased fruit number and
size
• ROS production
• DNA damage
• Disrupted carbohydrate
metabolism
• Alteration in lipid
composition
• Disruption to cytoskeleton

Susceptible Moderately susceptible Tolerant

All winter cereals, sorghum,

Boron Deficiency lucerne, canola, sunflower clover, maize
rice pea, bean

Wheat (cv. Binnawarra),

Wheat (cv. Halberd),
Boron Toxicity Barley (cv. Stirling) Barley (cv. Skiff), Sunflower
Sunflower (H. annus)
(H. tuberosis)
Boron transporters

• Passive (simple or
channel-mediated
diffusion)
• Active efflux
Boron uptake by roots
• NIP5; 1 =
acquaporin
• Noduline-26-
like intrinsic
protein
• BOR1 =
exporter

Tissue-specific
expression
Boron tolerance mechanisms

• Reduced B uptake
• Efficient B exclusion
• Vacuolar sequestration?
Genetic variability in B tolerance

• Bo1-derived cultivars in
wheat
• Another (unknown) allele at
chromosome 4AL
• Not been harnessed in any
breeding program so far
Fe solubility as a function of pH
Effect of HCO3- on Fe uptake
 Symptoms of Fe deficiency

Element with low mobility

Symptoms evident in young leaves

Fe Redox system (cytochromes; ferredoxin; FeSOD);

bridging element; metabolic control (aminoleulinic
acid)
 Adaptation to Fe deficiency
• Iron efficiency mechanisms
– Non specific:
• Formation of lateral roots
• Differentiation of transfer cells
• Preferential cation uptake
• Release of organic acids
• Increased microbial activity
– Specific
• High H+ excretion (strategy I plants)
• Release of phytosiderophores (strategy II plants)
Two strategies dealing with Fe deficiency
Strategy I plants: changes in rhizosphere pH
After growth of sunflower (strategy I plant) in nutrient solution,
roots were placed in agar containing the pH indicator bromocresol purple

Nitrate +Fe Ammonium +Fe Nitrate -Fe

Total root -0.8 +3.6 +5.6

Active root zone +3.6 +28.0
Net release of H+ (µmol H+ [g-1 root FW] h-1)

Römheld et al. (1984), Plant Physiol. 76, 603-606.

 Strategy II plants

• Release of phytosiderophores under Fe deficiency

• Phytosiderophores are non-proteinogenic amino acids:
derivatives of mugineic and avenic acids
 Fe deficiency responses

• FIT (FER-like Iron

Deficiency-Induced
Transcription Factor)
• The master regulator
• Activated in response
to low iron

• Interacts with other transcription factors (bHLHs and EIN3/EIL1)

• Induce iron acquisition genes (FRO2, IRT1, and FIT)
 Fe modulation of root growth

Upon destabilization of DELLA in epidermal cells, FIT becomes available to

activate iron acquisition genes and eventually iron uptake via IRT1. The local
increase in symplastic iron leads to upregulation of AUX1, an auxin importer, and
increased levels of auxin enhance lateral root growth.
Fe acquisition and translocation

• IRT (Iron Regulated Transporter)

• FRO2 (ferric reductase oxidase)
• PDR9 (phenolic transporter)
• OPT3 (Oligo Peptide Transporter)
• YSL (Yellow Stripe-Like
transporter)
 Essentiality of Zn
Zn Enzyme structure (carbonic anhydrase, alcohol • Over 1000
dehydrogenase, RNA polymerase); enzyme activity proteins
(isomerases, dehydrogenases); structural component associated with
of ribosomes; IAA synthesis; membrane integrity; zinc for
carbohydrate metabolism functionality
• Catalytic,
regulatory, and
structural roles
for proteins and
enzymes

• Zinc-finger transcription factors

• Zinc superoxide dismutase
Zn deficiency
• Sufficiency range is 15-50 µg
g-1 DW
• Stunted growth (“resetting”)
• Drastic decrease in a leaf size
(“little leaf”)
• Chlorosis and necrosis (ROS
damage)
• Observed in young leaves
Mechanisms of Zn uptake
ROOT UPTAKE: ZIP1
and ZIP2 transporters
from IRT family

XYLEM LOADING: HMA

primary active Zn pumps

VACUOLAR
SEQUESTRATION: MTPs
(Metal Tolerance Protein)
Zn transport into developing grain
• Mechanisms largely
unknown
• Essential for
biofortification
 Amelioration of alkaline soils
• Washing salts down the profile
• Adding Ca2+ ions

• Adding acids

• adding sulphur