from soil to seed: the quest for zinc-enriched rice
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From Soil to Seed
The Quest for Zinc-Enriched Rice
Sarah E.J. BeeboutSeptember 22, 2011
Why is zinc important for poverty mitigation?
1 Rice productivity•Zn deficiency:
•Stunting, poor tillering•Poor grain filling•Low yields
Prevalence?“Up to 50%” of rice soils are Zn-deficient (White and Zasoski, 1999)
2 Human nutrition
•Zn deficiency:
•Impairs cognitive development•Causes growth stunting
•Impairs immune system function•Greater susceptibility to respiratory infections, diarrhea
children
all ages
(Sandstead, 1989, 1994; Black, 2001)
(Ezzati et al., 2004)
0 2 4 6 8 10 12 14 16
TobaccoCholesterol
Blood pressureCooking stove pollution
SanitationUnsafe sex
Vitamin A deficiencyIron deficiencyZinc deficiency
Underweight
% Attributable DALYs
Burden of Disease: Least Developed Sub-region
Risk factors
Malnutrition
Prevalence?
DALY=disability adjusted life years;econometric index of time “lost” to disease
Brief history of Zn research on rice1970’s: • Zn-deficiency identified as a yield-limiting factor in
some soils (Katyal & Ponnamperuma, 1974)
• Zn fertilizer recommendations developed (Castro, 1977)
• Began screening germplasm for Zn-deficiency tolerance (Ponnamperuma, 1977)
1980’s and 1990’s: • More detailed chemical and physiological mechanisms
identified (Sajwan & Lindsay, 1986; Beggi et al., 1994)• Zn fertilizer recommendations improved (Savithri et
al., 1999)• Germplasm tolerance to moderate Zn-deficiency
improved (e.g. IR34, IR36, and many others)
Yield-limiting Zn deficiency: “SOLVED” by breeding• Inconsistent response to Zn fertilizer application• Very poor Zn fertilizer use efficiency (<1%)
Zinc fertilizers “not very popular with farmers”
Starting in the late 1990’s:biofortification=enriching food with human nutrients during plant growth
(in contrast to “fortification”=adding nutrients during food processing)
• New goal: breeding plants to improve human nutrition• CGIAR Micronutrients Project• HarvestPlus Challenge Program
Zn, Fe, vitamin Amultiple crops
Biofortification: Complements other approaches to human zinc nutrition
Dietary diversification
Fortification Supplementation
Biofortification:Increasing nutritional value of staple foods
Biofortification: Why rice? Why zinc?
Rice now: Average 16 ppm Zn (white)40-60% of calories for Asian poor10-20 % of daily Zn requirement
High-Zn rice:Natural variability up to 30 ppm Zn (unlike Fe)60-70% of daily Zn requirementImprove Zn status of 500 million people
(Hunt et al., 2002; Meenakshi et al., 2007)
Zinc target levels
seed
brown(de-hulled)
white(polished)
30 ppmfrom 20
24 ppmfrom 16
•most of Zn is retained (unlike Fe)
Convergence of Interests (2007)
Plant Zn nutrition
• with increasing rice yields:• Zn deficiency was
noticeable again• removal of biomass
was “mining” soil Zn
• desire to improve identification and correction of Zn deficiency
Biofortification• Zn fertilizer application
enriched grain Zn in wheat (Cakmak, 2007)
• desire in HarvestPlus to explore Zn fertilizer application to other crops
First soil science objective for biofortification collaboration:
Manage rice for better zinc uptake and higher grain zinc
