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The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
www.plantcell.org/cgi/doi/10.1105/tpc.111.tt0511 1
© 2013 American Society of Plant Biologists
Genetic Improvements in Agriculture
From Hunter Gatherer to Green Revolution and Beyond
www.plantcell.org/cgi/doi/10.1105/tpc.111.tt0511© 2013 American Society of Plant Biologists
Image credit: Smithsonian Institution Human Origins Program
•Life on earth is about four billion years old•Homo sapiens emerged as a species about 300,000 years ago
Cro magnon skull Homo sapiens
~ 30,000 years old
© 2013 American Society of Plant Biologists
•Human activities have caused vast changes in the physical, chemical, geological, atmospheric and biological realm of our planetCro magnon skull
Homo sapiens~ 30,000 years old
Image credit: Smithsonian Institution Human Origins Program
© 2013 American Society of Plant Biologists
We’ve removed mountains
1984 2009
The Hobet coal mine in Boone County, West Virginia spreads over 10,000 acres (15.6 square miles)
NASA images by Robert Simmon, based on Landsat 5 data from the USGS Global Visualization Viewer.
2 km
1987
2006
Photo credit: NASA/Goddard Space Flight Center Scientific Visualization Studio United States Geological Survey and Le Grand Portage
+ View movie of the dam's
construction
Three Gorges Dam, the world's largest hydroelectric power generator. The reservoir just upstream of the dam is more than 2 miles (3 km) across
...dammed rivers...
Dodo by Edouard Poppig, 1841
The dodo (Raphus cucullatus) was a large flightless bird indigenous to Mauritius. The arrival of humans led to the dodo’s extinction by the end of the 17th century.
....caused extinctions.....
© 2013 American Society of Plant Biologists
...modified other species in extraordinary ways
?
© 2013 American Society of Plant Biologists
... and modified plant genomes for thousands of years......
Image credits: P. Cos, Cacaphony, USDA, CIMMYT
© 2013 American Society of Plant Biologists
1850 1900 1950 20001800
Atmospheric CO2
500 1000 1500 20000
Global population
Now we face our biggest challenges
How do we feed more people without further damaging our planet?
Photo courtesy Earth Observatory NASA
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
www.plantcell.org/cgi/doi/10.1105/tpc.111.tt0511 2
© 2013 American Society of Plant Biologists
How do we feed more people with fewer resources available?
Fess et al. (2011) Crop breeding for low input agriculture: a sustainable response to feed a growing world population. Sustainability 3: 1742-1772Photo courtesy Earth Observatory NASA
© 2013 American Society of Plant Biologists
What is the role of plant breeding in addressing global challenges?
Photo credits: Xochiquetzal Fonseca/CIMMYT and IRRI
© 2013 American Society of Plant Biologists
• The Distant Past
• Crop plant domestication and beyond
• The Recent Past
• Hybrid seed
• The (First) Green Revolution
• Advances in breeding technologies
• Now and Into The Future
• Breeding for improved human health
• Breeding for drought tolerance
• Agricultural innovation in Africa
• The Second Green Revolution
GENETIC IMPROVEMENTS IN AGRICULTURE
© 2013 American Society of Plant Biologists
The Distant Past (>10,000 years ago to 1900)
Karol Schauer
• Homo sapiens originated 400,000 –250,000 years ago
• Major crops were domesticated ~ 10,000 –5000 years ago
• The development of human civilizations is correlated with the development of agriculture
© 2013 American Society of Plant Biologists
Plant domestication followed the end of the most recent glacial period
Allaby, R.G., Fuller, D.Q., and Brown, T.A. (2008) The genetic expectations of a protracted model for the origins of domesticated crops. Proc. Natl. Acad. Sci. USA 105: 13982-13986, copyright National Academy of Sciences USA
Glacial retreatGlacial advance
Wild gathering Pre-domestication cultivation
Domestication
Neolithic Revolution
thousand years ago
© 2013 American Society of Plant Biologists
How did people begin to cultivate plants?
X X
X
It is thought to have been a gradual change from seeking and following food sources
© 2013 American Society of Plant Biologists
X
X
How did people begin to cultivate plants?
It is thought to have been a gradual change from seeking and following food sources to semi settled migration
© 2013 American Society of Plant Biologists
X
How did people begin to cultivate plants?
It is thought to have been a gradual change from seeking and following food sources to semi settled migration and finally permanent settlements.
© 2013 American Society of Plant Biologists
Plants were domesticated in parallel in several regions
Reprinted by permission from Macmillan Publishers Ltd.: [Nature] Diamond, J. (2002). Evolution, consequences and future of plant and animal domestication. Nature 418: 700-707, copyright 2002.
Wheat, barley, pea, lentil~ 13,000 years ago
Rice, soybean ~ 9000 years ago
Rice, bean ~ 8500 years ago
Corn, squash, bean, potato ~ 10,000 years ago
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
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© 2013 American Society of Plant Biologists
Genetic modification arose as a consequence of cultivation
Natural variation within population
Image courtesy of University of California Museum of Paleontology, Understanding Evolution - www.evolution.berkeley.edu
Planting seeds from “good” plants increased their representation in subsequent generations
© 2013 American Society of Plant Biologists
During maize domestication cobsize increased
Photo © Robert S. Peabody Museum of Archaeology, Phillips Academy, Andover, Massachusetts. All Rights Reserved.
Cobs from archeological
sites in the Valley of Tehuacan,
Mexico
7000 years ago
500 years ago
© 2013 American Society of Plant Biologists
The hard casings around many grains were eliminated
Photo by Hugh Iltis; Reprinted from Doebley, J.F., Gaut, B.S., and Smith, B.D. (2006). The Molecular Genetics of Crop Domestication. Cell 127: 1309-1321, with permission from Elsevier.
