Professional Documents
Culture Documents
Proceedings
Oral Papers and Posters
Edited by Robert McIntosh
This document was prepared by the BGRI Secretariat, with financial support from the
Durable Rust Resistance in Wheat Project (http://wheatrust.cornell.edu).
The abstracts were edited by Dr. Robert McIntosh, Honorary Associate at the
Plant Breeding Institute of the University of Sydney.
www.globalrust.org www.globalrust.org
Proceedings
Oral Papers and Posters
Edited by Robert McIntosh
www.globalrust.org
Published by
Borlaug Global Rust Initiative
250 Emerson Hall
Cornell University
Ithaca, NY 14843
USA
htpp://www.globalrust.org
Email: bgri@cornell.edu
ISBN: 978-0-615-37399-7
The Borlaug Global Rust Initiative is grateful to the International Maize and Wheat Improvement Center
(CIMMYT) for serving as the kind host of the BGRI 2009 Technical Workshop, coordinating logistics, and assisting
with production of these proceedings.
The BGRI secretariat is located at Cornell University and is supported by the Bill & Melinda Gates Foundation.
Many other organizations and individuals contributed to the BGRI 2009 Technical Workshop, and we are very
grateful for their support.
Oral Papers
Table of contents
01. The Agricultural Research and Experimentation Board of the State of Sonora “A model for productive farmer-
researcher partnerships”
A. Gandara...........................................................................................................................................................................9
02. History and status of the wheat rusts
Robert A. McIntosh........................................................................................................................................................... 11
03. Using race survey outputs to protect wheat from rust
Robert F. Park, Thomas Fetch, Yue Jin, Mohinder Prashar, Zac Pretorius................................................................... 25
04. Using trap plot outputs to protect heat from rust
K Nazari, AH Yahyaoui, R Singh, T Fetch, Hodson, R Park............................................................................................. 33
05. The global cereal rust monitoring system
D.P. Hodson, K. Cressman, K. Nazari, R.F. Park, A. Yahyaoui . ...................................................................................... 35
06. Sequencing Ug99 and other stem rust races: progress and results*
Les J. Szabo , Christina Cuomo........................................................................................................................................ 47
07. Advances in host-pathogen molecular interactions: rust effectors as targets for recognition
Peter Dodds, Greg Lawrence, Rohit Mago, Michael Ayliffe, Narayana Upadhyaya,
Les Szabo, Robert Park, Jeff Ellis...................................................................................................................................... 49
08. Global stem rust surveillance in practice
Z.A. Pretorius, K. Nazari................................................................................................................................................... 55
09. Race nomenclature systems: Can we speak the same language?
T. Fetch Jr., Y. Jin, K. Nazari, R. Park, M. Prashar, Z. Pretorius....................................................................................... 61
10. Are rust pathogens under control in the Southern Cone of South America?
Silvia Germán, Marcia Chaves, Pablo Campos, Lidia de Viedma, Ricardo Madariaga............................................. 65
11. Recent experiences with global surveillance of wheat stripe rust
Mogens S. Hovmøller, Amor H. Yahyaoui, Annemarie F. Justesen............................................................................... 75
12. The development and application of near-isogenic lines for monitoring cereal rust pathogens
C.R. Wellings, R.P. Singh, A.H. Yahyaoui, K. Nazari, R.A McIntosh................................................................................ 77
13. Progress and prospects in discovery and use of novel sources of stem rust resistance
Y. Jin, M. Rouse, P.D. Olivera, B.J. Steffenson.................................................................................................................. 89
14. Cytogenetic manipulation to enhance the utility of alien resistance genes
M.O. Pumphrey, I.S. Dundas, S.S. Xu, Y. Jin, J.D. Faris, X. Cai, W.X. Liu, L.L. Qi, B. Friebe, B.S. Gill.............................. 93
15. Cloned rust resistance genes and gene based molecular markers in wheat: Current status and
future prospects.
Kota R, ES Lagudah, R Mago, H McFadden, PK Sambasivam, W Spielmeyer, L Tabe; B Keller, SG Krattinger,
LL Selter; S Herrera-Foesel, J Huerta-Espino, RP Singh; H Bariana, R Park, C Wellings; S Cloutier; Y Jin .............. 101
16. Molecular-genetic dissection of rice nonhost resistance to wheat stem rust
Michael Ayliffe, Yue Jin, Brian Steffenson, Zhensheng Kang, Shiping Wang, Hei Leung......................................... 103
17. Screening for stem rust resistance in East Africa
D. Singh*, B. Girma, P. Njau, R. Wanyera, A. Badebo, S. Bhavani, R.P. Singh,
J. Huerta-Espino, G. Woldeab and R. Ward.................................................................................................................. 111
18. Developing and optimizing markers for stem rust resistance in wheat
Long-Xi Yu, Zewdie Abate, James A Anderson UK Bansal, HS Bariana, Sridhar Bhavani, Jorge
Dubcovsky, Evans S Lagudah, Sixin Liu, PK Sambasivam, Ravi P Singh, Mark E Sorrells........................................ 117
iv 1. The Agricultural Research and Experimentation Board of the State of Sonora “A model for productive farmer-researcher partnerships” 9
Poster Abstracts
Table of contents
Theme 1: Rust Race Analysis & Surveillance
01. Genetic Diversity of Wheat Stem Rust Pathogen (Puccinia graminis f. sp. tritici)
Isolates from Ethiopia as Revealed by Microsatellites
B Admassu, W Friedt, F Ordon....................................................................................................................................... 229
02. Wheat Rusts Survey and Virulence of Puccinia graminis in Ethiopia
G Woldeab, B Girma, B Hundie, A Badebo, B Kassa, N Bacha, M Degefu, S Baze, F Handoro,
B Bitew, S Tmesgen, T Husein ........................................................................................................................................ 229
03. Puccinia striiformis f. sp. tritici Race Changes in the United States
X Chen, L Penman, A Wan, P Cheng.............................................................................................................................. 230
04. Population Structure of Wheat Disease Pathogens Causing Epiphytotics in Southern Russia
G Volkova, V Nadykta, L Anpilogova, L Kovalenco, E Sinyak...................................................................................... 230
05. Variability in Responses to Puccinia graminis pers. f. sp. tritici on Different Host Plants
ES Skolotneva, SN Lekomtseva...................................................................................................................................... 231
06. Occurrence of Wheat Rusts in Turkey During the 2008 Growing Season
Z Mert, F Dusunceli, K Akan, S Albustan, M Aydogdu, A Karakaya............................................................................ 232
07. Evolution of the Leaf Rust Pathogen on Durum Wheat in Northwestern Mexico
J Huerta-Espino, RP Singh, SA Herrera-Foessel, JB Pérez-López, P Figueroa-López ................................................ 232
08. Wheat Rusts in India - Pathogenic Changes
M Prashar, SC Bardwaj, SK Jain, YP Sharma, Jag Shoran........................................................................................... 233
09. Effective Rust Resistance Genes in Wheat under Moroccan Conditions
A Ramdani, M Jlibene, N Nsarellah, SM Udupa............................................................................................................ 233
10. Survey of Wheat Diseases in Morocco During the 2007-08 Growing Season
A Ramdani....................................................................................................................................................................... 234
