Breeding for resistance to

abiotic stresses
Plant growth and development is the product of the interaction between the genotype (genetic potential) and the
environment in which the plant grows. Any stress in the environment will adversely impact growth and
development. Plants perform well in environments to which they are well adapted. Yield potential was
previously defined as the highest yield attainable by a genotype growing in an environment to which it is
adapted and in which there is no environmental stress (i.e., optimum growing conditions).

Types of abiotic environmental stresses

Plant growth and development depend on biochemical processes (e.g., photosynthesis, respiration) that in turn
depend on factors in the environment in order to proceed optimally. As previously indicated, when conditions in
the environment are less than optimum, the plant experiences stress, which adversely affects its growth and
development, and ultimately its productivity and economic value. The common stresses that plants may be
exposed to in agroecological systems include the following:

1 Drought. This is the environmental condition caused by lack of rainfall.

2 Heat. Heat stress occurs when temperatures are high enough to cause irreversible damage to plant function.

3 Cold. Cold stress manifests when plants are exposed to low temperatures that cause physiological disruptions that
may be irreversible.

4 Salinity. Stress from salinity occurs when the dissolved salts accumulate in the soil solution to an extent that plant
growth is inhibited.

5 Mineral toxicity. Mineral toxicity occurs when an element in the soil solution is present at a concentration such that
plants are physiologically impaired.

6 Oxidation. Oxygen-free radicals (or activated oxygen) are known to cause degenerative conditions in plant cells.

7 Water-logging. Excessive soil moisture as a result of prolonged rainfall can cause anoxic soil conditions, leading to
roots suffering from lack of oxygen.

8 Mineral deficiency. Inadequate amounts of essential soil minerals for plant growth cause crop injury.

A. Drought stress

Drought occurs both above ground (atmospheric drought) and below ground (soil drought). The duration of
drought is variable, sometimes lasting for a short time and without severe adverse physiological impact,
sometimes lasting throughout an entire growing season or even years, resulting in complete devastation of crops.

Breeding for drought resistance

Drought resistance is highly cropping region specific. Drought resistance in crops is a major breeding objective
in dryland farming systems. In more advanced agricultural production systems, the goal of combating drought is
to obtain economic plant production in spite of the stress
Plant traits affecting drought response

Researchers have identified numerous stress response genes. However, these genes are not necessarily stress
adaptive in terms of drought resistance. Nonetheless, both adaptive and non-adaptive genes are important in
plant performance under drought stress. A good example of this fact is the role of genes that condition early
flowering. Whereas these genes are expressed in any environment (i.e., not drought induced), early flowering
may play a role in plant response to, and performance under, drought stress. Major plant traits that play a role in
plant drought-resistance response include phenology, development and size, root, plant surface, non-senescence,
stem reserve utilization, photosynthetic systems, and water use efficiency.

Mechanisms of drought resistance

Plant species differ in the stages at which they are most susceptible to drought stress. Some species are most
prone to stress damage during the early vegetative stage, while others are most susceptible during the pre- or
postanthesis stage, with others in between. Four general mechanisms may be identified by which plants resist
drought:

1 Escape. Using early maturing cultivars may allow the crop to complete its life cycle (or at least the critical growth
stage) before the onset of drought later in the season. The plants use the optimal conditions at the beginning of the
season to develop vigor.

2 Avoidance. Some plants avoid drought stress by decreasing water loss, for example by having cuticular wax or by
having the capacity to extract soil moisture efficiently.

3 Tolerance. In species such as cereals in which grain filling is found to depend on both actual photosynthesis during
the stage, as well as dry matter distribution from carbohydrates in pre-anthesis, terminal drought significantly reduces
photosynthesis. This shifts the burden of grain filling to stored carbohydrate as the source of dry matter for the
purpose. Consequently, such species may be more tolerant of postanthesis drought, being able to produce appreciable
yield under the stress.

4 Recovery. Because drought varies in duration, some species are able to rebound (recover) after a brief drought
episode. Traits that enhance recovery from drought include vegetative vigor, tillering, and long growth duration.

Breeding methods

In breeding for drought resistance, breeders may select for early maturity for use in avoiding the stress. Also,
genotypes with proven drought resistance may be used in hybridization programs to transfer the resistance genes
into superior cultivars.

B. Cold Stress

Plants use a variety of adaptive mechanisms to combat environmental stress caused by cold temperature. Seed
dormancy is a physiological condition that delays seed germination until the embryo has gone through an after
ripening period, during which certain biochemical and enzymatic processes occur for the seed to attain full
maturity.
When plants are exposed to gradually decreasing temperatures below a certain threshold, they acclimatize (low
temperature acclimation) to the stress, a process called cold hardening. In spite of various adaptations to cold,
plants may be injured through exposure to cold temperatures in a variety of ways, depending on the temperature
range. One type of injury, called chilling injury, occurs at exposure to temperatures between 20 and 0°C.