Teosinte, the wild relative of maize, has hard coverings over each grain. Humans selected against these during maize domestication.
© 2013 American Society of Plant Biologists
Decrease in branching and increase in seed size were also selected for
Image credit Nicolle Rager Fuller, National Science Foundation
© 2013 American Society of Plant Biologists
Seeds that don’t break off were selected
WildShattering grain“Brittle rachis”Advantage –maximizes seed dispersal
DomesticatedNon-shattering grain
“Tough rachis”Advantage –
facilitates harvesting
From Konishi, S., Izawa, T., Lin, S.Y., Ebana, K., Fukuta, Y., Sasaki, T., and Yano, M. (2006). An SNP caused loss of seed shattering during rice domestication. Science 312: 1392-1396. Reprinted with permission from AAAS.
© 2013 American Society of Plant Biologists
Many of our crops are products of extensive genomic rearrangements
From Dubcovsky, J. and Dvorak, J. (2007). Genome Plasticity a Key Factor in the Success of Polyploid Wheat Under Domestication. Science. 316: 1862-1866. Reprinted with permission from AAAS. Brassica figure from Adenosine
Common wheat is the result of interspecific hybridization between
three ancestors
Polyploid (multi-genome) plants are often bigger and so
selected for propagation
The brassicas share three genomes recombined in various ways
© 2013 American Society of Plant Biologists
Domestication through genome modification gave us modern crops
From Dubcovsky, J. and Dvorak, J. (2007). Genome Plasticity a Key Factor in the Success of Polyploid Wheat Under Domestication. Science. 316: 1862-1866. Reprinted with permission from AAAS.
PA CHIAMSERAUP
FORTUNA BESAR 15 M ARONG UNKNOWNPAROC
BLUE ROSE
BPI 76 REXORO SUPREM EKITCHILI SAM BA
SINAWPAGH
UNKNOWNCINA LATISAIL TEXAS RSBR GEB24
PATNA BLUE BONNETPETA
DGWG CP231 SLO 17 BENONG
IR86 CP SLO 17 SIGADIS
IR95IR127
IR8 CHOW SUNG IR262
IR1103 TADUKAN VELLAIKARIR400 TSAI YUAN CHUNG
IR1006 M UDGOTETEP
IR1163 IR238 TN1
IR1416 IR1641IR1402
IR22 TKM 6 IR746AIR1704
O. nivaraIR1870 IR1614
IR2006 IR579 IR747 IR24/ IR661 IR1721
IR773 A BPI 121 GAM PAI
IR1915 B IR1833 GAM PAI 15 IR1561 IR1737
IR1916 IR833 IR2040
IR2146 IR 2055IR2061
IR5236 IR5338 Ultimate LandracesGAM PAI TSAI YUAN CHUNG
IR5657 DEE GEO WOO GEN BENONGCINA Unknow n
IR18348 LATISAIL CHOW SUNGTADUKAN MUDGO
IR64 KITCHILI SAMBA TETEPPA CHIAM SINAWPAGHSERAUPBESAR 15 UNKNOWN (JAPANESE)NAHNG MON S 4 O. nivara (IRGC 101508)VELLAIKAR MARONG PAROC
CO 18
NAHNG M ON S4
NM S 4
IR 64
original rice genome
Mutations
Recombinations Translocations
Deletions
Inversions
One of the most widely grown crops, indica rice IR64 is the product of a complex breeding program that has caused extensive genomic modification, mutation, deletion and rearrangement
Slide courtesy of Ingo Potrykus
© 2013 American Society of Plant Biologists
The myth of natural food
The food we eat comes from plants already extensively modified from their original form. Even heritage varieties are extensively genetically modified.
Credit: Nicolle Rager Fuller, National Science Foundation
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
www.plantcell.org/cgi/doi/10.1105/tpc.111.tt0511 4
© 2013 American Society of Plant Biologists
The Recent Past –Scientific Plant Breeding
The twentieth century took us from gas lamps to Google and steamships to space shuttles
© 2013 American Society of Plant Biologists
6 billion
1.6 billion
1234567
1300 1400 1500 1600 1700 1800 1900 2000
World Population (billions)
The twentieth century took us from gas lamps to Google and steamships to space shuttles
And the world population
quadrupled in just over 100 years
The Recent Past –Scientific Plant Breeding
© 2013 American Society of Plant Biologists
The Recent Past –Scientific Plant Breeding
Improvements in plant propagation and breeding were needed to keep up with population growth
Photo credits: Gartons Plant Breeders
© 2013 American Society of Plant Biologists
Mendel and Darwin paved the way for scientific plant breeding
© 2013 American Society of Plant Biologists
The development of hybrid corn led to a big increase in yields
A BB x A A x B
The progeny of two genetically different parents often show enhanced growth – this effect is termed “hybrid vigor” (also known as heterosis)
Shull, G.H. (1909) A pure line method in corn breeding. Am. Breed. Assoc. Rep. 5, 51–59 by permission of Oxford University Press.
© 2013 American Society of Plant Biologists
Hybrid corn was rapidly adopted because of its increased yields
A BB x A A x B
Percentage of total corn acreage
Even though farmers had to purchase seed every year, increased yields more than
offset increased costs
Shull, G.H. (1909) A pure line method in corn breeding. Am. Breed. Assoc. Rep. 5, 51–59 by permission of Oxford University Press.