11. Diverse Stem Rust Races Found In A Single Field In Washington, USA
M Rouse, S Stoxen, L Szabo, X Chen, Y Jin..................................................................................................................... 234
12. Status of Wheat Rusts in Uzbekistan
Z Ziyaev, SK Baboev, Kh Turakulov, A Morgounov, Z Khalikulov, RC Sharma.......................................................... 235
vi 1. The Agricultural Research and Experimentation Board of the State of Sonora “A model for productive farmer-researcher partnerships” 9
37. Stem Rust Resistance in Aegilops tauschii Germplasm
M Rouse, E Olsen, M Pumphrey, BJ Steffenson, Y Jin................................................................................................... 249
38. Resistance to Wheat Stem Rust in Spelt Wheat (Triticum aestivum ssp. spelta)
PD Olivera, S Gale, L Wanschura, M Rouse, Y Jin ............................................................................................................ 249
39. Progress and Prospects in Discovery and Use of Novel Sources of Stem Rust Resistance
Y Jin, M Rouse, PD Olivera, BJ Steffenson..................................................................................................................... 250
40. Wheat-Stripe Rust Interactions Involving ‘Moro’ Resistance
DA Gaudet, X Wang, B Puchalski, F Leggett, A Kuzyk, A Laroche............................................................................... 250
41. Effect of Silencing Gene Yr10 for Stripe Rust Resistance in Moro Wheat
W Liu, A Laroche, Z-S Kang, DA Gaudet........................................................................................................................ 251
42. Cloning And Characterization of Avr1 Gene from Puccinia triticina
A Pacheco, H Zhang, DB Hays.......................................................................................................................................... 251
viii 1. The Agricultural Research and Experimentation Board of the State of Sonora “A model for productive farmer-researcher partnerships” 9
1. The Agricultural Research and Objectives
Patronato’s main objective is to provide moral
Experimentation Board of the State and financial support to agricultural research activities
of Sonora “A model for productive conduced by INIFAP-CIMMYT in Sonora, as well as to
other research Institutions whose work is relevant to
farmer-researcher partnerships” farmers. Patronato also coordinates, jointly with CIANO
and CIMMYT, agricultural research and development
A. Gandara1 and technology transfer programs within the state. In
addition, it grants fellowships to researchers, promotes
scientific meetings, and provides services to farmers,
History of Agricultural Research in Sonora
such as the production of certified seed of new,
The development of agricultural research in the
improved varieties developed by INIFAP-CIMMYT, or
Yaqui Valley was marked by series of significant events:
other research institutions.
1. Agricultural research in Sonora began in 1910,
when the Richardson Company, concessionaire of
the Mexican Government for land surveying and
Organizational structure
Patronato’s governing body is made up of a
development in the Yaqui Valley, established field trials
General Assembly, a Board of Directors, and four
at “Campo Ontagota”.
Regional Technical Committees. The Assembly is the
2. In the early thirties, the “El Yaqui” Experiment
supreme authority, made up of 38 farmers’ unions and
Station was established at the initiative of Rodolfo Elias
associations - private and communal farmers - formally
Calles, Governor of the State of Sonora, who passed a
established in the state.
law assigning the land in block 611 of the Yaqui Valley to
this purpose. It was agreed that the experiment station
was to be operated by the Ministry of Agriculture, Financial resources
through the Agricultural Research Institute (Instituto de Patronato’s main funding source is the farmers’
Investigaciones Agricolas, IIA). voluntary contributions in the form of a quota based
3. In 1943, the Office of Special Studies (OSS), on crop production per hectare, which is collected
a Rockefeller Foundation/Ministry of Agriculture at planting time. Others sources of income include;
cooperative program, initiated research in Mexico with marketing registered seed to farmers’ unions and
the participation of American and Mexican scientists. associations, collaborative agreements with research
Dr. Borlaug arrived in Mexico in September 1944 and institutions, and private and public companies,
he his colleagues, members of the research team, donations by philanthropic foundations and public and
established the first field trials for selecting wheat private institutions interested in promoting agricultural
lines with improved rust resistance at the “El Yaqui” development in the region.
Experiment Station. Since 1996, all federal and state government
4. In 1955, with the initiative and support of financial support for agricultural research is managed by
the Yaqui Valley farmers, the land in block 910 was Fundacion Produce Sonora.
appropriated to establish the Northwest Agricultural The Laboratory for Phytosanitary Diagnosis of the
Research Center (CIANO), within the structure of OSS; PIEAES is another important funding source.
its mission was to solve problems limiting production in
the region. Importance of Patronato’s research
5. In 1960, OSS and IIA merged, to form the National participation
Agricultural Research Institute (INIA), today INIFAP. In northwestern Mexico, the most outstanding
6. In 1964, Yaqui Valley farmers decided to create achievement over 45 years of Patronato-CIANO-CIMMYT
their own organization to provide continuous and association has been the development of the model
systematic financial support for agricultural research. for a fruitful working partnership. This partnership has
This initiative led to the creation of The Agricultural strengthened research programs and has led to the
Research and Experimentation Board of the State of development of advanced technologies to solve real
Sonora (PIEAES), known simply as Patronato. problems and enhance agricultural productivity and
sustainability in the state of Sonora. An example for this
association was the detection of a widespread epidemic
of leaf rust on all commercially grown durum wheat
1
Patronato para la Investigación y Experimentación Agrícola del Estado de varieties in the Yaqui and Mayo Valleys in March 2001.
Sonora, Ciudad Obregón, México
10 The Agricultural Research and Experimentation Board of the State of Sonora “A model for productive farmer-researcher partnerships”
2. History and status rust was often more serious when wheat was grown
adjacent to hedgerows of barberry (Berberis vulgaris).
of the wheat rusts The next important step in understanding cereal rusts
came in the late 19th Century when Eriksson and co-
Robert A. McIntosh workers demonstrated the existence of formae speciales
as variants of a single fungal species varying in ability
Abstract to parasitize different host groups. For example, the
The rusts have been ongoing problems for wheat wheat-attacking form of P. graminis, P. graminis f. sp.
production probably since domestication of the crop tritici (PGT), the rye-attacking form P. g. f. sp. secalis (PGS),
about 8,000 years ago. Epidemics vary in size and and the oat-attacking form, P. g. f. sp. avenae (PGA). All of
frequency with host genotype and environment, wet these forms are capable of completing their life cycles
years being ‘rust’ years. Although partial control in on the alternate (sexual stage) host, but differ widely
modern agriculture was achieved with resistant varieties, in crossability. For example, PGT and PGS hybridize
conditions favoring epidemics were made worse with readily, but neither is crossable with PGA. By far the
the intensification of production and greater resistance most attention has been given to these forms, and
gene uniformity in the host. The current Ug99 incident relationships with, and among, other forms are not so
illustrates the situation of very widely adapted successful well known, but likely some are more closely related to
genotypes grown across huge areas in the presence PGA than to PGT or PGS.
