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Screening and trait-based selection forScreening and trait-based selection for
drought resistance in ricedrought resistance in rice
R. Serraj & J. Cairns
OutlineOutline
• Introduction: the drought challengesIntroduction: the drought challenges
• Types of drought & functional definition of droughtTypes of drought & functional definition of drought
‘resistance/tolerance’ (DR)‘resistance/tolerance’ (DR)
• Considerations in designing a drought screeningConsiderations in designing a drought screening
programprogram
• Design & management of field drought experimentsDesign & management of field drought experiments
• Direct selection for yield under stressDirect selection for yield under stress
• Trait-based selection for drought resistanceTrait-based selection for drought resistance
• Examples and case studiesExamples and case studies
• ConclusionsConclusions
Challenges in manipulating drought
resistance (DR) genes
• Most physiological and metabolic processes are +
affected: cell growth, stomata, photosynthesis,
translocation,..
• Very large number of genes regulated by drought
• Large genetic populations required
• Even component traits are difficult to screen
• Logistical challenges for dealing with a large number of
genes even when identified
Drought stress and rice response
 Lowland/aquatic adaptation
 Shallow, compact root system
 Deep roots not efficient at water extraction
 Higher tissue sensitivity to water deficit?
 High sensitivity of grain set to stress
 Very sensitive to timing of stress
 Other mechanisms?
Rice tends to be more sensitive to
drought stress than other crops?
Timing, duration & intensity
of stress occurrence
Intermittent Drought
13-Jul 2-Aug 22-Aug 11-Sep 1-Oct 21-Oct 10-Nov
S(Waterlevel-cm)
-40
-35
-30
-25
-20
-15
-10
-5
0
5
10
15
Transplanting
Flowering
Maturity
Early Drought
13-Jul 2-Aug 22-Aug 11-Sep 1-Oct 21-Oct 10-Nov
S(Waterlevel-cm)
-40
-35
-30
-25
-20
-15
-10
-5
0
5
10
15
Transplanting
Flowering
Maturity
Intermittent drought
Early drought
13-Jul 2-Aug 22-Aug 11-Sep 1-Oct 21-Oct 10-Nov
S(Waterlevel-cm)
-40
-35
-30
-25
-20
-15
-10
-5
0
5
10
15
Flowering
Transplanting
Maturity
Late drought
Terminal drought
1st
message:
1. Know your target environment :
what type of stress is more
frequent?
Characterization of target environmentCharacterization of target environment
Available soil moisture (post-rainy season)Rainfall probability
• Use historical climate series, crop simulation & water
balance models
Breeding Target? Germination, crop establishment,
vegetative growth & tillering, flowering, grain filling
Example: variability of pearl millet TPEs
in N.W. India
How to define "drought tolerance" that can be realistically
pursued in a breeding program ?
Stable grain yield under stress
Adaptive responses to stress
Underlying biological mechanisms
Alleles coding for the mechanisms
“Resistance" has to be researchable, breakable into simpler
levels (e.g. floret sterility and ASI in maize, rooting effects vs. rooting
pattern..), should focus on heritable genetic variation, linked to
Functional definition of
‘drought resistance’
2nd
message:
1. Know your target environment :
What type of stress is more
frequent?
2. Yield stability under drought
should be the target
1. Artificially create stress in the "normal" growing season
+ avoids genotype x season problems
- excluding water is costly/difficult (reproducibility)
- no control of stress occurrence (timing, severity & duration)
2. Manage water-deficits in a non-growing season
+ application of uniform, repeatable & controlled stress
environment
+ maximize genetic component of the observed variation
- extrapolation of results to TPEs?
Drought screening environments
Combination of options?