© 2013 American Society of Plant Biologists
Norman Borlaug, “father of the green revolution”
Distinguished plant breeder and Nobel LaureateNorman Borlaug 1914-2009
One of the most significant accomplishments of 20th
century science was the development of lodging-resistant, high-yielding semi-dwarf grain varieties
© 2013 American Society of Plant Biologists
Improved green-revolution plants dramatically increased crop yields
The introduction of disease-resistant,
semi-dwarf varieties turning countries from
grain importers to grain exporters
Source: FAO via Brian0918
Dwarf wheat was developed at CIMMYT – the International Maize and Wheat Improvement Center
© 2013 American Society of Plant Biologists
CGIAR is an international organization of agricultural research groups
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
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© 2013 American Society of Plant Biologists
Rice breeding at IRRI also brought huge yield increases
0
0.5
1
1.5
2
2.5
3
3.5
4
4.5
1961 20001980
World rice yield (ton/ha) (FAO)
Photo courtesy IRRI
IR8, released in 1966, “…was to tropical rices what the Model T Ford was to automobiles.” It was
known as “miracle rice” because of its high yields.
© 2013 American Society of Plant Biologists
Crop productivity has kept pace with population because of increased
yields
Burney, J.A., Davis, S.J., and Lobell, D.B. (2010). Greenhouse gas mitigation by agricultural intensification. Proc. Natl. Acad. Sci. 107: 12052-12057.
Population (billions) Crop area (hectare)Crop production (gigaton)
Crop area has not increased as rapidly as crop production, because yields (food per hectare) have increased
Growing more food without using more land helps mitigate climate change and slow the loss of biodiversity
100% increase
>100% increase
~20% increase
© 2013 American Society of Plant Biologists
Modern plant breeders use molecular methods including DNA
sequencing and proteomics as well as field studies
Photo credits Scott Bauer USDA; CIMMYT; IRRI; RCMI; Duke Institute for Genome Sciences and Policy
© 2013 American Society of Plant Biologists
Advances in genetic technologies contribute to improved plants
• Marker assisted selection • Genome-wide association studies• Recombinant DNA technology and transgenic plants• Cisgenics and intragenics• Transgrafting• Precision genome editing
Photo credit: IRRI
© 2013 American Society of Plant Biologists
Marker assisted selection (MAS)
Phenotype: physical expression of traits
Genotype: sequence of all the genes in a genome
Photo credit LemnaTec; Anderson, L.K., Lai, A., Stack, S.M., Rizzon, C. and Gaut, B.S. (2006). Uneven distribution of expressed sequence tag loci on maize pachytene chromosomes. Genome Research. 16: 115-122.
© 2013 American Society of Plant Biologists
Marker assisted selection (MAS)
Phenotype: physical expression of traits
Genotype: sequence of all the genes in a genome
Selecting for DNA markers is faster than
selecting for phenotype
Photo credit LemnaTec; Anderson, L.K., Lai, A., Stack, S.M., Rizzon, C. and Gaut, B.S. (2006). Uneven distribution of expressed sequence tag loci on maize pachytene chromosomes. Genome Research. 16: 115-122.
© 2013 American Society of Plant Biologists
How markers work: Each generation, genes reassort or shuffle
Markers let us “see” which genes each
individual has inherited
© 2013 American Society of Plant Biologists
Example: Introgression of a disease resistance gene
Elite tomato Poor tomato but disease resistant (resistance gene indicated)
We want to add a disease resistance trait to an “elite” tomato plant.
© 2013 American Society of Plant Biologists
Example: Introgression of a disease resistance gene
We cross the two plants. Some of their progeny inherit the disease resistance trait, some don’t – how can we tell the difference?
Photo by Stephen Ausmus USDA
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
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© 2013 American Society of Plant Biologists
Example: Introgression of a disease resistance gene
We can use markers to look at their DNA and identify those with the resistance gene.It’s faster and easier than infecting them to see the phenotype
© 2013 American Society of Plant Biologists
Example: Introgression of a disease resistance gene
Is this an elite, disease-resistant tomato? No, half of its genes are from the poor tomato
© 2013 American Society of Plant Biologists
Example: Introgression of a disease resistance gene
We have to repeatedly cross back to the elite tomato, using markers to identify plants with the disease resistance gene
© 2013 American Society of Plant Biologists
Example: Introgression of a disease resistance gene
After several generations, elite, disease resistant tomato
Markers greatly accelerate breeding
programs
© 2013 American Society of Plant Biologists
MAS as a tool in production of submergence tolerant rice (Sub1)
Many rice-growing regions are prone to flooding. In Pakistan a 2010 a huge, deadly, flood submerged 17 million acres (69,000 km2) and destroyed much of the harvest
Photo credits: Abdul Majeed Goraya / IRIN; NASA Goddard
July 2010
Sept. 2010
50 km
© 2013 American Society of Plant Biologists
Submergence-tolerant rice can survive floods as long as 17 days
Sensitive rice – cannot survive prolonged flooding
Submergence-tolerant Sub1 rice – growth arrests during flooding, enhancing survival
FLOODING Water retreats
© 2013 American Society of Plant Biologists
Production of Swarna–Sub1:Cross Swarna with Sub1 donor
After several generations, Swarna-Sub1
Swarna
Sub1
Swarna is a high-yielding rice variety but sensitive to flooding
© 2013 American Society of Plant Biologists
Reprinted by permission from Macmillan Publishers Ltd. (NATURE) Xu, K., Xu, X., Fukao, T., Canlas, P., Maghirang-Rodriguez, R., Heuer, S., Ismail, A.M., Bailey-Serres, J., Ronald, P.C., and Mackill, D.J. (2006). Sub1A is an ethylene-response-factor-like gene that confers submergence tolerance to rice. Nature 442: 705-708. Photo couresy of Adam Barclay CPS, IRRI Photo.
Swarna –high yielding,flooding sensitive
Submergence tolerant parent
Swarna-Sub1
MAS allowed the Sub-1 trait to be rapidly introgressed into Swarna. The Swarna-Sub1 rice accounted for over ¼ of the rice planted in India in 2010.
