PP-504

Sarita Maam:

Plant Adaptive Mechanisms to Drought

Drought stress is one of the most severe abiotic stresses affecting global agriculture. It occurs
when water availability is reduced below the optimal level required for plant growth,
development, and reproduction. Plants, being sessile organisms (unable to move), have
developed multiple survival strategies to endure drought, which include escape, avoidance,
and tolerance mechanisms.
 (A) Drought Escape and Desiccation Avoidance
Mechanisms
These are proactive strategies plants use to avoid experiencing stress or reduce its damaging
effects when it occurs.

1. Concept of Stress Escape

Definition:

Drought escape refers to the ability of a plant to complete its life cycle before the onset of
drought. It is a temporal strategy that allows the plant to "escape" drought periods by
growing during favorable times.

How does it work?

Instead of developing resistance or tolerance, plants:

 Germinate rapidly when water is available

 Grow, reproduce, and complete their life cycle before the soil moisture drops
 Avoid the need to survive during drought altogether

Genetically Controlled:

This behavior is regulated by genes controlling flowering time, maturity period, and
vegetative growth rate.

Examples:

 Desert ephemerals: Plants in arid zones that grow quickly after rain and complete their
life cycle within a few weeks.
 Early-maturing crop varieties like:
o Wheat (e.g., HD 2967)
o Sorghum and Pearl millet for drylands

2. Exploiting Genetic Variability in Phenology

Phenology refers to the timing of key developmental stages of a plant: germination, flowering,
fruiting, and maturity.

Importance in Drought Escape:

  Early flowering and maturing genotypes are less exposed to terminal drought stress
(which usually occurs during flowering or grain filling).
 Exploiting this variability helps in breeding and selecting crop varieties suited to
specific drought-prone environments.

Examples of Phenological Variability:

 Sorghum: Varieties differ in flowering time, so farmers choose types depending on

rainfall patterns.
 Rice: Short-duration varieties like Sahbhagi Dhan are used in rainfed ecosystems.

(B) Drought Avoidance Mechanisms

Drought avoidance mechanisms enable the plant to maintain internal water balance (cell
turgor) despite external water scarcity. It allows continued physiological functions even under
limited water.

1. Maintenance of Cell Turgor

Cell turgor is the pressure exerted by water inside the cell against the cell wall. It's vital for:

 Cell expansion
 Stomatal opening
 Nutrient uptake
 Photosynthesis

✅Mechanisms for Turgor Maintenance:

i. Osmotic Adjustment:

 Cells accumulate small organic compounds (called osmolytes) like:

o Proline
o Glycine betaine
o Soluble sugars (trehalose, fructans)
 These compounds reduce the osmotic potential inside the cell, allowing water uptake
even from dry soil.
 This process does not interfere with cellular metabolism.

ii. Increased Cell Wall Elasticity:

 Elastic cell walls stretch more easily, enabling cells to hold water even when turgor
pressure is low.

iii. Reduction in Leaf Area:

  Smaller or fewer leaves reduce the surface area for transpiration, conserving moisture.

Crop Examples:

 Barley and wheat accumulate proline under stress.

 Chickpea shows excellent osmotic adjustment during flowering under water-deficit.

2. Water Mining by Root Characteristics

A key to drought avoidance is the ability of roots to extract water from deeper or larger soil
volumes.

Root Traits Involved:

i. Deep Root Systems:

 Roots grow vertically downwards to access deeper moisture reserves.

ii. Extensive Lateral Roots:

 Greater spread allows roots to explore more surface area for residual water.

iii. Root Density & Root Hair Proliferation:

 Denser roots and more root hairs enhance absorption efficiency.

iv. Faster Root Elongation:

 Helps reach moist zones more rapidly during early drought stages.

Example:

 Pearl millet, pigeon pea, and groundnut have root systems adapted to drought-prone
soils.

Moisture Conservation Mechanisms

These are mainly morphological or anatomical adaptations that help in reducing water loss.
 1. Regulation of Transpiration

Transpiration is the loss of water vapor through stomata. Under drought, plants try to limit this
to conserve water.

✅Traits That Help Reduce Heat Load:

i. Leaf Orientation:

 Leaves align vertically to reduce direct sunlight interception and overheating.

ii. Leaf Reflectance:

 Waxy or hairy leaves reflect solar radiation and minimize leaf temperature.

iii. Leaf Rolling or Folding:

 In crops like rice, leaves roll up during water stress to reduce the exposed area and trap
humidity.