Objectives of a screening system
1. Focus on TPE & adaptation to major stress constraints
2. Minimize problems in detecting heritable differences in
DR
3. Yield under stress as function of: yield potential, escape,
DR
4. Use drought sensitivity index (DSI) to distinguish DR
from escape & yield potential
5. DR as the ability to maintain yield components in stress
compared to non-stress:
• maintenance of grain number, grain size, biomass,
yield, harvest index
• Differentiate among genotypes
degree of yield reduction (+ 50%), manage ‘escape’ effects,
use period of max evapotranspiration (ET) demand, time
initiation of stress carefully, adjust for differences in escape
• Apply uniform stress (drought nursery approach)
“uniform” soil water profile, uniform pre-stress crop growth,
water application & water use
• Repeatable stress
standard planting time & management procedures, initiate
stress by phenological stage, stable water use environment,
dedicated field facility, well-established management
practices
Screening facility/systemScreening facility/system
• Phenology, morphology
• Growth stage at which stress occurs
• Severity and duration of stress
• Management levels and options
• Plot size, design
• Interacting factors
• Biotic stresses
• Crop quality, economics
Considerations for screeningConsiderations for screening
3rd
message:
1. Know your target environment : What type of
stress is more frequent?
2. Yield stability under drought should be the
target
3. Screening system should be
uniform, repeatable and differentiate
genotypes
• land plane field regularly
• minimize pre-stress differences
• band fertilizer with precision
• weed management over all year
• pest & disease control
• uniform stand (over-sow and
thin?)
Field management forField management for
drought screeningdrought screening
Control of field irrigationControl of field irrigation
Combinations: e.g. sprinkler irrigation in early stages, drip or surface
irrigation at later stage, precise final irrigation
Method Freq Advantages Disadvantages
Surface / Furrow ∆ /
3 x wk
Robust components
Efficient water use
Easy to measure
Inefficient water use
Requires land leveling
Hard to measure amount of water
Requires raised beds
Row spacing too wide for rice
Sprinkler 3 x wk Robust components
Efficient water use
Easy to measure
Uneven distribution pattern
same timing of stress for all entries
Large border areas needed
Leaks, blockages and wind
Drip daily Differential irrigation/ plot
Efficient water use
Uniform application
Expense of components (filters etc)
Limits weed control options
Labor cost: check quality of irrigation
1. Statistical design
• fit design to field constraints (alpha designs)
• adjust for primary & secondary gradients
• replicates & soil gradients – local spatial variation
• final irrigation differences
• sprinkler application gradients
2. Data organizing/checking:
• missing/duplicate data, expected range, detect and
correct outliers
3. Data analysis procedures:
• spatial analysis/adjustment, error variance and
heritability, best linear unbiased estimates
Design & data managementDesign & data management
4th
message:
1. Know your target environment : What
type of stress is more frequent?
2. Yield stability under drought should be
the target
3. Screening system should be uniform,
repeatable and differentiate genotypes
4. Field and irrigation management are
critical for drought screening (you
have the data you pay for…)
Soil moisture monitoring techniques:
What’s best?
IAEA 3-year study
comparing various
devices & different
cropping systems
• Soil Moisture Neutron Probe is the
preferred soil water measurement
technology:
1. measures larger volume of soil than others
- adv variability of soil water on the small
scale near to the access tube;
2. less sensitive to salinity or temperature -
large errors in capacitance or TDR systems
used in access tubes.
Alternatives (depending on needs and $):
– Gravimetry
– Tensiometers
– Thetaprobe
– Diviner-2000
Comparison of soil waterComparison of soil water
measuring techniquesmeasuring techniques
Screening Tools and Methodologies
• Line Source Irrigation
• Rain-out shelters
• Drained paddy (lowland)
• Continuous intermittent stress
(upland)
Line source moisture
gradient evaluation
Drought evaluation under
rain-out shelter
Dry-season managed environments
1. vegetative screening
Are they useful?
• can be useful as a comparative method IF
stands are good, water can be applied evenly
AND this is the type of stress that occurs in
the target environment
e.g. subsets of diverse IRRI germplasm
collection
Strategy:
• Select for lines that continue growth with early
water shortage (deep effective roots,
continued tillering) – avoid survival stage!