© 2013 American Society of Plant Biologists
Advances in genomics technologies facilitate breeding for complex traits
•Genome sequence data are available for more than 20 plant species•Molecular breeding and mapping tools are developed for many species•Genome-wide association studies help match genes to traits
Anderson, L.K., Lai, A., Stack, S.M., Rizzon, C. and Gaut, B.S. (2006). Uneven distribution of expressed sequence tag loci on maize pachytene chromosomes. Genome Research. 16: 115-122.
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
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© 2013 American Society of Plant Biologists
Genome sequence data are available for many important plants
Maize
© 2013 American Society of Plant Biologists
Phenotype analysis
Genotype analysis
Genome-wide methods make it possible to identify genes associated with complex traits, like yield or water use efficiency
Association analysis
Gene discovery
© 2013 American Society of Plant Biologists
This approach allows hundreds of genes with small effects to be
identified
Reprinted by permission from Macmillan Publishers Ltd. Tian, F., Bradbury, P.J., Brown, P.J., Hung, H., Sun, Q., Flint-Garcia, S., Rocheford, T.R., McMullen, M.D., Holland, J.B., and Buckler, E.S. (2011). Genome-wide association study of leaf architecture in the maize nested association mapping population. Nat Genet 43: 159-162.
In maize, grain yields are correlated with leaf angle and size. A genome-wide association survey (GWAS) revealed hundreds of single-nucleotide polymorphisms (SNPs) associated with these traits, providing invaluable information for breeders.
© 2013 American Society of Plant Biologists
GWAS reveals SNPs that contribute to disease resistance
Reprinted by permission from Macmillan Publishers Ltd Kump, K.L., Bradbury, P.J., Wisser, R.J., Buckler, E.S., Belcher, A.R., Oropeza-Rosas, M.A., Zwonitzer, J.C., Kresovich, S., McMullen, M.D., Ware, D., Balint-Kurti, P.J., and Holland, J.B. (2011). Genome-wide association study of quantitative resistance to southern leaf blight in the maize nested association mapping population. Nat Genet 43: 163-168.
Similar studies have led to the identification of genes contributing to other agronomicallyimportant traits including drought tolerance
© 2013 American Society of Plant Biologists
Genetic Modification (GM) is another breeding method
Elite tomato Disease resistant plant (need not be same species)
Elite, disease resistant tomato
Recombinant DNA (or GM) allows a single
gene to be introduced into a genome. This
method can be faster than conventional
breeding
© 2013 American Society of Plant Biologists
Source of gene (disease-resistant
plant)
Gene of interest
Isolate gene of interest using molecular biology methods
Recombine into recipient plant DNA
Once a gene is introduced into the plant genome it functions like any other gene
© 2013 American Society of Plant Biologists
Molecular breeding
Why use GM methods sometimes and molecular breeding others?
1. Desired trait must be present in population
2. Genetic resources must be available
3. Plant should be propagated sexually
Photo credits: Gramene.org
© 2013 American Society of Plant Biologists
Why use GM methods sometimes and molecular breeding others?
Molecular breeding
1. Desired trait must be present in population
2. Genetic resources must be available
3. Plant should be propagated sexually
GM
1. Gene can come from any source
3. Plant can be propagated vegetatively2. Genetic resources not
requiredPhoto credits: Gramene.orgPhoto credits: Gramene.org ETH Life International
© 2013 American Society of Plant Biologists
GM Example: Disease resistant banana via pepper gene
Resistant SusceptibleBanana bacterial wilt (Xanthomonas campestris pv. musacearum) is destroying plants in eastern Africa. Transgenic plants carrying a resistance gene from pepper are resistant to the disease
Tripathi, L., Mwaka, H., Tripathi, J.N., and Tushemereirwe, W.K. (2010). Expression of sweet pepper Hrap gene in banana enhances resistance to Xanthomonas campestris pv. musacearum. Molecular Plant Pathology 11: 721-731.
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
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© 2013 American Society of Plant Biologists
GM Example: Insect resistance through introduction of the Bt gene
Wild-type peanut plant Peanut plant expressing the Bt gene
Photo by Herb Pilcher USDA
© 2013 American Society of Plant Biologists
Bacillus thuringiensis (Bt) bacteria produce insecticidal proteins
Bacillus thuringiensis expressing Bt toxin
Plant cell expressing Bt
toxin
Bacillus thuringiensis expressing insecticidal Bt toxin can be sprayed onto plants
Or the plants can be engineered to express the Bt gene coding for Bt toxin
© 2013 American Society of Plant Biologists
The effect of Bt toxin is highly specific
Bacillus thuringiensis expressing Bt toxin
Plant cell expressing Bt
toxin
Intestine
The Bt toxin affects only some insects because to be effective it has to be processed and bind to a specific receptor protein
Processing
Receptor binding
Intestinal lumen
© 2013 American Society of Plant Biologists
The effect of Bt toxin is highly specific
Bacillus thuringiensis expressing Bt toxin
Plant cell expressing Bt
toxin
After binding, the insecticidal proteins assemble to form a pore in the lining of the insect intestine which kills the insect
Pore assembly
Intestine
© 2013 American Society of Plant Biologists
Photo credit: Doug Buhler, Bugwood.org
Plants compete with other plants for sunlight and nutrients. Many farmers use herbicides to eliminate weeds (undesired plants) from their fields.
Left – corn rows sprayed with herbicide to eliminate competing plantsRight – corn being choked by giant foxtail (Setaria faberi)
GM Example: Herbicide resistance
© 2013 American Society of Plant Biologists
Herbicide tolerant plants are environmentally friendly
Cerdeira, A.L. and Duke, S.O. (2006). The Current Status and Environmental Impacts of Glyphosate-Resistant Crops. J. Environ. Qual. 35: 1633-1658. Photo credit Hunt Sanders, University of Georgia, bugwood.org.
Farmers that plant herbicide-tolerant crop plants use less herbicide, herbicides that are less toxic, and till (plow) less, saving soil and fuel.