of an ongoing threat from a recently emerged widely During the 1910s, Stakman and co-workers in the
virulent and obviously highly aggressive pathotype of USA showed that PGT was comprised of pathotypes
the stem rust pathogen. This paper addresses some (phenotypes, races, strains) with the ability to attack only
of the history of cereal rusts and reviews underlying certain combinations of Triticum genotypes, treating
principles of host pathogen genetics, some of which are diploids, tetraploids and hexaploids as a group. Initially
being neglected in the period of modern genetics. such pathotypes were given the status of genetically
fixed entities more or less like species. Subsequently,
Keywords pathotype variability was demonstrated in all cereal
Cereal rusts • host : pathogen genetics • resistance • rust pathogen species and the number of pathotypes
pathogenicity simply depended upon the number of host lines
(later, differential genes or gene combinations), and
Introduction the number of variations in infection type considered
The cereal rusts can be serious diseases of the small significant for any one host line, keeping in mind
grain winter cereals, including wheat, rye, triticale, oats that the genetic bases of the various resistances
and barley. The rusts of wheat attract the most attention were unknown. Thus, to some extent, the numbers
because wheat is one of the two most important food of pathotypes depended on the amount of effort
crops for mankind. While all the cereal rust pathogens that investigators wished to invest in defining them.
can be grouped as ‘rust pathogens’ because the different Nevertheless, pathotype identification and surveys
(Puccinia) species have many similarities, there are also became, and remain, routine activities in rust research
clear differences in terms of life cycles, alternate hosts, laboratories worldwide.
host range and genetics. Although de Bary described the various spore
The rusts and powdery mildews have been constant, stages of P. graminis, P. coronata and P. triticina (PT)
irregular curses for farmers throughout the history of and identified their alternate hosts in Berberis spp.,
agriculture, but it was not until the 19th century that it Rhamnus spp. and Thalictrum spp., respectively, it
was known that they were caused by fungi – the rust was Craigie in Canada who in 1927 demonstrated
pathogens being Basidiomycetes. The discovery by de heterothallism and sexual reproduction in P. graminis,
Bary that Puccinia graminis was a heteroecious species soon followed by Waterhouse (1929) in Australia with
explained the much earlier observations that stem similar work with P. triticina.
University of Sydney Plant Breeding Institute Cobbitty, Private Bag 11, Camden,
NSW 2570, Australia E-mail: bobm@camden.usyd.edu.au
In the 1940s and 1950s, host : pathogen genetics Table 1 Experimental designs used in host : pathogen
was put on a sound footing with the work on flax rust genetics (after Browder 1971)
(an autoecious rust system) by Flor who, from genetic
studies in both host and pathogen, showed that the Genetics of Use
expression of resistance in a host plant was specifically
Host Pathogen
dependent upon the presence of a corresponding
gene for avirulence in the pathogen. Any genetic or Postulation of
Unknown Unknown
environmental (e.g. temperature) factor that prevented CGPs
the presumed direct or indirect interaction of the gene
Known
products of the corresponding gene pairs resulted Unknown Pathotype analysis
(fixed)
in a compatible disease response. I visualize a single
corresponding gene pair (CGP) (Fig. 1) as the first law Known Host gene
Unknown
of host : pathogen genetics. The interactions shown in (fixed) postulation
Fig. 1 assume homozygosity (or complete dominance)
Physiologic and
of both host resistance and pathogen avirulence alleles Known Known
environmental
(cereal rust pathogens on cereals are dikaryotic, but (fixed) (fixed)
studies
behave as diploids). Differential phenotypic responses
(infection types) can occur with incomplete dominance.
For example, Samborski (1963) showed that a rare Centers of origin and global spread of
intermediate response produced by a P. triticina isolate wheat rust pathogens
on line Transfer with Lr9 was due to heterozygosity of
the corresponding P9 locus in the pathogen. There is general agreement that the centers of
If the above model is extended to a second CGP, the origin of pathogens are usually the same as the centers
matrix rapidly becomes very complicated (Fig. 2) even of origin of the host species (Karnal bunt may be an
when assuming homozygosity. However, this is the basis exception). In the case of heteroecious pathogens
for the second law of host : pathogen genetics which is those areas should overlap with the alternate hosts. The
about the interaction of CGPs. When more than a single pathogens (or at least components of their populations)
CGP is involved, the outcome is a phenotype that is then move from those areas along with the host
as incompatible, or more incompatible, than the most species. To much of Europe, northern Africa and Asia,
incompatible of the individual participating CGPs. The urediniospores could have been wind-borne from
use of these laws leads to the four basic experimental the Fertile Crescent. To more distal areas of southern
designs in host : pathogen genetics outlined by Browder Africa, the Americas and Australia, pathogens may have
(1971) (Table 1). One of the outcomes of Flor’s work been wind-borne (at very low frequencies because
PATHOGEN
VARIANT CLONE 1 CLONE 2 CLONE 3 CLONE 4
DESCRIPTION A VIRULENT A VIRULENT VIRULENT VIRULENT
PHENOTYPE
H
O
S
T
the populations are quite different), but the original generally assumed that there is no exchange of inoculum
founding populations were more likely to have been between northern and eastern Africa and southern
transported in cereal hay (along with weed seed) used Africa, a recent report (Visser et al. 2009) indicates that
to feed animals. It seems highly unlikely that suitably pathotypes closely related to Ug99 are not only present
adapted or adaptable pathogen populations would in South Africa, but are evolving in a parallel manner, and
have been present on grasses (or alternate hosts) prior are current predominant pathotypes in that country.
to colonization. Moreover, since the diseases usually The introductions of P. striiformis to eastern
appeared soon after colonization, the target areas of Australia in 1979 and 1998 (barley grass stripe rust), to
cultivated cereals would have been far too small to Western Australia in 2002 and on to eastern Australia in
intercept what would have been extremely low levels of 2003, South Africa in 1996, and to the USA in 2000 are
wind-borne viable spores. likely examples of man-borne introductions. The 1979
The initial occurrences of stripe rust in South America introduction to Australia was probably from western
(Chile) and North America were suggested to have arisen Europe (based on pathotype identity), the barley grass
from native P. striiformis populations on indigenous grass rust and Western Australian introductions were likely
communities. The South American pathotypes were from the USA (based on visitor frequencies), and the
very similar to European forms, but no exotic source was introduction to South Africa was probably from Turkey or
suggested for North American pathotypes. neighboring areas (based on pathotype similarity). No
In contrast to North America, Australian cereal rust origin has been proposed for the recent new group in
researchers were always cognizant of their geographical the USA, but eastern Asia may be a possibility. The lack
isolation, and proposed the likelihood of introduced of commonality between differentials used in Europe,
inoculum for a number of significant pathotype changes. Australia and South Africa with those used in India, China
For example, Watson and De Sousa (1983) presented and in the USA confounds the problem of pathotype
both pathotype and meteorological evidence for the comparisons, especially when we know that many of
possibility of wind-borne spores from southern Africa, the genotypes used in each area carry combinations of
but origins of other instances of putative introductions (seedling) resistance genes rather than single genes.
of PGT and PT were never identified. While it has been Whereas many will argue the need for molecular markers
1
International Center For Agricultural Research in the Dry Areas, Tel Hadya,
Aleppo, Syria; 2International Maize and Wheat Improvement Center, Mexico, D.F.
Mexico; 3Agriculture and Agri-Food Canada, Cereal Research Centre, Winnipeg,
MN R3T 2M9, Canada; 4Plant Breeding Institute Cobbitty, University of Sydney,
Camden, NSW 2570, Australia.