• Select lines that tolerate delayed
transplanting – early vigor
(e.g. Batangas area: delayed rainfall)
URV2-DS-2006:900 entries x 3 reps
Dry-season managed environments
2. reproductive-stage screening
Management is complex: interaction of phenology x stress phase
Stress management
options should be
adjusted for cultivar
phenology:
• staggered planting,
• individual plot irrigation
Applying stress at flowering
• 14-20 days of water exclusion around flowering reduces yield
by 50-80 %
• when flag leaf auricle reaches auricle of the penultimate leaf
on the first tiller, ~7 days before 50% anthesis in control plots
Effects:
• Spikelet fertility
• Pollen formation
• Panicle exertion
• Pollen germination
• Fertilization
• Embryo development/abortion
Azucena panicles 03DSAzucena panicles 03DS
Last irrigation d57 d64 d69 d73 d78 d85 d90 d95 control
Direct selection for yield componentsDirect selection for yield components
• Heritability of yield under stress is not very different from
yield in NS in case of well-managed experiments
• Need to consider yield in good years as well as in drought
years – must have yield potential experiments
• Need to reduce yield by >50% in stress treatment to get
valuable variation
• Variation in maturity and yield potential can interfere
with drought responses!
5th
message:
1. Know your target environment : What type of stress
is more frequent?
2. Yield stability under drought should be the target
3. Screening system should be uniform, repeatable
and differentiate genotypes
4. Field and irrigation management are critical for
drought screening (You have the data you pay
for…)
5. Screening procedure = f (targets, needs &
means)
(No single size fits all…)
Unsuccessful for 3 major reasons :
1. Lack of effective integrative multidisciplinary approach
associating physiology, breeding, genetics, modeling..
2. Criteria more related to survival mechanisms than to
productivity
3. Too long & unranked “shopping list” of physiological
traits to be improved
Priority as a function of the target environment
Trait-based crop yield
improvement under drought
Putative physiological traits applied in
breeding for drought tolerance
• Emergence characteristics/vigor
• Nutrient acquisition/uptake efficiency
• Leaf development & photosynthesis
• Water use efficiency (WUE) component traits
• Deep root development
• Canopy temperature/stomata regulation
• Osmotic Adjustment/relative water content
• Hormonal control: ABA, GA , Ethylene
• Stay green/senescence
• Grain number maintenance
• Grain fill duration and rate
• Harvest index under drought
• Yield & its components under drought etc…
Simple scoring methods such as leaf drying and rolling, scored around
flowering stage can be a useful secondary traits for detecting differences
in plant water status
Breeding for drought-affected
environments
Options
• direct selection for yield & components, trait-
based selection, MAS, transgenics
Strategy
• direct selection for yield has promise (slow?)
• close attention to experimental design and
management
• donor parental lines with useful traits
• secondary traits can be helpful, but with careful
and integrated approach
Marker Assisted Breeding:
Genomic tools for applied plant breeding
 Choice of parental lines for crosses
 Diversity assessment
 Choice of breeding scheme
 Required degree of recombination
 Selection criteria
 Improve heritability
 Reduce time required
 Components of more complexly
inherited traits
 Pyramiding oligo-genes
 Minimizing undesirable correlations
 LD & Associative mapping
 Phenotyping is most critical
6th
message:
1. Know your target environment : What type of stress is
more frequent?
2. Yield stability under drought should be the target
3. Screening system should be uniform, repeatable and
differentiate genotypes
4. Field and irrigation management are critical for drought
screening (You have the data you pay for…)
5. Screening procedure = f (targets, needs & means)
(No single size fits all…)
6. Trait-based improvement needs
careful choice and requires integration
of breeding & other disciplines
ConclusionsConclusions
• No quick fix to drought stress:
“the pathway exists!”
• Small and steady steps +
integration and synergy
• Improvements: site & TPE
characterization (historical
weather data), more drought-
prone sites(?), Systems
analysis, DSI, soil hydrology,
database, trials replication,
careful and yield-based trait
dissection..