Heat Avoidance vs. Heat Tolerance:

Reducing leaf temperature helps in maintaining enzyme activity, membrane stability, and
photosynthetic efficiency during heat-drought combinations.

2. Stomatal Factors and Guard Cell Metabolism

Stomata play a crucial role in gas exchange and transpiration. Their regulation is vital under
drought.

➤ Stomatal Closure:

 Mediated by the hormone Abscisic Acid (ABA).

 Under water deficit, ABA accumulates and signals guard cells to close stomata, reducing
transpiration.

➤ Guard Cell Osmoregulation:

 Ion channels in guard cells regulate the movement of K⁺ and Cl⁻ ions, controlling water
inflow and stomatal aperture.
 Efficient ABA response in tolerant plants results in fast stomatal closure with minimal
loss of photosynthesis.
➤ Genetic Basis:

 Genes like OST1, SLAC1 are involved in ABA-mediated stomatal closure.

3. Moisture Conservation by Waxes (Cuticular Wax Layer)

The cuticle is a hydrophobic layer made of cutin and wax covering the epidermis of aerial parts.

✅Benefits of Thick Waxy Coating:

 Reduces non-stomatal transpiration

 Increases leaf reflectance (light-colored waxes reflect solar radiation)
 Acts as a barrier against pathogens and UV damage
 Helps in maintaining leaf surface integrity during drought

Examples:

 Sorghum: Known for thick epicuticular wax under stress

 Cowpea: Exhibits glossy leaves due to wax, aiding in drought tolerance

Overall Summary Table

Mechanism Type Trait/Process Function in Drought
Avoids drought by completing life cycle
Escape Early maturity, short lifecycle
early
Maintains water status and
Avoidance Deep roots, osmotic adjustment
physiological activity
Moisture Leaf wax, leaf rolling, stomatal Minimizes water loss from plant
Conservation closure surfaces
Stomatal Control ABA-induced stomatal closure Limits transpiration effectively
Anatomical Changes Hairy or waxy leaves Reduces heat load and water loss

Modern Implications and Use in Breeding

  Marker-assisted breeding and genomics are now used to identify and select for
drought-tolerant genes.
 Transgenic approaches: Overexpression of DREB genes (Drought-Responsive Element
Binding) leads to enhanced tolerance.
 CRISPR/Cas9 gene editing is being explored to modify key genes like NCED3 (in ABA
biosynthesis) for precise drought resistance.

Awadhesh Sir:

Stress Perception and Molecular Responses

of Plants to Drought Stress
Drought stress affects plants at various levels—physiological, cellular, and molecular. To cope
with it, plants have developed a multi-layered response system, beginning with perception of
stress signals, followed by signal transduction, gene expression, and activation of stress-
specific proteins and adaptive mechanisms.

This topic can be broken down into four major areas:

1. Stress Perception in Plants

What is Stress Perception?

Stress perception refers to a plant's initial recognition of an unfavorable change in the

environment—such as a drop in soil moisture or cellular dehydration.

How do plants perceive drought?

Plants lack a nervous system, but they use specialized membrane proteins and sensors located
in:

 Plasma membranes
 Chloroplasts
 Nuclei
 Endoplasmic reticulum

Key Components in Drought Perception:

 1. Receptor-like Kinases (RLKs):
o These are membrane-bound sensors that detect changes in turgor pressure,
osmotic potential, and cell wall structure.
o They act as the first messengers in drought signaling.
2. Histidine Kinases (HKs):
o Function as osmosensors (e.g., in the Arabidopsis AHK1 gene).
3. Mechanosensitive Channels:
o Sense cellular deformation or stretching due to water loss.
4. Reactive Oxygen Species (ROS):
o Drought causes imbalance in photosynthesis and respiration → leading to ROS
production (H₂O₂, O₂⁻).
o These ROS serve as secondary messengers for signal initiation.

2. Signal Transduction Pathways

Once stress is perceived, it must be communicated to the cell's nucleus for appropriate gene
expression. This stress signaling involves several steps and molecules.

Key Elements in Drought Signal Transduction:

(a) Second Messengers:

These molecules quickly transmit the signal inside the cell.

Messenger Role
Major second messenger; binds with proteins like calmodulin to
Ca²⁺ ions
activate transcription
Inositol Phosphates
Mobilizes calcium stores
(IP₃/IP₆)
Reactive Oxygen Species
Activates downstream kinases and defense proteins
(ROS)
Abscisic Acid (ABA) Most critical hormone for drought signaling

(b) Protein Kinase Cascades

What are Protein Kinases?