Mechanisms of resistance to low temperature

Like drought resistance, certain physiological or morphological adaptations can make plants either avoid or
tolerate stress due to low temperatures. The mechanism of low-temperature resistance may be
grouped into two.
1. Chilling resistance

The factors that confer resistance to chilling are believed to operate at the cell membrane level where they
influence membrane fluidity. Chilling-resistant seeds are known to imbibe moisture slowly. The presence of
phenols in the seed coat of legumes is implicated in conferring chilling resistance.

2. Freezing resistance

Several mechanisms are used by plants to resist freezing, including the following:

1 Escape. Like drought, cultural practices may be adapted by producers to prevent the vulnerable stage of growth
coinciding with the presence of the stress factor.

2 Avoidance. One of the injuries of low temperature results from the intracellular formation of ice following
nucleation of ice in the tissue. Water may remain supercooled without forming ice crystals. Certain compounds that
are capable of promoting ice nucleation are active at low temperatures. Bacteria such as Pseudomonas syringae are
capable of producing ice-nucleating proteins. The first field test of a bioengineered organism was the testing of “ice
minus”, a microbe genetically engineered to be incapable of producing the bacterial protein that causes ice nucleation.
This was intended to be an approach for helping frost-sensitive plants survive frost.

3 Tolerance. Freezing tolerance occurs when a plant is able to withstand both intracellular and extracellular ice
formation.

Breeding for tolerance to low-temperature stress

Whereas the genetics of low-temperature tolerance have been studied to a reasonable degree, breeders have only
had minimal success in applying research results to practical breeding. Breeding superhardy cultivars remains a
challenge.

1 Exploitable genetic variability for low-temperature tolerance has been largely exhausted within the existing gene
pools of most species.

2 A large number of genes with small effects and complex interaction are assumed to determine the phenotypic
expansion of low-temperature tolerance, making selection difficult.

3 Current methodologies for measuring low-temperature tolerance give poor resolution of small phenotypic
differences.

4 Measures of low-temperature tolerance lack the precision for single-plant analysis and many are destructive, making
selection procedures complicated.

5 Poor expression of low-temperature tolerance in alien genetic backgrounds has prevented the expansion of gene
pools through interspecific and intergeneric transfers (e.g., the superior low-temperature tolerance of rye is suppressed
in the wheat background).

C. Salinity stress

Soil salinity constraints to crop production occur in an estimated 95% million hectares worldwide. Salinity is
the accumulation of dissolved salts in the soil solution to a degree that inhibits plant growth and development.
Plant growth is inhibited in salt-affected soils because the high salt concentration in the soil solution inhibits the
process of water absorption by osmosis (osmotic stress). When excessive amounts of salt enter the transpiration
stream, plant cells may be injured. Plants that are tolerant of high soil salt concentration are called halophytes.

Breeding for salt tolerance

A common approach to breeding salt tolerance starts with assembling and screening germplasm for salinity
tolerance. The selected genotypes are used as parents to transfer the trait to desired cultivars, followed by
selecting desirable recombinants from the segregating population. This approach has yielded some success in
species such as rice, wheat, and lucerne. The challenge in breeding for salt tolerance is how to measure salinity
tolerance. Screening is commonly based on the growth of plants under salt stress. Two distinct mechanisms
exist for salinity tolerance:
1 Tolerance to the osmotic effect of the saline solution (the osmotic effect makes it harder for plants to extract water
from the soil).

2 Tolerance to the salt-specific nature of the soil saline solution (a high sodium concentration makes it difficult for the
plant to exclude NaCl while taking up other ions).

Selection for these ions was used in breeding rice and lucerne cultivars with high salt tolerance. Traits for
osmotic effects are related to growth (e.g., leaf elongation, root elongation, shoot biomass, leaf area expansion),
the latter two being the most effective indices. Molecular maker technology and genetic engineering techniques
are being used in salt-tolerance breeding efforts. Salinity tolerance has been found in the wild species of crops
such as tomato, pigeon pea, and common bean.

D. Heat stress

Heat stress may be defined as the occurrence of temperatures hot enough for a sufficient time to cause
irreversible damage to plant function or development. A heat-resistant genotype is one that is more productive
than another genotype in environments where heat stress occurs. Heat tolerance is the relative performance of a
plant or plant process under heat compared with the performance under optimal temperatures.
Excessive heat in the soil affects the emergence of seedlings of both cool and warm season crops causing
reduced crop stands. High temperatures tend to accelerate reproductive development. This may be part of the
reason why the potential grain yields of warm season crops (e.g., rice, cowpea)

Breeding for resistance to heat stress

Breeding for resistance to heat stress has not been as widely addressed as other environmental stresses that
plants face in crop production.
An approach to breeding heat resistance that is deemed by some to be more efficient is to select for specific
traits that confer heat tolerance during reproductive development. To do this, genotypes with heat tolerance have
to be discovered. This involves screening large accessions from germplasm collections. These genotypes can
then be crossed with desirable cultivars if they lack the yield and other plant attributes desired.

E. Mineral toxicity stress

Plants obtain most of their nutrient requirements from the soil, largely from the products of weathering of
mineral rocks or the decomposition of organic matter. Uptake in improper amounts may lead to toxic
consequences to plants.