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R. serraj. Screening and trait based selection for drought resistance in rice

  • 1. Screening and trait-based selection forScreening and trait-based selection for drought resistance in ricedrought resistance in rice R. Serraj & J. Cairns
  • 2. OutlineOutline • Introduction: the drought challengesIntroduction: the drought challenges • Types of drought & functional definition of droughtTypes of drought & functional definition of drought ‘resistance/tolerance’ (DR)‘resistance/tolerance’ (DR) • Considerations in designing a drought screeningConsiderations in designing a drought screening programprogram • Design & management of field drought experimentsDesign & management of field drought experiments • Direct selection for yield under stressDirect selection for yield under stress • Trait-based selection for drought resistanceTrait-based selection for drought resistance • Examples and case studiesExamples and case studies • ConclusionsConclusions
  • 3. Challenges in manipulating drought resistance (DR) genes • Most physiological and metabolic processes are + affected: cell growth, stomata, photosynthesis, translocation,.. • Very large number of genes regulated by drought • Large genetic populations required • Even component traits are difficult to screen • Logistical challenges for dealing with a large number of genes even when identified
  • 4. Drought stress and rice response  Lowland/aquatic adaptation  Shallow, compact root system  Deep roots not efficient at water extraction  Higher tissue sensitivity to water deficit?  High sensitivity of grain set to stress  Very sensitive to timing of stress  Other mechanisms? Rice tends to be more sensitive to drought stress than other crops?
  • 5. Timing, duration & intensity of stress occurrence Intermittent Drought 13-Jul 2-Aug 22-Aug 11-Sep 1-Oct 21-Oct 10-Nov S(Waterlevel-cm) -40 -35 -30 -25 -20 -15 -10 -5 0 5 10 15 Transplanting Flowering Maturity Early Drought 13-Jul 2-Aug 22-Aug 11-Sep 1-Oct 21-Oct 10-Nov S(Waterlevel-cm) -40 -35 -30 -25 -20 -15 -10 -5 0 5 10 15 Transplanting Flowering Maturity Intermittent drought Early drought 13-Jul 2-Aug 22-Aug 11-Sep 1-Oct 21-Oct 10-Nov S(Waterlevel-cm) -40 -35 -30 -25 -20 -15 -10 -5 0 5 10 15 Flowering Transplanting Maturity Late drought Terminal drought
  • 6. 1st message: 1. Know your target environment : what type of stress is more frequent?
  • 7. Characterization of target environmentCharacterization of target environment Available soil moisture (post-rainy season)Rainfall probability • Use historical climate series, crop simulation & water balance models Breeding Target? Germination, crop establishment, vegetative growth & tillering, flowering, grain filling
  • 8. Example: variability of pearl millet TPEs in N.W. India
  • 9. How to define "drought tolerance" that can be realistically pursued in a breeding program ? Stable grain yield under stress Adaptive responses to stress Underlying biological mechanisms Alleles coding for the mechanisms “Resistance" has to be researchable, breakable into simpler levels (e.g. floret sterility and ASI in maize, rooting effects vs. rooting pattern..), should focus on heritable genetic variation, linked to Functional definition of ‘drought resistance’
  • 10. 2nd message: 1. Know your target environment : What type of stress is more frequent? 2. Yield stability under drought should be the target
  • 11. 1. Artificially create stress in the "normal" growing season + avoids genotype x season problems - excluding water is costly/difficult (reproducibility) - no control of stress occurrence (timing, severity & duration) 2. Manage water-deficits in a non-growing season + application of uniform, repeatable & controlled stress environment + maximize genetic component of the observed variation - extrapolation of results to TPEs? Drought screening environments Combination of options?