Soybean
*
*
© 2013 American Society of Plant Biologists
Gene flow through pollen movement has to be monitored and controlled
Howard F. Schwartz, Colorado State University, Bugwood.org
There have been confirmed cases of gene transfer from crops to weeds and vice versa.• What consequences are
expected from gene flow?• How can gene flow be
minimized? • How can consequences be
mitigated?
© 2013 American Society of Plant Biologists
Emerging technologies circumvent some concerns about transgenics
In the conventional approach to transgenic plant production, a large piece of DNA, derived from several sources, is inserted randomly into the genome
Open Reading Frame
5′ UTR 3′ UTRPromoter
Plant genome
Transgene cassette
The promoter, 5′ and 3′ untranslated regions (UTRs) and protein coding region may all come from different sources
All the cells carry the
transgene
© 2013 American Society of Plant Biologists
Cisgenics: Genes from the same or closely-related species
Schouten, H.J., Krens, F.A. and Jacobsen, E. (2006). Cisgenic plants are similar to traditionally bred plants. EMBO Rep. 7: 750-753.
Apple gene
Apple geneApple gene
That might mean that little foreign DNA is introduced (“cisgenic”)
Some traits can be modified by the introduction of a cisgene –a gene from the same or closely-related species
Or, bacterial and viral DNA may be included, but no protein-coding regions from other organisms (“intragenic”)
Small amounts of DNA from T-DNA borders may be incorporated
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
10/17/2013
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© 2013 American Society of Plant Biologists
Cisgenics can add or silence genes
RNA silencing
ON
OFF
Resistance gene
One application of cisgenics is to add resistance genes
Advantages: Avoids lengthy backcrossing processParticularly useful for plants propagated vegetatively, such as potato or apple
Disadvantages: Gene must exist in gene pool
Gene silencing can be induced by introduction of antisense or hairpin RNA,
or overexpression of an endogenous gene
Silencing construct
The gene that causes cut apples to turn brown can be silenced
See for example Arctic Apple
© 2013 American Society of Plant Biologists
Transgrafting: Wild-type tissues grafted to transgenic tissues
Transgrafting: A non-transgenic shoot can be grafted onto transgenic root, so the food products and pollen don’t carry the transgene
Transgenes can include those conferring root-specific traits e.g., pathogen resistance or nutrient uptake
Or, the roots can produce mobile factors such as small RNAs or proteins that affect the shoot’s phenotype
See Liang, D., White, R.G. and Waterhouse, P.M. (2012). Gene silencing in Arabidopsis spreads from the root to the shoot, through a gating barrier, by template-dependent, nonvascular, cell-to-cell movement. Plant Physiol. 159: 984-1000; McGarry, R.C. and Kragler, F. (2013). Phloem-mobile signals affecting flowers: applications for crop breeding. Trends Plant Sci. 18: 198-206.
© 2013 American Society of Plant Biologists
Several new methods are available for precise genome engineering
From Jinek, M., Chylinski, K., Fonfara, I., Hauer, M., Doudna, J.A. and Charpentier, E. (2012). A programmable dual-RNA–guided DNA endonuclease in adaptive bacterial immunity. Science. 337: 816-821 with permission from AAAS; see also Bhaya, D., Davison, M. and Barrangou, R. (2011). CRISPR-Cas Systems in Bacteria and Archaea: Versatile small RNAs for adaptive defense and regulation. Annu. Rev. Genet. 45: 273-297.
Zinc-finger nucleases (ZFNs) and transcription activator-like effector nucleases (TALENs) are proteins that can produce double-strand DNA breaks that when repaired introduce site-specific mutations or insertions
The clustered regularly interspaced short palindromic repeats (CRISPR) / CRISPR-associated (Cas) system uses RNAs to target nucleases to specific sites; when repaired, site-specific mutations or insertions are introduced
© 2013 American Society of Plant Biologists
ZFNs and TALENs can be programmed to cleave specific sites
Reprinted from Gaj, T., Gersbach, C.A. and Barbas, C.F. (2013). ZFN, TALEN, and CRISPR/Cas-based methods for genome engineering. Trends Biotech. 31: 397-405 with permission from Elsevier.
ZFNs recognize 3-basepair sequences. A series of four linked ZFNs has a 12-bp binding specificity
TALENs recognize DNA via a series of 33 – 35 amino acid domains that each recognizes a single DNA basepair
The CRISPR / Cas system used RNA for targeting: cheaper than proteins
© 2013 American Society of Plant Biologists
Repair of the resulting double-strand breaks causes mutations / insertions
Reprinted by permission from Macmillan Publishers Ltd: from Nekrasov, V., Staskawicz, B., Weigel, D., Jones, J.D.G. and Kamoun, S. (2013). Targeted mutagenesis in the model plant Nicotiana benthamiana using Cas9 RNA-guided endonuclease. Nat Biotech. 31: 691-693.
Targetted cleavage
Template DNA
Repair in the presence of template DNA can lead to insertion of new sequences
Repair of the ds breaks can lead to small insertions and deletions
Wild-type sequence
Repair products showing deletions, insertions, and replacements
© 2013 American Society of Plant Biologists
These and other new approaches are becoming increasingly important
Reprinted by permission from Macmillan Publishers Ltd. From Lusser, M., Parisi, C., Plan, D. and Rodriguez-Cerezo, E. (2012). Deployment of new biotechnologies in plant breeding. Nat Biotech. 30: 231-239.
