Drought Tolerance of Crops: Progress and Challenges

Dr. Viswanathan Chinnusamy

Principal Scientist
Division of Plant Physiology
IARI, New Delhi 110012
Email: viswanathan@iari.res.in

OUTLINE

Introduction to drought stress

Target Environment
Mechanisms of drought tolerance
Physiological and molecular genetic
basis of drought tolerance
Yield stability under drought
Challenges:
o MIADE
o Realistic, accurate and high
throughput Phenotyping
o Molecular genetics of crop plants

ABIOTIC STRESSES

Excess of non-essential
environmental factors

Water - deficit, hypoxia, anoxia

Temperature- Low, High
Nutrients deficiency, toxicity
Light - low, high
pH acidity, alkalinity

Ionic toxicity - salinity,

alkalinity, heavy metals
Air pollution
UV-B radiation

Growth

Growth

Scarcity or excess of essential

environmental factors

Quantity of factor

Quantity of factor

Crop Yield

Drought

Drought = moisture deficit stress

water stress
PWP

FC

Water
logging

Soil moisture
Meteorological drought: Deficit in precipitation over a long term average

Agricultural drought: Soil moisture deficit that leads to reduction in growth,

development and yield of crops

Target environment
Terminal

Intermittent

Water availability

FC
Water scarcity

PWP

Yield

Crop growth stage

25

50
Water Availability

100

Drought Stress Tolerance is No More a Myth

It is very common to state that drought tolerance is a very complex
trait and it is not tractable for genetic improvement.

It is often believed that drought tolerance is a nebulous term that

becomes more nebulous the more closely we look at it, much as
a newspaper photograph does when viewed through a
magnifying glass (Passioura 1996).
This opinion emerged rather due to a global view of yield response of
plants under illdefined experimental conditions or agricultural
situations. Hence the results are confusing and non-reproducible.
Omics of plant stress response revealed massive change in
epigenome, transcriptome, proteome and metabolome in response to
drought that further strengthened the opinion that drought tolerance is
a very complex trait.

Complex traits are amenable for easy modification

GA regulated genes are more than 1100 in Arabidopsis seedlings.

Yield improvement in green revolution is brought about by single gene mutation that
resulted in gibberellin (GA) deficiency in rice (semi-dwarf1/sd1) and GA insensitivity
in wheat (Reduced height/Rht).

"Clearly, recently we have seen some very promising advances in terms of

drought tolerances in crop plants," says Nguyen. "Now it's a question of how
to optimize the system."

Tester is also optimistic: Ultimately, "I think there will be a palette of genes from
which breeders and crop scientists will select for putting together the
drought tolerance for a particular region. (Pennisi E. 2008)

Drought Tolerance
Drought Resistance

The mechanisms causing minimum loss of yield in a water

deficit environment relative to the maximum yield in a water
constraint free management of the crop.
Drought Tolerance

Constitutive
mechanisms

Drought Tolerance

Acquired
mechanisms

Dehydration
Avoidance

Dehydration
tolerance

Yield Stability

where:

GY = WU x WUE x HI
WU= water transpired by the crop
WUE = water use efficiency (=biomass/unit water transpired)
HI = harvest index (economic yield/total biomass)
Passioura (1977)

Drought Tolerance
Dehydration Avoidance
Water Mining (WU)

Dehydration tolerance

Minimizing
water loss

Root traitsTranspiration -Stomata

Morphological &
LAI (Leaf area, no.
anatomical changes Leaf rolling & drying)
Aquaporins
leaf reflectance
OA
characters (wax load
and pubescence, leaf
angle, leaf rolling)

+ ABA
-CK, Ethylene

Cellular tolerance

WUE &
EUW

Phenotypic &
developmental
plasticity

Metabolic homeostasis
Xanthopyll cycle
Photorespiration
Maintenance respiration
Osmoprotection
Cell membrane stability
Oxidative Stress Mgmt.
Stress proteins

Activity &
Efficiency of
rubisco at low Ci

Phenology
Flowering
Development

Recovery, growth, yield

+ ABA
+CK, Auxin, GA,

Traits & Genes:

Water Mining

QTLs for Root System Architecture

Root-ABA1, a major constitutive
QTL, affects maize root architecture
and leaf ABA concentration

