Part 3: Physiological Adaptations to

Adverse Chemical Conditions

#Topics to be covered
– High input, low input approach to plant nutrition
– Nutrient Use efficiency
– Tolerance to excessive supply of mineral elements
– Acid mineral soil – physiological effects of excessive AL3+,
Mn4+ levels
– Al3+ tolerance and Mn4+ tolerance
– Water logging and flooded soils (i.e., soil chemical factors,
plant responses to water logging)
– Salinity
# High input, low input approach to plant nutrition

• Soil fertility problems can be solved in either of two ways:

A) High Input approach

 Here the objective is to adjust the soil to fit crop requirement i.e.,
adjusting soil fertility to factors such as pH and soil nutrient
availability to optimum levels for a given spp or variety.
 The High Input approach has been successful in temperate regions
but not for tropical or sub-tropical regions.
 This is due to the fact that in temperate regions soil chemical
conditions naturally change over a long period of time.
 Tropical and subtropical regions experience extremes in soil
acidity and nutrient deficiency which cant be easily remedied
 In the tropics and subtropics, soil alkalinity, soil acidity, Al3+ and
Mn4+ toxicity are widespread problems.
B) The Low Input Approach

 Aims to fit plants to the soil conditions rather than trying to supply
their optimum nutrient requirements.

 This has lead to the development of genotypes adapted to specific

ecological conditions.

 Breeding for high nutrient efficiency and tolerance to conditions

such as Al3+ and Mn4+ toxicity, water-logging and salinity is very
prevalent in the tropics.
#The concept of f nutrient use efficiency

 NUE is the ability of a crop to grow and produce dry matter and
economic yield under low fertility conditions.

 For a given genotype it is reflected by the ability to produce high

yield in a soil that is limiting in mineral nutrients as measured
with a standard genotype (grain).

 Experiments have been done for comparing yield or % yield

reduction in genotypes supplied with insufficient amounts of
mineral nutrients e.g. phosphorous use efficiency of pasture spp.
#Manganese Toxicity

 Effects on plants:
 Affects metabolic processes such as enzyme activities and organic compounds
 Sterility
 Symptoms similar to Fe excess
 Mn4+ toxicity often occurs with Al3+ toxicity

 Signs:
 Yellowish brown spots between leaf veins, extending to the whole interveinal area
 Brown spots on veins of lower leaf blades and leaf sheaths
 Drying of leaf tips eight weeks after planting
 Chlorosis of younger leaves
 Stunting
 Reduced tillering
 Reduced grain yield

***Mn toxicity can be determined by testing the plant and soil.

#Mn toxicity can be caused by one or more of the following:

 Large concentration of Mn2+ in the soil solution because of low soil

pH (<5.5)

 Poor and unbalanced crop nutrient status. Low root oxidation:

– deficiencies of Si, K, P, Ca, or Mg, and
– substances that inhibit respiration (e.g., H2S, FeS, organic acids).

 Application of urban or industrial waste with large Mn content.

#Soils where Mn toxicity can occur are as follows:

 Acidic soils (pH <5.5);

 Lowland soils containing large amounts of easily reducible Mn
 Acid sulfate soils
 Areas affected by Mn mining (e.g., Japan)

Some explanations:
 The Mn concentration in soil solution can increase at low soil pH or when redox
potential is low due to flooding.
 A large concentration of Mn in plant tissue changes metabolic processes (e.g., enzyme
activities and organic compounds).
 This then leads to visible Mn toxicity symptoms such as chlorosis (photo-oxidation of
chlorophyll) or necrosis (accumulation of oxidized phenolic compounds, e.g.,
anthocyanin).
 Crop varieties differ in their susceptibility to Mn toxicity, for instance, Mn toxicity
rarely occurs in rice.
 Excessive amounts of Mn in solution can lead to excess Mn uptake in cases where
exclusion or tolerance mechanisms in roots are not functioning adequately.
#The major adaptive mechanisms by which some plants
overcome Mn toxicity are as follows:

1. Mn stress avoidance: Release of O2 from roots (root oxidation

power) to oxidize Mn2+ in the rhizosphere. Differences in root
anatomy and morphology as well as toxic substances (H2S) affect
root oxidation power.