E-mail: k.nazari@cgiar.org
Actual confirmed observations of Ug99, obtained wise expansion of the Ug99 range had continued and
after the first postulated migration routes were the pathogen had now penetrated the major wheat
produced have been supportive, and not contradictory, producing areas of the Middle East. Fig. 2 shows current
of the initial GIS-based predictions. In 2006, reports of known locations.
stem rust were received from a site close to New Halfa Recorded known locations of Ug99 over time
in eastern Sudan, and subsequently from at least two illustrate the mobility of the pathogen and highlight the
sites in Western Yemen in the same year. Analysis of rust possibilities for long-distance air-borne transmission.
samples collected from these locations subsequently Obviously, understanding, and if possible predicting,
confirmed the presence of Ug99 (race TTKSK). The likely air-borne movements is going to be a critical
observed range of expansion of Ug99 indicated step- component of any cereal rust monitoring system.
wise movements following regional winds. Crossing This is a non-trivial task, as predicting air-borne
of the Red Sea into Yemen was considered particularly particle movement is a challenge due to the inherent
significant as several lines of evidence indicated that complexity and variability of the underlying system.
this might prove to be a gateway for onward movement Hence it must be borne in mind that there will always
into important wheat areas of the Middle East and Asia. be considerable uncertainties associated with any
In 2008, confirmatory race analysis data were obtained such pathogen prediction studies. These uncertainties
from stem rust samples that had been collected at two also have implications regarding any assumptions of
sites in Iran – Borujerd and Hamadan – at the end (July) “fixed repeatable pathways”; such hypotheses must
of the 2007 wheat season (Nazari et al. 2008). Step- be approached with the utmost caution as significant
deviations may well occur. It is not a given that the new
The two confirmed Ug99 (race TTKSK) sites in data (i.e. using wind data models to find potential
western Iran - Borujerd and Hamadan – as reported by sources for known final destinations) also highlighted
Nazari et al. (2008) were not exactly on the trajectory these same areas as potential zones of origin for on-
paths predicted by the HYSPLIT model, but were in ward spore movements that terminated in Borujerd and
close proximity. However further analysis, using daily Hamadan. Finally, unconfirmed but credible reports of
rainfall estimates in combination with the trajectory Ug99 were obtained from a site near Shiraz in Iran earlier
models, has provided circumstantial evidence of an in the wheat season of 2007. This location is coincident
intermediate staging location either in southern Iran or with one of the postulated intermediate staging zones
southern Iraq. Rainfall events are important as they are derived from the combined model outputs.
the principle means by which spore deposition occurs All of these foundation activities, i.e. the
(Rowell and Romig 1966). On two occasions during the compilation of key datasets, the mapping of known
period Dec. 2006 to Jan. 2007 significant rainfall events Ug99 sites, and the use of wind and rain models to
were exactly coincident in time and space with wind understand potential movements, provide a basis upon
trajectories originating from confirmed Ug99 sites in which a fully operational monitoring system can be
Yemen. It is feasible that these rainfall events had the developed. Obviously these activities do not constitute
potential to deposit rust spores in either southern Iraq a fully functional monitoring system and several
or around the gulf coast of Iran. Backwards trajectory additional key components are required.
No. Surveys
This means there is a fundamental requirement for
geo-referenced data inputs to drive the entire system.
To ensure a global overview there is also a need for a
centralized database from which integrated analysis can
be undertaken. Finally, for the system to have utility it
must deliver timely and targeted information to a range
of end-users, i.e. scientists, decision-makers and policy-
makers. A simplified schematic overview of the cereal
rust monitoring system is given in Fig. 4.
2007 2008
1
USDA ARS Cereal Disease Laboratory, St. Paul, MN; 2Broad Institute,
Massachusetts Institute of Technology and Harvard University, Cambridge, MA
Avirulence locus Product size (aa) Cys rich # gene family members Cognate R genes
AvrL567 150 no 12 L5, L6, L7
AvrM 260-384 no 6 M
AvrP4 95 yes 3 P4
AvrP123 117 yes >2 P, P1, P2, P3
1
Department of Plant Sciences, University of the Free State, Bloemfontein 9300,
South Africa; 2ICARDA, PO Box 5466, Aleppo, Syrian Arab Republic.
E-mail: pretorza.sci@ufs.ac.za
Ethiopia (DZ)* Ethiopian Institute of Agricultural Research, Debre Zeit Dr Ayele Badebo Huluka
South Africa (Bfn)* University of the Free State, Bloemfontein Prof ZA Pretorius
South Africa (Bhm)* ARC-Small Grain Institute, Bethlehem Dr Tarekegn Geleta Terefe
100
80
60
40 Personal
Institute
20
Paraguay
Canada
Uruguay
Australia
Brazil
Mexico
Ethiopia (DZ)
S Africa (Bfn)
Argentina
Armenia
Chile
China
Ethiopia (A)
Georgia
India
Iran
Kenya
Morocco
Pakistan
Sudan
Syria
S Africa (Bhm)
USA
Five countries, viz. Australia (PBI Cobbitty, University information being obtained from these collections, the
of Sydney), Canada (Agriculture and Agri-Food, mortality of East African wheat stem rust samples sent to
Winnipeg), India (DWR, Flowerdale, Shimla), Mexico the USA was unacceptably high (Y. Jin pers comm).
(CIMMYT) and USA (USDA-ARS CDL, St. Paul) have The numbers of entries in differential sets ranged
institutional histories of more than 60 years in stem rust from eight (Iran) to 50 (CIMMYT). Cereal rust laboratories
race analysis. Personal experience ranged from 0 to 35 in Argentina, Brazil, Canada, China, Mexico, Pakistan,
years (Fig. 1). Some respondents mentioned that they South Africa, Uruguay and USA use, or plan to use, the
lack stem rust experience, but have been trained in leaf or North American (NA) (Roelfs and Martens 1988; Roelfs
stripe rust analysis. Respondents from Pakistan and South et al. 1993; Jin et al. 2008) stem rust set. Other countries
Africa (ARC Bethlehem) indicated that their institutions such as Armenia, Ethiopia (Ambo), Morocco and Syria
have a significant history of stem rust work, but that they use the ICARDA set which includes the NA differential
personally lacked experience in this field. Considering lines. Not all countries using the NA system have the
both personal and institutional experience, countries same number of entries or the same genotype per Sr
such as Australia, Canada, China, Georgia, Mexico, South gene and it is possible that the expression of resistance
Africa and USA have had good continuity in stem rust genes in dissimilar backgrounds may influence race
race surveys. designation. Of some concern is the fact that more than
In terms of the number of stem rust samples one version of the NA race code exists in the literature.
processed during 2006 to 2008, the disease was most Jin et al. (2008) proposed an official fifth set (Sr24, 31, 38
prevalent in Canada (684 isolates), Georgia (422), and McNair 701) to the NA differential series (Roelfs et
USA (273) South Africa (218) and Australia (200) (Fig. al. 1993). However, a fifth set consisting of Sr7a, 8b, 13
2). Although no response was received from Yemen, and McNair, previously added by Canadian rust workers
32 isolates of P. graminis f. sp. tritici were pathotyped (Fetch and Dunsmore 2004), is still being used (Admassu
at the CDL in St. Paul and Agriculture and Agri- et al. 2009). Papers using the NA nomenclature thus
Food Canada in Winnipeg. In a study completed in have to be clear on which Sr gene set was used.