Approach:
Test Zn deficiency management recommendations on grain Zn
From Zn deficiency literature
Plant symptoms were worse with:•long-term flooding•high soil organic matter
(Forno et al., 1975)
use up O2 in soil=redox potential decreases
•but Zn2+ does not change form with redoxchanges (unlike Fe3+ to Fe2+)
Zn fertilizer has limited usefulness in flooded soils:
0102030405060
-5 15 35 55 75 95
Low (native)
Medium (10 kg/ha)
High (50 kg/ha)
Avai
labl
e so
il Zn
(mg/
kg)
Days
(Johnson-Beebout et al., 2009)
Hypotheses applying Zn-deficiency research to biofortification
1. Higher grain Zn if soil is non-flooded (than if continuously flooded)
2. Higher grain Zn with later (than basal) Zn fertilizer application
3. Higher grain Zn of genetically biofortified lines with optimized agronomic management (than without)
Greenhouse experiment (2008)Objective:To test the effect of the timing of Zn fertilizer
application on grain Zn content, relative to:– Plant growth stage– Flooding/draining periods
Design and management: Jack Jacob, Efren Laureles, Oliver Castillo
Implementation and data collection:Jerone Onoya, Max Alumaga, Angel Bautista, Briccio Salisi
Results: Soil
0
10
20
30
40
-20 0 20 40 60 80 100 120Avai
labl
e so
il Zn
(mg/
kg)
Days after transplanting
Flooded
Mid-season drainage
Late-season drainage
Zn fertilization1-week drainage periods
Soil Zn is highest during drainage periods, regardless of when applied
(as expected based on Zn-def research)
Results: Grain Zn (IR69428)
25
30
35
40
Z0 ZB ZM ZL
Zn c
once
ntra
tion
(mg/
kg) continuous flooding
mid-season drainagelate-season drainage
•IR69428-6-1-1-3-3 (biofortified) was above target Zn level under all conditions of this experiment
•Zn was increased from 31 to 37 ppm by adding Zn during a late season drainage period
brow
n ric
e
Field experiment in Zn-deficient field in Pila,Laguna (DS 2009)
Farmer: Domingo NidoDesign and management: WencyLarazo, Oliver Castillo, Dennis Tuyogon
Implementation and data collection:Jerone Onoya, Angel Bautista, Ike Reyes, Juan Puno, Briccio Salisi
Objective:To test the effect of the different amounts of Zn
fertilization, with:– Continuous flooding– Alternate wetting and drying (AWD)
Results: Soil
-300-200-100
0100200300400500600
Eh (m
V)
Redox potentialAlternate Wetting and DryingContinuous Flooding
0
1
2
3
4
5Soil Zn availability
soil
Zn (m
g/kg
)
AWD vs. CF:
•higher redox (i.e. more oxidized)•higher Zn availability
Results: Grain Zn, NSIC 158
15
20
25
AWD CF
a
b
15
20
25
Z0 Z1 Z2 Z3
aabc
bc
Zn co
ncen
trat
ion
(mg/
kg)
brow
n ric
e
•water management effect : AWD>CF by 2 ppm
•small Zn fertilizer effect (<1 ppm), only at highest Zn application rate
“Aha!” moments
• water management (only as dry as “safe AWD”) affected grain Zn of a non-biofortified variety
…investigate AWD further
• even with optimized management, NSIC 158 only had 23 ppm Zn, compared with 37 for IR69428
…management can’t replace breeding
Discussion with breeders
“We have further increased grain Zn content by improving water and zinc fertilizer management…”
“But only in 1 (biofortified) genotype”“…and the soil science hypotheses have been
confirmed, as well as their relevance to grain Zn.”