© 2013 American Society of Plant Biologists
Photo credit: IRRI
Future Challenges
© 2013 American Society of Plant Biologists
Breeders can use more than one technology to address a challenge
PROBLEM
SOLUTIONPublic –private
partnerships
MAS breeding
Gene pyramiding
Improved agronomic practices
GM technology
Genome-wide
association
© 2013 American Society of Plant Biologists
Breeding plants for β-carotene (pro-vitamin A) enrichment
Image sources: Petaholmes based on WHO data;
Vitamin A deficiency is a leading cause of blindness
The Plant Cell, October 2013 © 2013The American Society of Plant Biologists
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© 2013 American Society of Plant Biologists
Enhanced β-carotene content in food can prevent vitamin A
deficiency
Vitamin Aβ-carotene
β-carotene is converted to vitamin A in the human body
•Many staple foods are poor sources of β-carotene so many people do not get adequate vitamin A in their diet
© 2013 American Society of Plant Biologists
Synthesis, storage and breakdown all affect β-carotene content
Vitamin A
To increase beta-carotene levels in plants, you need more synthesis, more storage or less catabolism
GGPP
phytoene
lycopene
β-carotene
Synthesis
StorageCatabolism
or conversion
to other forms
Chromoplasts –organelles that store
carotenoids
Photo credit: University of Wisconsin
© 2013 American Society of Plant Biologists
β-carotene makes the rice
look golden
Vitamin A
Photo credit: Golden rice humanitarian board
β-carotene
The β-carotene enriched foods
shown here have been
produced using GM and non-GM
approaches
There is no inherently right or wrong way to enhance plant
nutritional quality
© 2013 American Society of Plant Biologists
Biofortified plants are improving nutrition for many
The non-profit organization HarvestPlus focuses on the development of biofortified crops for the developing world, including a provitamin A enriched sweet potato that is currently being grown by half a million families. Other biofortification projects are underway to increase levels of protein, iron, zinc, antioxidants and other beneficial components in food.
Sources: HarvestPlus; CIMMYT
© 2013 American Society of Plant Biologists
Breeding for drought tolerance
Water use efficiency is a complex trait that involves hundreds of genes
Photo credit: J.S. Quick, Bugwood.org
© 2013 American Society of Plant Biologists
Food production for one person for one day requires 3000 liters of water
Comprehensive Assessment of Water Management in Agriculture. 2007. Water for Food, Water for Life: A Comprehensive Assessment of Water Management in Agriculture. London: Earthscan, and Colombo: International Water Management Institute.
Global water withdrawal (km3/yr)
70% of water usage is for agriculture
© 2013 American Society of Plant Biologists
The incidence of major droughts is on the rise
China experienced a major drought in 2011
Extreme
Severe
Moderate
Image credits: USDA Foreign Agricultural Service; IRRI; NASA earth observatory, USDA Farm Service Agency
Major droughts and heat waves in China, Russia, the U.S. and Australia have impacted food production and raised prices
Russia experienced heat waves, drought and wildfires in 2010
© 2013 American Society of Plant Biologists
In 2011 seed companies released water-optimized corn
Both of these varieties were developed using modern molecular breeding methods
without the use of recombinant DNA
© 2013 American Society of Plant Biologists
Agricultural innovation in Africa –breeding crops for sub-Saharan
Africa
0-1
1-2
2-3
3-4
4-5
5-6
6-7
7-8
Fertility rateBirth rates are high and crop yields are low so many African countries are not currently self-sufficient
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© 2013 American Society of Plant Biologists
Many African countries experience a very high rate of undernourishment
Source: FAOSTAT
© 2013 American Society of Plant Biologists
The challenges to food production in Africa are immense
• Lack of infrastructure, especially irrigation and access to transportation networks
• High incidence of diseases • Lack of available fertilizers• Lack of education and support for
farmers• Lack of economic supports and market
stability• Agricultural subsidies in other countries
affect market value
© 2013 American Society of Plant Biologists
Maize is a staple crop in Africa but very sensitive to drought damage
Less than 10% of crop land in sub-Saharan Africa is irrigated, making agriculture production highly susceptible to drought
Photo credit: Anne Wangalachi/CIMMYT Map Source – FAO Aquastat 2005
Irrigation as percentage of cultivated area
© 2013 American Society of Plant Biologists
As a consequence of climate changes, droughts are expected to increase
Image credit: United Nations Economic Commission for Africa, 2008 Africa Review Report on Drought and Desertification
In some African countries, yields from rain-fed agriculture, which is important for the poorest farmers, could be reduced by up to 50% by 2020.
-(FAO 2010)
Water-efficient maize optimized for growth in sub-Saharan Africa (WEMA) has been developed through a
combination of breeding and GM methods
WEMA is being developed as a public-private partnership that includes international
and regional plant breeding institutes,
philanthropic groups and Monsanto
Photo credits: Anne Wangalachi/CIMMYT
Water Efficient Maize for Africa (WEMA) was developed through a public-private partnership
© 2013 American Society of Plant Biologists
Plant breeding can support African agriculture
African farmers need access to high yielding,
drought tolerant, disease resistant
plants. Most food is grown by small-
scale farmers with little mechanization. Cassava, cowpea and banana are
important crops and the focus of
intensive breeding programs.
Photos courtesy if IITA
© 2013 American Society of Plant Biologists
African governments are working together to support agriculture
“AGRA is a dynamic, African-led partnership working across the African continent to help millions of small-scale farmers and their families lift themselves out of poverty and hunger”. A major thrust of these efforts is to develop Africa’s human capacity through education, innovation and technology transfer.
Alliance for a Green Revolution in Africa
Source: AGRA
© 2013 American Society of Plant Biologists
In the next 50 years, we will have to produce as much food as we have yet produced in human history
Photo credit: © UNICEF/NYHQ1998-0891/Giacomo Pirozzi
© 2013 American Society of Plant Biologists
We have many paths to follow
PROBLEMS
SOLUTIONS
Increased water use efficiency
Improved nutrient content
Higher yields
Stress tolerance
Disease resistance
Improved fertilizer uptake
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© 2013 American Society of Plant Biologists
Breeding crops for a second green revolution
Many people are calling for a second green revolution, to develop plants that minimize environmental degradation while enhancing human health.