Germany, 152 Ethiopian wheat stem rust isolates were Australia and Georgia use the Stakman system plus
characterized (Admassu et al. 2009). Similarly, due to a additional tester lines, which differ between the two
lack of infrastructure, race analyses of stem rust samples countries, and Iran and India have their own differential
collected in Kenya (99 isolates), Ethiopia (56) and Sudan sets. However, data obtained from the Georgian and
(1) were done in St. Paul and Winnipeg. Despite valuable Indian sets can be converted to the NA code. The ARC
800
700
600
500 2008
400 2007
300 2006
200
100
0
Ethiopia (NA)
Paraguay
Kenya (NA)
Sudan (NA)
Canada
Uruguay
Australia
Brazil
Mexico
S Africa (Bfn)
Argentina
Armenia
Chile
China
Ethiopia (A)
Georgia
India
Iran
Morocco
Pakistan
Syria
S Africa (Bhm)
USA
Ethiopia (Germ)
Small Grain Institute in South Africa allocates a ‘2SA’ disease in China, stem rust has been controlled through
race number, but the current differential set also allows resistance breeding and now occurs mainly on spring
conversion to a NA code. wheat in some regions. In a study conducted in
Most respondents indicated that they reselect or Germany, Admassu et al. (2009) reported 22 stem rust
increase their differential entries annually and 64% races from 152 collections made in Ethiopia in 2006. No
indicated that they often observe mixtures within virulence for Sr24 was detected but TTKS was commonly
differentials. Even laboratories starting with seed found. At Ambo problems were encountered with poor
multiplication of ‘new’ stocks may be at risk of impurities seed germination and mixed differentials resulting in
because most likely their lines were obtained from incomplete data sets. However, virulence for Sr31 was
established institutions where these entries have been common. The respondent from Georgia mentioned that
increased many times. An international effort to purify barberry occurs in certain regions, but did not provide
differential lines and establishment of a single stock for information indicating its potential contribution to racial
each entry from which subsets can be distributed, will diversity in stem rust.
greatly enhance the accuracy of global data. In Australia, stem rust has been rare in commercial
Respondents from Australia, Argentina, Brazil, wheat in recent years. Most samples come from
Canada, China, India, Iran, Mexico, Morocco, Pakistan, experimental plots and only three races were detected
South Africa, Uruguay and USA mentioned that they in the most recent survey. This low incidence is
have access to air-conditioned greenhouses, an isolated attributed to resistance breeding using Sr genes which
facility for raising seedlings uncontaminated by rust remain effective, individually, or in combinations. The
spores, efficient inoculation and incubation facilities, and South American countries reported that, in general, stem
the capacity to sub-culture stem rust isolates. In general rust has not been a problem for many years. Several
these facilities are sufficient for conducting reliable institutes are currently increasing differential lines and
cereal rust research. Furthermore, efforts are currently should start with race analysis in 2010. In Argentina,
underway in Kenya and Ethiopia to improve their rust representative isolates from 2000 onwards are available
research facilities. Together with trained personnel and for testing once the system has been optimized.
appropriate seed stocks and rust isolates, stem rust race In North America, one race (QFCSC) has dominated
data, in addition to those from established laboratories, the Great Plains, Eastern USA and Canada in recent years.
should be available from several countries in future. Unique races were found in the Northwest US, most
Based on recent survey data, most diversity in stem likely from a sexual population (Y. Jin pers. comm.). In
rust populations appears to occur in China, Ethiopia Mexico, diversity in wheat stem rust is uncommon and
and Georgia. Despite being a historically important only one race has lately been identified.
Table 2 Differential genotypes recommended for use in pathotyping Puccinia graminis f. sp. tritici
1
INIA La Estanzuela, CC 39173, Colonia, CP 70000, Uruguay; 2EMBRAPA Trigo,
Passo Fundo, Brazil; 3INTA Bordenave, Bordenave, Argentina; 4MAG DIA CRIA,
Encarnación, Paraguay; 5INIA CRI Quilamapu, Chillán, Chile.
E-mail: sgerman@inia.org.uy
66 Are rust pathogens under control in the Southern Cone of South America?
number of races are generally present every year. first race virulent on BRS 194, widely grown in Brazil from
The prevalent races change dramatically over time, in 1988 to 1994 (Chaves et al. 2009). MDT-10,20 and related
accordance with the area sown to cultivars susceptible race MFT-10,20 (Brazilian designation B55, Chaves 2007)
to different pathotypes. After short periods of time, with have been prevalent in Brazil since 2005 and since first
few exceptions, resistance of new cultivars is overcome detected in Argentina and Uruguay in 2007. These races
by new virulent races of the pathogen. are also present in Paraguay. Related races MDT-10,20 and
Leaf rust surveys and race identifications are MFT-10,20 are virulent on a wide range of commercial
performed annually in Argentina, Brazil and Uruguay. cultivars, and have caused severe epidemics on the most
Samples from Chile and Paraguay are also analyzed for popular cultivars in Brazil and Paraguay, as well as other
race identification. During 2004-2007 races MCP (Long popular cultivars from Argentina, Brazil and Uruguay.
and Kolmer 1989) with additional virulence on Lr10 MCT-10/MHT-10 was frequently found in Argentina during
(MCP-10), MDR with additional virulence on Lr10 and 2004-2006. Race MCD-10,20 (previously prevalent across
Lr20 (MDR-10,20) and MFP, also virulent on Lr20 (MFP-20), the region) and MCP-10,20 were also identified in samples
were present in high frequencies in Argentina (Campos from Chile and Paraguay. Race MCD-10,20, was isolated
2008) and Uruguay (Table 1). MCP-10 was associated from samples collected during 2007 and 2008 in Chile
with severe epidemics on Klein Don Enrique (Lr26 and and analyzed in Uruguay and the USDA-Cereal Disease
additional resistance +; Antonelli 2003) in Argentina and Laboratory (J. Kolmer pers comm). Some races isolated
Uruguay, MDR-10,20 was associated with epidemics on from durum wheats were different from races previously
INIA Torcaza (Lr10, Lr24 +; Germán et al. 2005; Demichellis identified in the region (J. Kolmer, pers. comm.). The most
et al. 2008) and INIA Churrinche (Lr10, Lr24; Demichelis significant changes in the leaf rust population during
et al. 2008) in Uruguay, and MFP-20 was associated with 1996-2003 affecting 10 cultivars represented an estimated
epidemics on INIA Tero (Lr17, Lr24; Germán et al. 2005) loss of US$172 million to Southern Cone wheat farmers
in Uruguay. MFP-20 (Brazilian designation B56) was the (Germán et al. 2004).