  • 12. Objectives of a screening system 1. Focus on TPE & adaptation to major stress constraints 2. Minimize problems in detecting heritable differences in DR 3. Yield under stress as function of: yield potential, escape, DR 4. Use drought sensitivity index (DSI) to distinguish DR from escape & yield potential 5. DR as the ability to maintain yield components in stress compared to non-stress: • maintenance of grain number, grain size, biomass, yield, harvest index
  • 13. • Differentiate among genotypes degree of yield reduction (+ 50%), manage ‘escape’ effects, use period of max evapotranspiration (ET) demand, time initiation of stress carefully, adjust for differences in escape • Apply uniform stress (drought nursery approach) “uniform” soil water profile, uniform pre-stress crop growth, water application & water use • Repeatable stress standard planting time & management procedures, initiate stress by phenological stage, stable water use environment, dedicated field facility, well-established management practices Screening facility/systemScreening facility/system
  • 14. • Phenology, morphology • Growth stage at which stress occurs • Severity and duration of stress • Management levels and options • Plot size, design • Interacting factors • Biotic stresses • Crop quality, economics Considerations for screeningConsiderations for screening
  • 15. 3rd message: 1. Know your target environment : What type of stress is more frequent? 2. Yield stability under drought should be the target 3. Screening system should be uniform, repeatable and differentiate genotypes
  • 16. • land plane field regularly • minimize pre-stress differences • band fertilizer with precision • weed management over all year • pest & disease control • uniform stand (over-sow and thin?) Field management forField management for drought screeningdrought screening
  • 17. Control of field irrigationControl of field irrigation Combinations: e.g. sprinkler irrigation in early stages, drip or surface irrigation at later stage, precise final irrigation Method Freq Advantages Disadvantages Surface / Furrow ∆ / 3 x wk Robust components Efficient water use Easy to measure Inefficient water use Requires land leveling Hard to measure amount of water Requires raised beds Row spacing too wide for rice Sprinkler 3 x wk Robust components Efficient water use Easy to measure Uneven distribution pattern same timing of stress for all entries Large border areas needed Leaks, blockages and wind Drip daily Differential irrigation/ plot Efficient water use Uniform application Expense of components (filters etc) Limits weed control options Labor cost: check quality of irrigation
  • 18. 1. Statistical design • fit design to field constraints (alpha designs) • adjust for primary & secondary gradients • replicates & soil gradients – local spatial variation • final irrigation differences • sprinkler application gradients 2. Data organizing/checking: • missing/duplicate data, expected range, detect and correct outliers 3. Data analysis procedures: • spatial analysis/adjustment, error variance and heritability, best linear unbiased estimates Design & data managementDesign & data management
  • 19. 4th message: 1. Know your target environment : What type of stress is more frequent? 2. Yield stability under drought should be the target 3. Screening system should be uniform, repeatable and differentiate genotypes 4. Field and irrigation management are critical for drought screening (you have the data you pay for…)
  • 20. Soil moisture monitoring techniques: What’s best? IAEA 3-year study comparing various devices & different cropping systems
  • 21. • Soil Moisture Neutron Probe is the preferred soil water measurement technology: 1. measures larger volume of soil than others - adv variability of soil water on the small scale near to the access tube; 2. less sensitive to salinity or temperature - large errors in capacitance or TDR systems used in access tubes. Alternatives (depending on needs and $): – Gravimetry – Tensiometers – Thetaprobe – Diviner-2000 Comparison of soil waterComparison of soil water measuring techniquesmeasuring techniques
  • 22. Screening Tools and Methodologies • Line Source Irrigation • Rain-out shelters • Drained paddy (lowland) • Continuous intermittent stress (upland)
  • 25. Dry-season managed environments 1. vegetative screening Are they useful? • can be useful as a comparative method IF stands are good, water can be applied evenly AND this is the type of stress that occurs in the target environment e.g. subsets of diverse IRRI germplasm collection Strategy: • Select for lines that continue growth with early water shortage (deep effective roots, continued tillering) – avoid survival stage! • Select lines that tolerate delayed transplanting – early vigor (e.g. Batangas area: delayed rainfall) URV2-DS-2006:900 entries x 3 reps
  • 26. Dry-season managed environments 2. reproductive-stage screening Management is complex: interaction of phenology x stress phase Stress management options should be adjusted for cultivar phenology: • staggered planting, • individual plot irrigation
  • 27. Applying stress at flowering • 14-20 days of water exclusion around flowering reduces yield by 50-80 % • when flag leaf auricle reaches auricle of the penultimate leaf on the first tiller, ~7 days before 50% anthesis in control plots Effects: • Spikelet fertility • Pollen formation • Panicle exertion • Pollen germination • Fertilization • Embryo development/abortion Azucena panicles 03DSAzucena panicles 03DS Last irrigation d57 d64 d69 d73 d78 d85 d90 d95 control
  • 28. Direct selection for yield componentsDirect selection for yield components • Heritability of yield under stress is not very different from yield in NS in case of well-managed experiments • Need to consider yield in good years as well as in drought years – must have yield potential experiments • Need to reduce yield by >50% in stress treatment to get valuable variation • Variation in maturity and yield potential can interfere with drought responses!