2. Mn stress tolerance: Retention of Mn in root tissue (oxidation and

accumulation of Mn2+ in cell walls). Concentration of excess Mn
in metabolically inactive forms.
#Measures to prevent Mn toxicity
1. Seed treatment:
 In a temperate climate, coat seeds with oxidants (e.g., Ca peroxide) to improve
germination and seedling emergence by increasing the supply of O2.

2. Water management:
 Source of water (should be clean).
 Avoid flooding.

3. Soil-Fertilizer management:
 Balance use of fertilizers (NPK or NPK + lime) to avoid nutrient stress as a source of Mn
toxicity.
 Apply sufficient K fertilizer.
 Apply lime on acid soils to reduce the concentration of active Mn.
 Do not apply excessive amounts of organic matter on soils containing large concentrations
of Mn.
 Use less-acidifying ammonium fertilizers (e.g., urea as N source)

4. Straw management:
 Recycle straw or ash to replenish Si and K removed from the field.
 An adequate Si supply prevents Mn toxicity of crop plants by decreasing plant Mn uptake
 Si also increases the internal tolerance for an excessive amount of Mn in plant tissue.
#Treating Mn toxicity:

a) Apply lime to alleviate soil acidity on upland soils.

b) Apply silica slabs (1.5 to 3 t ha-1) to alleviate Si deficiency

#Aluminum Toxicity

 Inhibits root growth & shoot growth by inducing nutrient (i.e., Mg,
Ca and P) deficiencies, drought stress, and phytohormone
imbalances.

 Signs
 Orange-yellow to white interveinal chlorosis on leaves.
 Poor growth or stunted growth.
 Yellow to white mottling of interveins is followed by leaf tip death and leaf margin
scorch.
 Necrosis of chlorotic areas during severe Al toxicity.
 Stunted and deformed roots in susceptible cultivars.

 Causes
 Excess Al3+ concentration in soil solution is caused by low soil pH (<5).
 The concentration of Al in soil solution depends on soil pH as well as the concentration
of organic and inorganic compounds that can form complexes with Al.
#Al toxicity occurs on the following soils:

 Acid upland soils with large exchangeable Al content.

 Al toxicity often occurs together with Mn toxicity.

 Acid sulfate soils (e.g., Thailand).

 Flooded soils with pH <4 before Fe toxicity symptoms appear.

#Genotypic differences in susceptibility to Al toxicity in rice
are as follows:

 Al stress avoidance, due to the exclusion of Al from sensitive

sites or reduced Al3+ activity in the rhizosphere, thus reducing
the Al inhibition of Ca2+ and Mg2+ influx.

 Al stress tolerance, due to high tissue tolerance of Al,

immobilization of Al in nontoxic forms
#Flooding/Waterlogging Tolerance

Getting enough oxygen to the roots is the key parameter that determines
whether the plant lives or dies.
#Explanation
 In waterlogged soils, diffusion of gases through soil pores is strongly inhibited
by water content that it fails to match the needs of growing roots.

 A slowing of oxygen influx is the principal cause of injury to roots, and the
shoots they support.

 Small amounts of oxygen dissolved in the floodwater in equilibrium with the air
is a little over 3%.

 This small amount is quickly consumed during the early stages of flooding by
aerobic micro-organisms and roots.

 In addition to imposing oxygen shortage, flooding also impedes the diffusive

escape and/or oxidative breakdown of gases such as ethylene or carbon dioxide
that are produced by roots and soil micro-organisms.

 This leads to accumulations that can influence root growth and function, for
e.g., accumulated ethylene may slow root extension, while carbon dioxide in the
soil can severely damage roots of certain species.
#How roots survive anaerobic conditions/adaptation
 During natural waterlogging of the soil, anoxia will be preceded
by a period of partial oxygen shortage (hypoxia).

 This will last for as long as it takes for dissolved oxygen in the
floodwater to be consumed by roots and other aerobic soil
organisms.

 This hypoxic interlude can act as a training period by improving

the ability of the roots to survive subsequent anoxia by inducing
biochemical acclimation or anatomical acclimation.
#Adaptations of Hydrophytes

1. Development of lots of air spaces in their tissues.

 Anatomical acclimatation through aerenchyma formation
 The presence of large interconnected intercellular gas-filled spaces that often extend
from the shoots to near the root tip (aerenchyma) is feature shared by most, although
not in all species, that grow well in waterlogged areas
2. Roots in water-clogged soil
are called ‘Breathing Roots’
or ‘Pneumatophores’.

• These roots grow to reach the

surface above the water in
order to obtain O2 to survive.