Table 1 Race frequencies of Puccinia triticina isolates collected in Argentina, Brazil and Uruguay during 2004-2007
MCP-10 2000 46.0 16.9 29.0 3.0 2000 28.0 15.4 1.5
MCT-10 B34 4.0 17.5 25.0 2.0 1989 2.0 3.7 3.1 2.9 1992 1.7 2.2
MDP-10,20 B58 2005 1.9 7.0 9.0 2007 2.5 2004 4.0 10.9 18.7 19.0
MDR-10,20 2004 1.0 16.9 8.0 8.0 2003 17.0 32.0 20.1 3.3
MFP-20 B56 2004 3.9 11.0 10.0 2005 0.7 5.2 2.5 2005 1.7 28.4 4.1
MDT-10,20 B55 2007 21.0 2005 26.4 66.5 73.0 2007 24.0
No isolates 96 138 197 299 191 244 100 175 134 121
68 Are rust pathogens under control in the Southern Cone of South America?
Field resistance is the most important criterion rust susceptible cultivars were grown on 35% and
for selection in populations derived from crosses with 15% of the wheat areas in Argentina and Uruguay,
adapted materials. Molecular markers for Lr34 (Lagudah respectively, during 2007, and this figure increased to
et al. 2006), used in Argentina to confirm the presence almost 30% in Uruguay during 2008. Except for a few
of this gene in advanced lines, will also be used in Brazil Brazilian cultivars, which are also grown in Argentina
and Uruguay. Trp1 and Trp2 markers for the APR present and Uruguay, there is no available information on the
in Toropí (S. Brammer et al. unpublished) are under stem rust reactions of cultivars used in Brazil, Chile and
validation and will be used to screen breeding lines Paraguay to races present in the region. The increasing
in Brazil. In contrast with APR sources from CIMMYT, areas of susceptible cultivars may result in inoculum
Toropí has intermediate resistance to Fusarium head increase and development of stem rust epidemics
blight (caused by Fusarium spp.), which can be severe in In Argentina, the frequent presence of natural
Argentina, Brazil, Paraguay and Uruguay when favorable stem rust infections in summer breeding nurseries at
weather conditions occur. Molecular markers for minor Balcarce allows selection and characterization of stem
genes conferring APR to leaf rust will be particularly rust reactions to local races. In Uruguay, an artificially
useful to screen populations or lines from crosses inoculated late sown nursery is used to characterize
involving one parent with intermediate or high resistance the stem rust field reactions of new breeding lines
to leaf rust conferred by major genes. Homogeneous and commercial cultivars. Information of seedling
adapted advanced lines have been selected from crosses reactions to local races is also available in Argentina
involving sources of APR to leaf rust and adapted high and Uruguay. Selection for resistance is performed in
yielding materials and are being used to start a second breeding materials where segregation for resistance
cycle of selection for APR to leaf rust. Selected lines will is present, but no efforts are directed to identify and
eventually be released as commercial cultivars. systematically use sources of resistance to the local
Although both the economic importance of stripe pathogen population.
rust and pathogen variation have decreased in the The most important genes conferring resistance
last decade, breeding for resistance has remained a in the regional germplasm are Sr24 and Sr31
long term objective of the INIA-Chile wheat breeding (Campos and López 2008; Germán et al. 2007). A
program. Since susceptible materials are annually high proportion of the Uruguayan, Argentinean and
completely destroyed by the disease, indicating that Brazilian wheat germplasm carries the associated
favorable environment and compatible pathogen/ genes Lr24 and Lr26 (Campos 2008; Germán et al.
host combinations prevail, the use of sources of 2007). Only limited information of other Sr genes
durable resistance to stripe rust is a priority. Due to present in modern germplasm is available.
the association between APR to leaf rust and APR to
stripe rust (Singh 1992; Singh et al. 2003), selection for Facing the threat of Ug99 and derived races
resistance to leaf rust will indirectly increase the level of Widespread severe stem rust epidemics
resistance to stripe rust in the germplasm developed in associated with new pathogen races virulent to most
eastern countries of the Southern Cone. commercially grown cultivars have occurred in the
Stem rust resistance was a major wheat breeding Southern Cone (Germán et al. 2007). Likewise, since
objective when the disease was prevalent. The absence most varieties used by farmers are susceptible in Kenya,
of the disease for many years not only decreased the region is facing a potential threat of significant
opportunities for selection, but also led to changed epidemics if Ug99 or derived races were accidentally
priorities for breeding programs. As a result, susceptible introduced (Campos and López 2008; Germán and
cultivars were released in Argentina and Uruguay. Stem Verges 2005; Germán et al. 2007).
Table 2 Numbers and percentages of stem rust resistant and moderately resistant lines identified in Kenya, 2005-2008
70 Are rust pathogens under control in the Southern Cone of South America?
International testing in Kenya and Ethiopia made Homogeneous adapted F6 lines combining
possible by the Borlaug Global Rust Initiative has allowed resistance to Ug99 and derived races and resistance
highly relevant testing and selection for resistance to leaf rust were selected in Uruguay (Table 4). LE2304
in South American breeding materials. An increased is a sister line of INIA Tero, and likely carries the same
number of wheat materials from the region have been seedling resistance to stem rust. Parula and Genaro*3/
tested at the Kenyan Agricultural Research Institute- Parula posses APR to leaf rust and stripe rust, and Parula
Njoro Plant Breeding Resarch Center since 2005 (Table also has high levels of APR to stem rust. Lines R07 F5-
2). The reduced proportion of resistant materials in 2006 3027 and R07 F5-3037 derived from the cross Genaro*3/
and 2007 relative to 2005 was due to the new variant Parula//LE 2252 were more resistant to stem rust in
virulent for Sr24 (Jin et al. 2008). However, a number of Kenya than Genaro*3/Parula.
cultivars and lines continued to be resistant (Table 3). The Sources of APR to stem rust identified in east
genetic bases of the resistances are unknown. Africa were distributed in the International Stem Rust
Several sources of resistance to stem rust identified Resistance Screening Nurseries (ISRRSN). Entries that
in North East Africa, including APR sources such as do not carry Sr31 or Sr24, but were resistant in Kenya,
Pavon 76 and Parula (Singh et al. 2008) have been used were also resistant or moderately resistant when tested
in crosses with locally adapted cultivars in Argentina, in Uruguay. These sources of resistance are probably
Brazil, Paraguay and Uruguay. Other materials with APR also effective against other races present in the region.
to leaf rust distributed through collaborative research Therefore research efforts to introduce stem rust resistance
projects among programs in the Southern Cone were to Ug99 and derived races into the regional germplasm will
also resistant in Kenya (e.g. Suz6/Opata (Table 4), BR23// also increase the level of resistance to current local races of
CEP19/PF85490). the pathogen.
Table 4 Field stem rust (Kenya) and leaf rust reactions (Uruguay) of F6 Uruguayan breeding lines
72 Are rust pathogens under control in the Southern Cone of South America?
Hacke, E. 2007 Mejoramiento genético para el control Rajaram S, Campos A (1974) Epidemiology of wheat
de las principales enfermedades que afectan al trigo rusts in the western Hemisphere. CIMMYT, México DF.
candeal. In: Acevedo E, Silva P (eds) Trigo candeal: Research Bulletin no. 27
calidad, mercado, zonas de cultivo. Universidad de Singer P (2008) Monitoramento da sensibilidade de raças
Chile, [s.l.]. Serie Ciencias Agronómicas no 12 de Puccinia triticina a fungicidas do grupo dos tiazóis
Jin Y, Pretorius ZA, Singh RP, Fetch T Jr (2008) Detection e estrobilurinas. In: II Reunião da Comissão Brasileira
of virulence to resistance gene Sr24 within race TTKS de Pesquisa de Trigo e Triticale, 2008, Passo Fundo, RS,
of Puccinia graminis f. sp. tritici. Plant Dis 92:923-926 Brasil. Painel técnico: fungicidas triazóis no controle da
Lagudah ES, McFadden H, Singh RP, Huerta Espino J ferrugem da folha do trigo. CBPTT, Passo Fundo. www.