Advances in genetics make that possible.
Photo credit: IRRI
© 2013 American Society of Plant Biologists
What risk assessments are performed on GM crops?
Before release into the environment, GM crops are subject to risk-assessment and risk-management measures to evaluate:• Risks to human health (including toxicity and allergenicity)• Risks of evolution of resistance in target pathogens or pests• Risks to non-target organisms • Risks from movement of transgenes
© 2013 American Society of Plant Biologists
Will genes from GMOs contaminate wild populations?
John William Waterhouse: Pandora - 1896
When Pandora opened the forbidden box she released
evil into the world
Pollen can move DNA between plants. To minimize this possibility, GM crops have to be grown prescribed distances away from closely related plants. Technological methods to reduce this risk are being developed.
© 2013 American Society of Plant Biologists
Will anti-insecticidal genes harm unintended targets?
Image credit jons2
The evidence shows that the planting of GE crops has largely resulted in less adverse or equivalent effects on the farm environment compared with the conventional non-GE systems that GE crops replaced (National Academies 2010)
© 2013 American Society of Plant Biologists
Photo credit: CIMMYT.
Partnerships including national agricultural research institutions, non-government and community-
based organizations, regional research networks, and private
companies work together to develop seeds that are suited to
local conditions and are affordable for local farmers
> 45% of corn yields are often lost to insects
Will GMOs take away choice and exploit small farmers?
© 2013 American Society of Plant Biologists
Are GM crops safe to eat?
YES
Photo credit: Neil Palmer/ CIAT
All GM plants are subject to extensive testing and
regulatory oversight and no detrimental health effects
have been identified
Bt corn is less prone contamination by fungi which
produce toxins linked to cancer and birth defects
GM biofortification can ensure that all children get adequate levels of protein, vitamins and mineral nutrients.
GM is a safe and beneficial tool in the
quest to sustainably feed the growing population
© 2013 American Society of Plant Biologists
Scientists worldwide endorse GM as an important tool for breeding
“Both genetic improvementand better crop management
are vital and both should beresourced in parallel.” - 2009
“The ASPB believes strongly that, with continued responsible regulation and oversight, GE will bring many significant health and environmental benefits to the world and its people.” - 2006
© 2013 American Society of Plant Biologists
GMOs in the newsRothamsted’s response to activists
Aphid alarm pheromone (E)-β-farnesene
At Rothamsted Research in England, wheat plants were engineered to produce a chemical alarm signal that deters aphids
Source: Rothamsted Research; Sense about Science.
Activists threatened to destroy the experimental plants
Rothamsted scientists and supporters responded with public appeal, video, debate and petition with more than 6000 signatures
Ultimately, no plants were destroyed and the experiment was completed successfully
© 2013 American Society of Plant Biologists
GMOs in the newsAnti-GMO activist Lynas apologizes
In January 2013, Lynas apologized for his prior anti-GM activities, saying that he’d been “completely wrong to oppose GMOs”, and urging others to reconsider their opposition
Source: Mark Lynas
Mark Lynas is a well-known author and journalist
whose work has focused on the environmental
effects of climate change
“So my message to the anti-GM lobby … is this. You are entitled to your views. But you must know by now that they are not supported by science.”
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© 2013 American Society of Plant Biologists
GMOs in the newsFalse claims about GMO threats
Daily Mail: “Uncovered, the 'toxic' gene hiding in GM crops: Revelation throws new doubt over safety of foods”
-(21 Jan 2013)
Reuters: “Monsanto corn study in France finds tumors and organ damage in rats”
-(19 Sept 2012)
Natural News: “GMOs may be the new thalidomide”-(19 Sept 2012)
Daily Mail: “Cancer row over GM foods as study says it did THIS to rats... and can cause organ damage and early death in humans”
-(19 Sept 2012)
© 2013 American Society of Plant Biologists
Anti-GM activists intentionally propagate fear and misinformation
The more a technology is associated with “risk” in the media, the greater the public experiences it as “risky” – this is called “the social amplification of risk”
Sjoberg, L. (2004). Principles of risk perception applied to gene technology. EMBO Rep. 5: S47-S51. Frewer, L.J., Miles, S. and Marsh, R. (2002). The Media and Genetically Modified Foods: Evidence in Support of Social Amplification of Risk. Risk Analysis. 22: 701-711. Frewer, L. (2003). 10. Societal issues and public attitudes towards genetically modified foods. Trends in Food Science & Technology. 14: 319-332.
© 2013 American Society of Plant Biologists
Example: A retracted study blaming vaccines for autism had a big impact
A since-retracted report that the MMR vaccine causes autism led to a dramatic decrease in children being vaccinated, leading to disease outbreaks and an increase in mortality from these diseases
Plotkin, S., Gerber, J.S. and Offit, P.A. (2009). Vaccines and Autism: A Tale of Shifting Hypotheses. Clinical Infectious Diseases. 48: 456-461; Data from Health Protection Scheme UK; Cottrell, S. and Roberts, R.J. (2011). Measles outbreak in Europe. BMJ. 342: d3724.
1996 2000 2005 2010
1500
1000
500
50000
10000
20000
30000
40000
Reported cases of measles and mumpsin England (1996 – 2010)
94
90
86
82
Percentage of children who completed their MMR vaccination course in England (1996 – 2010)
Wakefield et al., 1998
© 2013 American Society of Plant Biologists
GMOs in the news “The Séralini affair”
Sept. 19, 2012 Gilles-Eric Séralini released a report that claimed rats fed GM corn died 2 – 3 times more than control rats
Sept.– Nov. 2012
The European Food Safety Authority
reports that Séralini’s study “does not meet acceptable scientific
standards”
European Food Safety Authority (2012) Final review of the Séralini et al. (2012a) publication on a 2-year rodent feeding study with glyphosate formulations and GM maize NK603 as published online on 19 September 2012 in Food and Chemical Toxicology EFSA Journal 2012;10(11):2986
Long term toxicity of a Roundup herbicide and a Roundup‐tolerantgenetically modified maizeSéralini, G.E., Clair,E., Mesnage, R., Gress, S., Defarge, N., Malatesta, M., Hennequin, D., and Spiroux de Vendômois, J.