  • 29. 5th message: 1. Know your target environment : What type of stress is more frequent? 2. Yield stability under drought should be the target 3. Screening system should be uniform, repeatable and differentiate genotypes 4. Field and irrigation management are critical for drought screening (You have the data you pay for…) 5. Screening procedure = f (targets, needs & means) (No single size fits all…)
  • 30. Unsuccessful for 3 major reasons : 1. Lack of effective integrative multidisciplinary approach associating physiology, breeding, genetics, modeling.. 2. Criteria more related to survival mechanisms than to productivity 3. Too long & unranked “shopping list” of physiological traits to be improved Priority as a function of the target environment Trait-based crop yield improvement under drought
  • 31. Putative physiological traits applied in breeding for drought tolerance • Emergence characteristics/vigor • Nutrient acquisition/uptake efficiency • Leaf development & photosynthesis • Water use efficiency (WUE) component traits • Deep root development • Canopy temperature/stomata regulation • Osmotic Adjustment/relative water content • Hormonal control: ABA, GA , Ethylene • Stay green/senescence • Grain number maintenance • Grain fill duration and rate • Harvest index under drought • Yield & its components under drought etc…
  • 32. Simple scoring methods such as leaf drying and rolling, scored around flowering stage can be a useful secondary traits for detecting differences in plant water status
  • 33. Breeding for drought-affected environments Options • direct selection for yield & components, trait- based selection, MAS, transgenics Strategy • direct selection for yield has promise (slow?) • close attention to experimental design and management • donor parental lines with useful traits • secondary traits can be helpful, but with careful and integrated approach
  • 34. Marker Assisted Breeding: Genomic tools for applied plant breeding  Choice of parental lines for crosses  Diversity assessment  Choice of breeding scheme  Required degree of recombination  Selection criteria  Improve heritability  Reduce time required  Components of more complexly inherited traits  Pyramiding oligo-genes  Minimizing undesirable correlations  LD & Associative mapping  Phenotyping is most critical
  • 35. 6th message: 1. Know your target environment : What type of stress is more frequent? 2. Yield stability under drought should be the target 3. Screening system should be uniform, repeatable and differentiate genotypes 4. Field and irrigation management are critical for drought screening (You have the data you pay for…) 5. Screening procedure = f (targets, needs & means) (No single size fits all…) 6. Trait-based improvement needs careful choice and requires integration of breeding & other disciplines
  • 36. ConclusionsConclusions • No quick fix to drought stress: “the pathway exists!” • Small and steady steps + integration and synergy • Improvements: site & TPE characterization (historical weather data), more drought- prone sites(?), Systems analysis, DSI, soil hydrology, database, trials replication, careful and yield-based trait dissection..

Notas do Editor

  1. Challenges in
  2. Various soil moisture monitoring equipments are available in the market, choosing the right device can be a daunting task. The SMNP has been the method for irrigation scheduling for many years, however, licensing, training of users and safety regulations pertaining to the radioactive source make their use preventive and expensive in some situations. The Joint FAO/IAEA Div. Nuclear Applications in Food and Agriculture decided that it was time to do a thorough comparison, comparing the SMNP with other major sensors such as the TDR, capacitance sensors and soil water potential devices.
  3. Advantages: simple, repeatability, dependable and reliable fast, economical, non-destructive temporal soil moisture changes can be monitored relatively easy to install large volume of soil is measured, providing more reliable measurements of soil moisture than most methods in large scale studies moisture changes with depth can be determined The conclusion of this comparative study was that most of the devices examined, worked well some of the time but all of them performed quite badly in some circumstances. The group was aware that the choice of a water measurement technology is often made for economic, convenience, and other reasons and there was a need to be able to get the best results from any device. The choice of a technology is sometimes not made by the ultimate user, or even if it is, the main constraint may be financial rather than technical.