Root-yield-1.06, a major constitutive QTL

for root and agronomic traits in maize
across water regimes

Giuliani et al. 2005. J Exp Bot. 56: 30613070

Landi et al. 2010. J Exp Bot. 61: 35533562

PWRKY6::CKXox

Root-specific manipulation of single gene expression can increase root traits

Plant Physiology, May 2010, Vol. 153, pp. 185197,

CL, Culm length

PL, panicle length
NP, number of panicles per hill
NSP, number of spikelets per panicle
TNS, total number of spikelets
FR, filling rate
NFG, number of filled grains
TGW, total grain weight
1,000GW, 1,000 grain weight.
NT

RCc:OSNAC10

Application of omics technologies has contributed to the development

of stress-tolerant crops in the field
cDNA microarray of drought stress response in rice SNAC1 with 5.6 fold
enhanced expression was isolated from the upland rice cultivar IRAT109.

WT

35S::SNAC1

Hu et al. PNAS USA 103: 1298712992

The hrd-D mutant showed more

secondary roots in Arabidopsis

Genes for root traits

+ ABA

+ Auxin

- CK

- Ethylene

Traits & Genes: Osmotic Adjustment

High OA

Low OA

QTLs for osmotic adjustment in rice

Garg et al. 2002. 99:15898-15903

Traits & Genes: WUE

Crop

No. of QTLs

References

Wheat

10 QTLs affecting per plant WUE (Total dry

matter/ amount of water used by per plant)

Zhang et al., 2002

Rice

7 QTLs located in 5 chomosomal regions.

Xu et al., 2009

ERECTA, a putative leucine -rich repeat receptor-like kinase (LRR-RLK), known for its
effects on inflorescence development, is a major contributor to a 13C QTL on chr2.

ABA pathway engineering:

Minimization of water loss &
Enhancement of cellular tolerance

Traits & Genes: Transpiration control & Cellular tolerance

aba2

WT

ABA: Water mining Regulate primary root growth; Minimization of transpiration

Stomatal Control; Osmotic adjustment regulates gene expression; Cellular tolerance

vp14

Isomerase/A
BA4

WT
WT

NCED3-ox

ABA2

BG1

WT

aao3

90% ~30% RH,

10 min

/aba3

ABA Signal Transduction

Negative regulators and effectors are shown in red for clarity. Colors in boxes
represent relative expression level of a gene before and after ABA treatment
(+ABA).
Plant Cell 16:596-615 (2004)

Drought tolerance of rd29A:anti-AtFTB canola during flowering

The DH12075 (a) and YPT2-RD29AantiAtFTP (b) plants were subjected to a
4-day drought treatment starting on day 8
after flowering. The pictures were taken
on day 8 of re-watering after the drought
stress

Seed yields of WT and transgenic canola in 2003 (a) and 2004 (b) confined field trials
In (a) the solid bars represent
seed yields for the twoirrigation condition, and the
dotted bars represent seed
yields for the one irrigation
condition.
Irrigation
was
conducted
during
the
flowering period.
Wang et al. 2005. Plant J. 43: 413424

Oh etconfers
at. 2005. Arabidopsis
CBF3/DREB1A
ABF3moss
in transgenic
rice plants
ABA
dessication
tolerance and
from
to higher
increased tolerance to abiotic stress without stunting growth. Plant
Physiology 138:341-351

Red Arrow indicates re-watering

Science Breakthrough 2009

Identification of the ABA receptors and its mechanism of action
2009 Sciences TOP 10

These results are a boon for plant biologyand possibly beyond. The PP2C and the
ABA receptors both belong to highly conserved families of proteins whose roles in other
organisms may become clearer now that their function in plants has been nailed down

Traits & Genes: Transpiration control & Cellular tolerance

P
ABF3

Hubbard K E et al. (2010) Genes Dev. 24:1695-1708

Overexpression of ABA Receptor enhances drought tolerance

Vaccination ?
Identification of new agonist chemicals
Engineered receptors
activated
by
the
agrochemicals

Wheat seedlings grown in vermiculite

and severely desiccated under drought.
The seedling on the right received 0.1
mol of ABA in the irrigation water
before the onset of stress. Control
seedlings received normal irrigation
water before on the left.