Soil nutrient elements

Metals occur naturally in soils, some of which are beneficial and essential for plant growth and development,
while others are toxic. However, at extreme conditions of soil reaction, excessive amounts of some elements
become available. Some micronutrients are required in only trace amounts; their presence in large quantities in
the soil solution may be toxic to plants. Some of the known toxicities of metallic elements occur at low pH (high
acidity) and include iron and aluminum toxicities.

Aluminum toxicity

Aluminum (Al) is one of the most abundant elements in the earth’s crust. One of the most important metal
toxicities of economic importance to crop production is that which occurs when aluminum concentrations are
greater than 2–3 ppm. At acid pH, Al3ions predominate in the soil. Aluminum is not an essential nutrient for
plants. At a pH of 5 or less, aluminum inhibits plant growth by interfering with cell division in root tips and
lateral roots, increasing cell wall rigidity, reducing DNA replication, decreasing respiration, and other effects. In
some cases, excess aluminum induces iron deficiency in some crops (e.g., rice, sorghum, wheat).

Breeding for aluminum tolerance

Aluminum-tolerant genotypes have been identified. Based on the patterns of aluminum accumulation in plant
tissue, three groups of aluminum-tolerant plants may be identified: (i) those with an apparent exclusion
mechanism allowing lower accumulation of aluminum in their roots than aluminum-sensitive plants (e.g., wheat,
barley, soybean); (ii) those with less aluminum in the shoot but more in the roots (e.g., wheat, barley, potato);
and (iii) those with high aluminum accumulation in the shoot (e.g., pine trees). Research in wheat suggests the
possibility of more than one aluminumtolerance gene and more than one aluminum-tolerance mechanism.

F. Mineral deficiency stress

Concepts associated with mineral deficiency

Mineral deficiencies or toxicities are widespread. A report from the Centro Internacional de Agricultura
Tropical (CIAT) in Peru estimates that about 60% of the soils in the common bean production regions of the
world have some soil mineral problem. Soils that are high in calcareous minerals tend to have high amounts of
basic elements (e.g., Ca, Mg, K) that tend to raise soil pH. A high soil pH in turn causes mineral deficiency
problems (e.g., Fe, Zn, P). Common mineral deficiency symptoms are summarized in Table 21.4. Zinc
deficiency in common bean has been reported in production areas such as southern Idaho and Michigan.

Breeding efforts

Cultivars vary in their sensitivity to zinc deficiency. Sensitive cultivars take up and store less zinc in various
plant parts and the seed than resistant cultivars. Researchers in common bean identified a zinc deficiency
resistant cultivar, “Matterhorn”, and subsequently determined that a single dominant gene, Znd, conditioned
resistance to soil zinc deficiency.

G. Oxidative stress

Concepts associated with oxidative stress

Oxygen free radicals (or activated oxygen) have been implicated in a variety of environmental stresses in plants.
They are involved in many degenerative conditions in eukaryotic cells (e.g., peroxidation of lipids, cross-linking
and inactivation of protein, and mutation in DNA).
Numerous sites of oxygen activation occur in the plant cell. These sites are highly controlled and coupled to
prevent the release of intermediate products. It is presumed that such a control or coupling breaks down when a
plant is under stress, resulting in leaking of activated oxygen. Injuries to the plant occur when the production of
activated oxygen exceeds the plant’s capacity to detoxify it. Symptoms of oxidative stress include loss of
osmotic responsiveness, wilting, and necrosis.

Applications and breeding efforts

Several herbicides are designed to function by the involvement of activated oxygen. These herbicides promote
the accumulation of metabolic intermediates and the energy used to create singlet oxygen, which kills the plant.
These herbicides are described as photobleaching (e.g., p-nitrodiphenyl ethers). Other herbicides that depend on
light and chlorophyll are paraquat and diquat (both bipyridylium herbicides). So far, few plants have been
selected for tolerance to oxygen free radicals.

H. Flood stress (water logging)

Whereas some plants are adapted to water-logged conditions (e.g., flooded rice culture), most plants need well
drained soils to grow properly.

Concepts associated with water-logging stress

In soybean, stress due to water logging can reduce crop yield by 17–43% when it occurs at the vegetative stage,
and by about 50–56% if the stress occurs at the reproductive stage. Floods are often caused by excessive rainfall
due to a prolonged seasonal rainfall. The excessive amount of water quickly creates anoxic (oxygendeficient)
soil conditions causing flood-sensitive plants to suffer anoxia or hypoxia. Fermentation occurs in plant roots
under such conditions. The photosynthetic capacity of plants is significantly inhibited. Floodtolerant species
have certain adaptive mechanisms, such as the formation of aerenchyma and adventitious roots. Some studies
indicate that root tissue survival under hypoxia depends on the fermentation rate and sufficient sugar supply to
maintain cell energy and membrane function.
Breeding efforts

Tolerance to water-logging appears to be quantitatively inherited. QTLs for tolerance to water-logging have
been reported in rice and soybean.