Enzymes that phosphorylate target proteins, altering their function or activity.

Key Kinases in Drought Response:

 1. Mitogen-Activated Protein Kinases (MAPKs):
o A cascade of MAPKKK → MAPKK → MAPK
o Amplifies signal and transmits it to transcription factors
o E.g., MPK3, MPK6 in Arabidopsis
2. Calcium-Dependent Protein Kinases (CDPKs):
o Activated by elevated intracellular Ca²⁺
o Regulate stress-specific gene expression
3. SnRK2s (Sucrose Non-Fermenting 1-related protein kinases):
o Key regulators of ABA-dependent signaling
o SnRK2.2/2.3/2.6 are activated by ABA and help in stomatal closure

(c) Role of ABA in Signal Transduction

 ABA is synthesized in roots and leaves during drought.

 Binds to PYR/PYL/RCAR receptors, which in turn:
o Inhibit PP2C phosphatases (negative regulators)
o Activate SnRK2s, leading to gene expression and stomatal closure

3. Expression of Stress-Specific Regulatory Genes

After signal transduction, the next step is to turn on genes that help the plant adapt to drought.
These genes are categorized as:

(a) Regulatory Genes

These are “master switches” that control the expression of multiple downstream genes.

Key Regulators:

1. Transcription Factors (TFs):

o Bind to specific DNA sequences in gene promoters
o Activate or repress gene transcription

4. Drought-Specific Transcription Factors (TFs)

These proteins regulate stress-responsive genes. Each TF family recognizes specific DNA
elements like DRE, ABRE, or MYB/MYC binding sites.
 TF Family Description Examples Function
DREB (Dehydration Responsive Binds to DRE/CRT DREB1A, ABA-independent
Element Binding) elements DREB2A pathway
AREB/ABF (ABA-Responsive Binds to ABRE ABA-dependent
ABF1, ABF2
Element Binding) elements regulation
General stress AtMYB60, Root and stomatal
MYB and MYC
regulators AtMYC2 regulation
Involved in cell Improves tolerance
NAC OsNAC6
division and stress in rice
Pathogen and drought WRKY33, Multi-stress
WRKY
responses WRKY75 protection

5. Functional Genes Associated with Adaptive

Mechanisms
Functional genes encode products that directly help plants survive drought. These are not
regulators but act at the physiological or biochemical level.

Types of Functional Genes:

1. Osmoprotectant Biosynthesis Genes:

 Proline: P5CS (Δ¹-pyrroline-5-carboxylate synthetase)

 Glycine betaine: BADH (Betaine Aldehyde Dehydrogenase)

2. LEA (Late Embryogenesis Abundant) Proteins:

 Stabilize proteins and membranes during dehydration

 Highly hydrophilic and accumulate under stress

3. Dehydrins:

 Type of LEA proteins

 Prevent protein denaturation and enzyme inactivation

4. Aquaporins (Water Channel Proteins):

 Regulate water movement across membranes

 Adjust cell water potential rapidly

5. ROS-Scavenging Enzymes:

 Superoxide dismutase (SOD), Catalase (CAT), Ascorbate peroxidase (APX)

  Prevent oxidative damage

6. Chaperone Proteins (e.g., HSPs):

 Maintain proper protein folding under heat and drought

Summary Table
Category Examples Function
Stress Sensors RLKs, HKs Detect water deficit
Signal Molecules Ca²⁺, ROS, ABA Transmit stress signal
Kinases MAPKs, SnRK2 Activate downstream targets
Transcription Factors DREB, NAC, MYB, ABF Regulate stress gene expression
Functional Genes P5CS, LEA, HSPs, SOD Provide tolerance mechanisms

Applications in Agriculture
How is this useful in crop improvement?

1. Biotechnological Interventions:
o Overexpression of TFs like DREB1A enhances drought tolerance.
o CRISPR-based gene editing targets ABA receptors or TFs.
2. Marker-Assisted Selection:
o Genes like DREB2, NCED3, AREB1 serve as markers for drought resistance.
3. Synthetic Biology:
o Engineering synthetic circuits for timely expression of protective genes.

Conclusion
The molecular response of plants to drought is a complex yet well-orchestrated process,
starting from perception to gene expression. Understanding this helps scientists design drought-
resilient crops through genetic engineering, molecular breeding, and genome editing.