• Marshes or Swamps
3. Fast upward shoot elongation

 Another developmental effect of flooding, that supplements the aerenchyma

system, is the promotion of shoot extension and/or increased uprightness of
submerged leaves

 The resulting increase in effective plant height, that can begin after a delay of less
than 2 h of inundation, improves access to aerial or dissolved oxygen or to light for
the generation of photosynthetic oxygen.

4. Replacement rooting

 Dying roots can be replacement by roots positioned near or at the better-aerated soil
surface.

 There are three mechanisms for generating these replacement root systems.

1. Stimulation of the outgrowth of root primordia already present within the shoot base.

2. Induction of a new root system that involves initiation of root primordia and their
subsequent outgrowth e.g. in Helianthus.
5. Biochemical acclimatation

 The mechanism by which cells initially sense the partial oxygen

shortage that triggers this acclimation is unknown.

 Following sensing of partial oxygen deficiency, genes coding for

so-called anaerobic proteins (actually hypoxically-induced
proteins or HIPs) are up-regulated at transcriptional and post-
translational levels while others coding for many aerobic proteins
remain expressed and translated up to the point when the cells
finally become anoxic.
#Soil chemical factors of Water logging
 Trapped carbon dioxide may form bicarbonate ions that can lead to iron unavailability and
chlorosis.

 Warm temperatures and ample supplies of organic matter will accelerate the development
of these potentially damaging soil conditions.

 These changes come about because of microbial respiration that utilizes inorganic ions as
alternative electron acceptors to oxygen in order to sustain energy generation.

 The changes are associated with measurable decreases in redox potential.

 Facultative anaerobes first chemically reduce nitrate, converting it to nitrite, nitrous oxide
and nitrogen gas (denitrification), rendering nitrate unavailable to roots.

 As the reducing intensity of the soil increases further, obligate anaerobes chemically reduce
oxides of Mn4+ and Fe3+ to form highly soluble Mn2+ and Fe2 that may enter roots and
interfere with enzyme activities and damage membranes.

 If flooding is prolonged further, anaerobic bacteria may then convert SO42- to H2S, a poison
of respiratory enzymes and non-respiratory oxidases.

 Acidic soils that are low in iron are especially likely to contain free and undissociated H2S.
#Salinity
 Salinity occurs through natural or human-induced processes that result in the
accumulation of dissolved salts in the soil water to an extent that inhibits plant
growth.
 Sodicity is a secondary result of salinity in clay soils, where leaching through either
natural or human-induced processes has washed soluble salts into the subsoil, and
left sodium bound to the negative charges of the clay.
 A saline soil is defined as having a high concentration of soluble salts, high enough
to affect plant growth.
 Salt concentration in a soil is measured in terms of its electrical conductivity.
 Salt stress can be a major challenge to plants & it limits agriculture all over the
world, particularly on irrigated farmlands.
 Salinity resistance is a quantitative trait, and it can be breed.
 Many factors interact with salinity & this complicates studies on the effects of
salinity, for e.g., humidity, temperature, light, irrigation & soil fertility.
 Drought and salinity are connected because in many regions, raising plants requires
irrigation.
 Irrigation water contains Ca, Mg and Na and as irrigation water evaporates and
transpires, Ca & Mg transpire, leaving Na dominant in the soil.
 The uptake of ground water by plant roots can also increase the salinity of ground
water or the soil around the roots due to the exclusion of salt.
#Effect of Salinity on Plants
 Salts in the soil water may inhibit plant growth for two reasons.

 First, the presence of salt in the soil solution reduces the ability of the plant to take
up water, and this leads to reductions in the growth rate & this is referred to as the
osmotic or water-deficit effect of salinity.

 Second, if excessive amounts of salt enter the plant in the transpiration stream, there
will be injury to cells in the transpiring leaves and this may cause further reductions
in growth & this is called the salt-specific or ion-excess effect of salinity.

 Osmotic stress and ion toxicity are the problems stemming from salt stress and the
resulting decrease in chemical activity causes cells to lose turgor.

 Cell growth depends on turgor to stretch the cell walls and lack of turgor implies
danger for cell survival.

 The plant’s defense against this salinity attack requires osmotic adjustment, and, to a
certain degree, this can be done through synthesis of intracellular solutes.
#Mechanisms of salt tolerance
#Examples of salt tolerant plants
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