et al (2006) Molecular genetic characterization of cnpt.embrapa.br/rcbptt/2rcbptt/index.htm. Accessed
the Lr34/Yr18 slow rusting resistance gene region in 9 Mar 2009
wheat. Theor Appl Genet 114:21-30 Singh RP (1992) Genetic association of leaf rust resistance
Long DL, Kolmer JA (1989) A North American system gene Lr34 with adult plant resistance to stripe rust in
of nomenclature for Puccinia recondita f.sp. tritici. bread wheat. Phytopathology 82:835-838
Phytopathology 79:525-529 Singh RP, Huerta-Espino J, William M (2003) Resistencia
Maciel JLN, Chaves MS (2008) Desempenho do durable a roya de la hoja y roya amarilla del trigo:
principio ativo tebuconazole no controle da genética y mejoramiento en el CIMMYT. In: Kohli MM,
ferrugem da folha do trigo. In: II Reunião da Díaz de Ackermann M, Castro M (eds) Estrategias
Comissão Brasileira de Pesquisa de Trigo e Triticale, y metodologías utilizadas en el mejoramiento de
2008, Passo Fundo, RS, Brasil. Painel técnico: trigo: un enfoque multidisciplinario, 8-11 oct 2001,
fungicidas triazóis no controle da ferrugem da folha La Estanzuela, Colonia, Uruguay. CIMMYT-INIA,
do trigo. CBPTT, Passo Fundo. www.cnpt.embrapa. Montevideo, pp109-118
br/rcbptt/2rcbptt/index.htm. Accessed 9 Mar 2009 Singh RP, Hodson DP, Huerta-Espino J, Jin Y et al (2008)
Madariaga R, Mellado M, Becerra V (2004) Significance Will stem rust destroy the world´s wheat crop? Adv
of wheat yellow rust (Yr) genes in Chile. In: Proc 11th Agron 98:271-309
Int Cereal Rusts and Powdery Mildews Conf Abstr Singh RP, Germán S, Huerta-Espino J (2009) Genetic
John Innes Center, Norwich, England, UK ppA2.38 control of wheat rusts in Latin America: current
Madariaga R, Matus I (2008) Razas de la roya de la hoja status and future challenge. In: Proc 15º Congreso
del trigo en el ciclo 2007 – 2008 en Chile: abstract. Latinoamericano de Fitopatología, 18º Congreso
In: 59o Congreso Agronómico de Chile, 07 – 10 Chileno de Fitopatología, 2009, Santiago, Chile.
Octubre 2008, La Serena, Sociedad Agronómica de Fitopatología: libro de resúmenes. ALF, Santiago. pp.
Chile. pp. 46 39-42
Mellado M (2007) El trigo en Chile. INIA, Centro Stakman EC, Stewart DM, Loegering WQ (1962)
Regional de Investigación Quilamapu, Chillán, Chile. Identification of physiologic races of Puccinia graminis
Colección Libros INIA no. 21 var. tritici. US Dept Agric ARS, Washington DC. E 6/7
Peterson RF, Campbell AB, Hannah AE (1948) A Zoldan SM, Barcellos AL, Sousa CNA de (2000) Detecção
diagrammatic scale for estimating rust intensity on do gene Lr13 de resistência à ferrugem da folha em
leaves and stems of cereals. Can J Res 26 (Sec C): plântulas de trigo. Fitopatol Bras 25:512-516
496-500 Zoldan SM, Barcellos AL (2002) Postulation of genes (Lr)
for resistance to leaf rust in Brazilian wheat cultivars.
Fitopatol Bras 27:508-516
1
University of Aarhus, Faculty of Agricultural Sciences, Department of
Integrated Pest Management, Flakkebjerg, 4200 Slagelse, Denmark;
2
International Center for Agricultural Research in the Dry Areas, PO Box 5466,
Aleppo, Syria.
Email: mogens.hovmoller@agrsci.dk
78 The development and application of near-isogenic lines for monitoring cereal rust pathogens
Table 1 Avocet S Near-isogenic Lines developed and released in three cohort
Gene Source NILs97 NILs 98 NILs 99
Yr1 Chinese 166 Yr1/6* Avocet S Yr1/6* Avocet S Yr1/6* Avocet S
Yr5 T. spelta album Yr5/6* Avocet S Yr5/6* Avocet S Yr5/6* Avocet S
Yr6 Oxley Yr6/6* Avocet S Yr6/6* Avocet S
Yr7 Lee Yr7/6* Avocet S Yr7/6* Avocet S Yr7/6* Avocet S
Yr8 Compair Yr8/6* Avocet S Yr8/6* Avocet S Yr8/6* Avocet S
Yr9 Clement Yr9/6* Avocet S Yr9/6* Avocet S Yr9/6* Avocet S
Yr10 Moro Yr10/6* Avocet S Yr10/6* Avocet S Yr10/6* Avocet S
Yr15 T. dicoccoides (V763-251-wb) Yr15/6* Avocet S Yr15/6* Avocet S Yr15/6* Avocet S
Yr17 Shortim/VPM1 Yr17/3* Avocet S Yr17/6* Avocet S Yr17/6* Avocet S
Yr18 Jupateco R Yr18/3* Avocet S Yr18/3* Avocet S
Yr24 Meering2*//K733/T. tauschii (CPI 18911) Yr24/3* Avocet S
Yr26 Haynaldia villosa derivative (C94.153) Yr26/3* Avocet S
Yr27 Opata 85 YrSk/3* Avocet S YrSk/3* Avocet S
YrSP Spaldings Prolific YrSp/6* Avocet S YrSp/6* Avocet S
Avocet S Avocet S Avocet S
Supplemental Stocks
Yr2 Kalyansona
Yr18 Jupateco R Jupateco R Jupateco R
Jupateco S Jupateco S Jupateco S
Turkey Lebanon Syria India China Kenya Uganda Ecuador Chile Mexico
NIL 1998 1998 1998 1999 1999 1999 1998 1999 1998 1998 1999 1999 1999 1998 1998 1998 1998 1999
T. T. Xindu Chengdu Yaan Santa
Adana Izmir Ankara Haymana Gissar Terbol Ludhiana Sichuan Njoro Kalingree Carillanca Toluca
Hadya Hadya Sichuan Sichuan Sichuan Catalina
Yr1 /
R R R R - 5MR tR R 60S 40S 65S 40S 100S R R R 30MS 1S
6* Avocet S
Yr5 /
60 R,20 R 10R 10S 5MR tR R R R R R R 10MR 1MR R R R
6* Avocet S
Yr7 / 40MR-
80 60 80S 80S - 90S 95S 80S 40S 100S 20S 50S 40S 40MS 40S 70S 50MS
6* Avocet S 100S
Yr8 /
5 R R 30MS 90 30S 30M 60MS 30S - 20R 5R 5R 10M 5MS R R R
6* Avocet S
Yr9 /
80 60 70S 70S - 95S 95S 90S R R 100S 100S 100S 40S 40S 80S 90S 100S
6* Avocet S
Yr10 /
R R R R - 5MR 5R R R R R R R 10MR 1MR R R R
6* Avocet S
Yr15 /
R R R R R 5MR tR R - R R R R R 1MR R R R
6* Avocet S
Yr17 / 30MS-
R R R R 90S 70S - R 40S 10R 90S 10MR 1MS R 40MS 1MR
6* Avocet S S
Avocet S 100 100 90S 90S 80S 95S 95S 90S 80S 80S - 100S - 90S 30MS 90S 100S 100S
The development and application of near-isogenic lines for monitoring cereal rust pathogens
Avocet R
100 100 90S 90S R 95S 90S 80S 30S 100S 100S 100S 100S 80S 40S 90S 90S 100S
(YrA)
Jupateco R
40 20 50MS 60S 90 80S 80S 70MR - R 20S R 5R 40S R 30 10R 30M
(Yr18)