Food Chem. Toxicol. 50: 4221–4231September 19, 2012
http://dx.doi.org/10.1016/j.fct.2012.08.005
Dozens of letters were written to the journal editor pointing out deficiencies in
the research
© 2013 American Society of Plant Biologists
• Sample sizes too small to be statistically significant
• “Cherry picking”: only some data presented
• Inappropriate test organism
• A dubious non-disclosure agreement that prevented reporters from consulting other experts about the research
The Séralini affair: Major flaws in the study include:
© 2013 American Society of Plant Biologists
Is a diet that contains Roundup or Roundup-tolerant corn toxic?
GM+R
GM
GM
R
Some rats had their diet supplemented with non-GM corn
Some rats had their diet supplemented with GM corn GM
Some rats had their diet supplemented with GM corn that had been sprayed with Roundup
Some rats had their diet supplemented with non-GM corn and had Roundup added to their water
To investigate toxicity, rats were given special diets for the duration of their lives (up to 680 days)
GM The proportion of corn in their diet is indicated by the size of the inner circle
© 2013 American Society of Plant Biologists
Experimental design: 1 control, 9 treatments
33% of food from non-GM corn
11%, 22% or 33% of food from GM corn
CO
NT
RO
L
11%, 22% or 33% of food from GM cornsprayed in the field
with Roundup
33% of food from non-GM corn, and water supplemented with 0.1 –5 x106 ppb Roundup
0.1 ppb
9 x105
ppb5 x106
ppb
GMGMGM GM+R
GM+R
GM+R
(The maximum tolerance for Roundup in corn is 1 x 103 ppb) EPA
© 2013 American Society of Plant Biologists
Experimental design: 10 male and 10 female rats in each group
♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀♂ ♀ ♂ ♀ ♂ ♀
0.1 ppb
9 x105
ppb5 x106
ppb
GMGMGM GM+R
GM+R
GM+R
© 2013 American Society of Plant Biologists
Results: Survival at 600 (♂) or 680 (♀) days
♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀♂ ♀ ♂ ♀ ♂ ♀
0.1 ppb
9 x105
ppb5 x106
ppb
GMGMGMGM+R
GM+R
GM+R
What do these results indicate about toxicity of GM corn or Roundup?
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The mortality data show no statistically significant treatment effects
Panchin, A.Y. (2013). Toxicity of roundup-tolerant genetically modified maize is not supported by statistical tests. Food Chem. Toxicol, 53: 475; Schorsch, F. (2013). Serious inadequacies regarding the pathology data presented in the paper by Séralini et al. (2012). Food Chem. Toxicol. 53: 465 – 466.
7 heads3 tails
5 heads5 tails
4 heads6 tails
3 heads7 tails
2 heads8 tails
4 heads6 tails
5 heads5 tails
Different values do not indicate real and reproducible differences
For example, how does the color of the hat affect the outcome of the coin toss? Obviously, it has no effect; the different numbers are a result of random variation
When Séralini’s data were analyzed, no significant differences were found between control and treatment groups
© 2013 American Society of Plant Biologists
“Cherry picking” means showing only a subset of data
Sanders, D., Kamoun, S., Williams, B. and Festing, M. (2013). Letter to the editor. Food and Chemical Toxicology. 53: 450-453; iPetitions.
It is unethical to exclude data that don’t fit the hypothesis
Hundreds of scientists have called on the authors to release their complete data set. The authors so far have refused to release the rest of the data
“All data cannot be shown in one report, and the most relevant are described here.”
-Séralini, 2012
Hiding conflicting data is also described as “sweeping it under the rug”
© 2013 American Society of Plant Biologists
7 heads3 tails
2 heads8 tails
If only some data are shown, the inherent variability of the study is hidden and differences are exaggerated
“Cherry picking” data skews data interpretation
“Wearing a blue hat causes 3.5 more
‘heads’ than wearing a pink hat”
Is this interpretation justified?
Of course not
© 2013 American Society of Plant Biologists
The authors show photos of tumors only on GM-fed rats
Look at the tumor on this rat that ate
GM corn!
Old Sprague-Dawley rats typically develop tumors no matter what they eat, and they did in the Séralini study, but showing a control rat with a tumor would dilute the association between GM and tumor formation
© 2013 American Society of Plant Biologists
As a consequence, many journalists passed on the press release as legitimate news
Séralini’s paper was released with a non-
disclosure agreement, meaning no experts could be consulted
Séralini manipulated the press embargo
Many articles are released to the press under an embargo, which allows the journalists time to research their articles
This usually involves getting an opinion from an independent
expert
© 2013 American Society of Plant Biologists
Scientists need to ensure that that debate, legislation and policy decisions are based on well-grounded
scientific evidence
“The majority of agricultural scientists, including myself, anticipate great benefits from biotechnology…
Genetic modification of crops is not some kind of witchcraft; rather, it is the progressive harnessing of the forces of nature to the benefit of feeding the human race…
We must be on guard against politically opportunistic, pseudo-scientists…
-Norman Borlaug, Nobel Laureate
Sanders, D., Kamoun, S., Williams, B. and Festing, M. (2013). Letter to the editor. Food and Chemical Toxicology. 53: 450-453; Borlaug, N.E. (2000). Ending World Hunger. The Promise of Biotechnology and the Threat of Antiscience Zealotry. Plant Physiol. 124: 487-490.
Most scientists support the continuing assessment of all new crops, including those developed by GM tools, and support informed public debate about their use