“But only in 1 genotype”
New objective for biofortificationcollaboration:
Understand how genotypes differ in grain Zn in differing soil environments
Approach:
Test Zn management strategies on multiple genotypes to determine basic physiological Zn uptake characteristics
Greenhouse experiment with 2 biofortified genotypes (2009)
Design and management:Oliver Castillo, Efren LaurelesData analysis: Francis Rubianes
Implementation and data collection: Jerone Onoya, Max Alumaga, Angel Bautista, Ike Reyes, Briccio Salisi
Objective:To test the effects of Zn fertilizer and water
management (without late season drainage) on:– IR69428 (as in earlier greenhouse experiment)– IR68144
Results: Grain Zn
25
30
35
40
no added Zn Zn at mid-tillering Zn at heading
Zn c
onc
(mg/
kg )
brow
n ric
eIR69428
IR68144
ab
c c cd
d
•no effect of 1-week mid-season drainage on grain Zn this time •IR69428 responded to late-season Zn application•IR68144 responded to vegetative stage Zn application
Implication:•IR69428 continues taking up Zn during grain-filling•IR68144 does not
Remobilization vs. Root Uptake
Zn
Zn
•Get lots of Zn into plant early
•Ensure sufficient Zn supply during grain-filling
Previous lit: no remobilization in rice (unlike wheat)
•Two aerobic rice genotypes: Zn-65 tracer studies showed no movement of labeled Zn from leaf to grain (Jiang et al., 2007)
•Nipponbare: Zn-62 tracer imaging study showed Zn moving from root to grain in 3.5 h, bypassing the flag leaf (Tsukamoto et al., 2006)
Physiologists join the collaborative biofortification effort
Starting point:• Several genotypes tolerant of extreme Zn
deficiency, with physiological data for some (RK Singh, Glenn Gregorio, Matthias Wissuwa, Abdel Ismail)
…but unknown grain Zn• Several genotypes with high grain Zn
(Parminder Virk, Tajinder Singh, Deepinder Grewal)
…but unknown Zn-deficiency tolerance
Post-doctoral work of Somayanda Impa
Hypotheses:Root Zn uptake mechanisms
active at
Genotypecategory
Seedling Grain-filling
Zn-deficiency tolerant ***
Zn-biofortified:remobilization ***
Zn-biofortified: direct uptake ***
Management and Data Collection: MJ Morete, Randell Eusebio
Results preview
Early season Znefficiency
(%)
Grain Zn attributable to different uptake mechanisms
(%)
Genotype Category RemobilizationDirect Uptake
RIL-46 Zn-deficiency tolerant 88 11 89
IR74 Zn-deficiency susceptible 49 32 68
IR68144 Zn-biofortified: remobilization
58 100 0
IR82247 100 100 0
SWHOO 98 100 0
IR69428 Zn-biofortified: direct uptake
93 0 100
Joryoongbyeo 75 0 100
IR64 Popular check 90 31 69
Foliar Zn application (DS2011)Physiology and Zinc Management
Rationale: Foliar Zn application would bypass soil chemistry challenges
Hypotheses:1) “Remobilization” genotypes are more likely than “direct
uptake” to respond to foliar Zn by increasing grain Zn2) Any genotype is more likely to respond in a Zn-deficient soil
Design and management: Ranee Mabesa
Implementation and data collection:Jerone Onoya, Rowell Mayores, Nards Baclao
Results: Grain ZnZn-sufficient site Zn-deficient site
Genotype Category >30 ppm Zn?Responsive to foliar Zn?
>30 ppmZn?
Responsive to foliar Zn?
NSIC222Popular check
No No No No
IR64 Almost (+spray) Yes No Yes
IR68144 Biofortification: remobilization Yes (+spray) Yes Almost
(+spray) Yes
IR83668
Biofortification: unknown
Yes No No No
IR85800 No No No No
IR91152AC-79 Yes No (no filled grains)
IR83317AC-124 Yes (+spray) Yes No Yes
IR91143AC-4 Yes (+spray) Yes No Yes
IR83317AC-25Zn-deficiency
tolerant
No No No No
IR64196 No No (not planted)
IR75862 (not planted) No No
Results: Grain ZnZn-sufficient site Zn-deficient site
Genotype Category >30 ppm Zn?Responsive to foliar Zn?
>30 ppmZn?
Responsive to foliar Zn?