that can be
off-the-shelf

Yield stability under field drought stress conditions

Reproductive stage stress tolerance

Spikelet Fertility

Molecular basis of spikelet fertility under drought stress needs to be studied

Regulated Expression of an Isopentenyltransferase Gene (PSARK::IPT) in

Peanut Significantly Improves Yield Under Field Drought Conditions
Expression
of
IPT
in
senescence
associated
promoter in leaves increases
drought tolerance

NT

PSARK::IPT

Qin et al. 2011. Plant Cell Physiol 52: 1904-1914

Plant nuclear factor Y (NF-Y) B subunits improved corn yields under drought

Nelson et al. 2007. PNAS USA 104: 16450-16455

Bacterial RNA Chaperones Confer Improved Grain Yield in Maize under WaterLimited Conditions

Castiglioni et al. 2008. Plant Physiology 147: 446455

QTL for Yield under drought

Bernier J, Kumar A, Ramaiah V, Spaner D, Atlin G (2007). A large-effect QTL for grain
yield under reproductive-stage drought stress in upland rice. Crop Sci. 47:505-516
Vandana x Way Rarem qtl12.1
This QTL accounted for an increase in grain yield, harvest index, and biomass yield under
stress, and it was detected over seasons under field conditions. The QTL accounted for
51% of the genetic variance in yield under drought
QTL qtl12.1 increases water uptake in upland rice:
Bernier et al. 2009. Field Crops Research 110: 139-146
qtl12.1 has a large and consistent effect on grain yield under upland drought stress
conditions, in a wide range of environments (21 field trials)
Bernier et al. 2009. Euphytica 166:207217

Kumar R, Venuprasad R, Atlin GN (2007). Genetic analysis of rainfed lowland rice

drought tolerance under naturally-occurring stress in eastern India: heritability and
QTL effects. Field Crops Res.103:42-52.
A QTL on chromosome 1 that accounts for 32% of the variation in yield under drought
stress in rainfed lowland rice.

Trends in Plant Science,

June 2011, Vol. 16, No. 6

Yang et al. 2010. Mol Plant 3: 469490

Yang et al. 2010. Mol Plant 3: 469490

CHALLENGE - 1
Large scale phenotyping under natural field A OPEN CHALLENGE

Controlled environment phenotyping: Phenomics

CHALLENGE - 2

Minimum Information about a Drought Experiment (MIADE)

1. Agronomic conditions of crop culture: Soil type, pH and Ec; nutrition;
spacing between plants
2. Soil water status: soil matric potential, amount and interval of irrigation
3. The crop growth stage at which the stress was imposed
4. Duration of stress
5. Plant water status: RWC or water potential and osmotic potential
6. Phenology, Yield and yield components
7. Weather data on rainfall, temperature and VPD

CHALLENGE - 3
Crop Functional Genomics
Rate limiting traits/processes/genes

1. Genotypes:
Germplasm Cores Mini-cores
Mutants T-DNA/Transposon tagged lines
2. Genomics:
OMICS and Bioinformatics
3. Efficient and Easy Transformation Protocols
Rate limiting step in gene function validation

Rate limiting traits/processes/genes

Yield
Yield
Component

Germplasm

Mutants

Phenotyping

Phenotyping

Component Traits
Pathways

Contrasting
genotypes

Genes

Association
mapping (LD)
WGA

Gene
cloning

QTL Mapping

GENES

MAS Breeding

Transgenics

Genotype with improved stress tolerance

CHALLENGE - 4
Yet to learn of the alphabets of the stress matrix

Potential combinations of environmental

stresses that can affect crops in the field

CHALLENGE - 5
Systems biology
OmicsCellular
networksGrowth models

YIELD

CHALLENGE - 6

Collaboration & Translation of research findings in to products

Drought tolerance can be improved

Drop by drop
Gene by Gene
Trait by Trait

Be SPECIFIC Target Environment, Pyramid Traits

? X

Thanks

One who solves the problem of WATER is worth two NOBEL Prizes
one for SCIENCE and one for PEACE
- John F. Kennedy

viswanathan@iari.res.in