NSIC222Popular check
No No No No
IR64 Almost (+spray) Yes No Yes
IR68144 Biofortification: remobilization Yes (+spray) Yes Almost
(+spray) Yes
IR83668
Biofortification: unknown
Yes No No No
IR85800 No No No No
IR91152AC-79 Yes No (no filled grains)
IR83317AC-124 Yes (+spray) Yes No Yes
IR91143AC-4 Yes (+spray) Yes No Yes
IR83317AC-25Zn-deficiency
tolerant
No No No No
IR64196 No No (not planted)
IR75862 (not planted) No No
Discussion with breeders: Divided by a common language
“G (genotype) x E (environment) interactions”
Soil scientists: opportunity for optimizing management(goal: absolute yield target)
Plant breeders: inconsistent genotype performance(goal: find best-performing genotypes)
(Brooks, 2010)
“Aha!” momentsHarvestPlus Rice Meeting (2009)
One breeder presented:• grain Zn data from 12 sites• soil data from each site for every routinely measured
parameter (pH, texture, exchangeable bases, etc. )• no correlations of grain Zn with any of the soil data
•routine soil lab testing is not helpful because it doesn’t measure redox-relevant parameters
…there must be a better way to assess E
Another new objective for biofortification collaboration
Determine which E variables have greatest impact on GxE patterns
New approach Design GxE trials collaboratively, with
locations chosen for Zn characteristics
Zinc GxE experiments (DS2011)
Five locations in Philippines, representing:• Zn-depleted (IRRI)• Excessively submerged (Bukidnon, Mindanao)• Peaty (Bay, Laguna)• Calcareous (Bohol)• Zn-sufficient (PhilRice, Nueva Ecija)
Experiment design: Deepinder Grewal, Glenn Gregorio, Sarah BeeboutExperiment management: Eric Clutario, Andy Sajise, Francis Rubianes
At each site:Two sets of plant breeding trials:• Zn biofortification advanced lines, with and without
Zn fertilizer (from Deepinder)• Zn-deficiency tolerant advanced lines (from Glenn)
Environmental data collection:• Water status throughout• Background soil characterization• Detailed soil monitoring in lab
Plant data collection:• Grain Zn• Grain yield
No results yet (data being compiled)
Stages of interdisciplinary collaboration
1. Discussion of experiment results:– seminars, project meetings– “aha!” moments that helped set research
objectives
2. Input into experiment design:– breeders nominating genotypes for Zn
management trials– soil scientists recommending Zn fertilizer and
water management for plant breeding trials
(Brooks and Johnson-Beebout, in press)
3. Joint experiments:– soil scientists influence design of breeding trials
(location, + Zn fertilizer, data collection)– breeders choose genotypes by standard breeding
protocols (compromising on the number) – division of labor for implementation
Challenges of stage 3• time: coordinating three research groups• money: sorting out budgets fairly• respect: sufficient mutual understanding of the
other disciplines to listen to design ideas
Conclusions about plant breeding strategies
1. Pre-release varietal GxE trials: use sites differing in Zn availability
minimum set: depleted, submerged, calcareous
2. Screen popular (old) varieties at same sites to rank Zn-deficiency tolerance
3. Biofortification: select for both remobilization and direct uptake
– improve GxE performance – make management more flexible
Conclusions about Zn management
1. Trend towards drier water management will improve Zn uptake
2. Standard Zn fertilizer recommendation (10 kg Zn/ha applied basally) is not helpful
•Better for researchers: apply Zn to non-flooded soil at maximum tillering•Better for farmers: ??? first need is to predict likelihood of Zn deficiency
AcknowledgementsFunding: HarvestPlus; Swiss Agency for Development and
Cooperation (SDC), through their Research Fellow Partnership Program (Impa’s post-doctoral fellowship)
Interactive discussions about Zn research: Gerard Barry, Achim Dobermann, Matthias Wissuwa, Ismail Cakmak, Wolf Pfeiffer, Untung Susanto, Andrew Green
Jason and Miriam Soils Group
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