Jupateco S 60 60 70S 80S 90 90S 85S 80MS - R 20S 5R 90S 50MSS R 50 50MR 90M
Kalyansona 10 100 10MS 80S 90 90S 70S 80S - R 5R 10R 65S 30M 5MR 40M 20MS 40M
virulent avirulent
Table 3 Responses of the 1998 Avocet S NIL set to Pst at various locations
Turkey Lebanon Syria Iraq Iran
1999 1999 1999 2000 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000
NIL
Jolgeh Agh Jafar
Tel Neishabur Torogh Kalaleh Gorgan Ultan Gharakhil Alarogh
Eskisehir Terbol Tel Hadya Tuwaitha Rokh Ghala Abad
Hadya Khorasan Khorasan Golestan Golestan Ardabil Mazandaran Ardabil
Korasan Golestan Ardabil
Yr1 / 6*
R 5MR R R R 80S 60S 90S R R R tR tR R R
Avocet S
Yr5 / 6*
70S 5MR R R R R R R R R R tR tR 5R R
Avocet S
Yr6 / 6* 90S
- - 80S 85S 100S - - - - 80S 100S - - -
Avocet S
Yr7 / 6*
60S 90S 90S 80S 100S 70S 80S 80S 80S 20S 80S 90S 60S 20S 40S
Avocet S
Yr8 / 6*
30S 30S TMR 10MS 5R R R 70S 60S R R 10S 40S R 30S
Avocet S
Yr9 / 6*
50S 95S 90S 75S 100S 80S 50S 50S 100S 30S 70S 80S 100S 100S 70S
Avocet S
Yr10 / 6*
R 5MR R R R R R R R R R 20MR 20MR R 10MS
Avocet S
Yr15 / 6*
R 5MR R R R R R R R R R tR t MR 5R R
Avocet S
Yr17 / 6*
10MR 90S R 5MR 5R 30S 80S 40S R R 20S tR 20S 10S 30S
Avocet S
Yr18 / 3*
- - 20MR 20MR 65S - - - - - - - - - -
Avocet S
YrSp / 6*
- - R R R - R - - - R tR - - -
Avocet S
Yr27 / 3*
- - 20MR/MS 20MS 25MR - 25S - - - R 20MR - - -
Avocet S
Jupateco R
20MS 80S - 30MS _ 20S R 60S 50S 5R 5MS 30MR 30MR 40MS 30S
(Yr18)
-
Jupateco S 20S 90S 50S _ 50S 60S 60S 100S 15S 40S 80S 100S 100S 20S
Avocet R
90S 95S 90S 90S 100S 90S 70S 80S 100S 80S 60S 100S 100S 100S 70S
(YrA)
Avocet S 80S 95S 60S 80S 100S 80S 80S 90S 100S 20S 100S 100S 100S 100S 80S
81
82
Table 3 (cont)
NIL 2001 2001 2001 2001 2001 2001 2001 2000 2000 2000 2000 1999 1999
Yr6 / 6* Avocet S tR 90S 80S 70S 90S 20-50S 40S 80S 20S 60MS 5MR 40 80
Yr7 / 6* Avocet S tR 90S 80S 80S 90S 30S 50S 90S 20S 100S 5R 40 80
Yr9 / 6* Avocet S 20S 70S 60S 70S 70S 0-TMS 20S 60S 40S 5R 100S 10 40
The development and application of near-isogenic lines for monitoring cereal rust pathogens
Yr27 / 3* Avocet S tR tR tR tMS 20MR tR 20S 10MS R 10R R 10 40
Jupateco R (Yr18) 5MR tMR tR tMR 20MR 5-10MS tR 20MS 60S 90S R _ _
Jupateco S 5MR 5MR tRMR 40MS 60S 5MS 30S 50S 70S 100S 5MS _ _
Avocet R (YrA) 100S 100S 80S 80S 100S 90S 80S 80S 60S 100S 5 80S 80S
Avocet S 50S 90S 60S 50MS 50S 20-50S 60S 70S 70S 20MR 40 60S 80S
virulent avirulent
Table 4 Responses of the 1999 NIL set to Pst at various locations
Yr5 / 6* Avocet S R 0 0 0 R R 0 tR R 0 R
Yr6 / 6* Avocet S 15 MSS 10 10 87 30MSS 60S 60s 100S 50S 30S 90S
Yr7 / 6* Avocet S 12 MSS 65 60 87 70S 70S 70s 100S 90S 60S 90S
Yr10 / 6* Avocet S R 0 0 0 R R 0 R R 0 R
Yr15 / 6* Avocet S R 0 0 0 R R 0 R R 0 R
Yr18 / 3* Avocet S 10 MSS 35 40 40 tR MSS 40S 30s 60MRMS 70S 60RMR 90S
Yr26 / 3* Avocet S 1 MSS 5 0 33 R tR/20S Ts-10s tR, 70RMR 15MR 15RMR 70S
Jupateco R (Yr18) 10 MSS 10 20 47 5MR 40S 40S 15R 40S 20RMR 60S
Avocet R (YrA) 7 MSS 0 0 7 5R/MR 80S 70S 10R 100S 20SMS 90S
virulent avirulent
84 The development and application of near-isogenic lines for monitoring cereal rust pathogens
Discussion Breeding for resistance
Relevant and effective pathogenicity surveys Pathogen survey data should assume an integral
are important components of breeding programs role in the research and selection methods adopted
aimed at incorporating resistance to obligate plant within regional breeding programs, i.e. the survey
pathogens. Surveys monitor the distribution of current should retain the most appropriate pathotypes for use in
pathotypes, are directed at the early detection of new screening and for detection of new sources of resistance.
avirulence/virulence combinations of importance to Where pathotype isolates cannot be maintained, the
agriculture, and if results can be related genetically to project should identify the locations most likely to
cultivar genotypes, contribute to decisions on cultivar represent a conducive environment combined with
recommendation. Surveys also allow the selection of a regionally appropriate spectrum of Pst avirulence/
isolates of known pathogenic profile for use in screening virulence. An example encountered was the evident use
activities, and the accumulated historical data provide of relatively avirulent pathotypes in selection nurseries
valuable insights into pathogen epidemiology and in the main breeding center in Iran during the 2000
disease management. season, when pathotypes virulent for Yr9 were evidently
The requirements for pathogenicity assessments causing problems in other areas of the country. This
that are reliable, low cost and less dependent on situation subsequently changed and Iranian breeders
experienced observers with environmentally controlled and pathologists now select for resistance in the
facilities provided the momentum to re-examine presence of the most virulent pathotypes.
the materials and methods of field-sown differential
nurseries. Although this concept was introduced for Pst Recognition of new pathotype introductions
studies in western Europe by Zadoks in the 1960s, there The occurrence of Pst in regions clearly conducive
has been limited practical success due to difficulties to epidemic development, but previously free of the
associated with the availability of suitable materials disease, has resulted in significant national issues over
of known genotype for disease and pathogenicity the past 20 years. This progressive extension of the
monitoring. Phenological variation among the geographical distribution of P. striiformis has served to
traditional differential sets used in seedling- based highlight the increased potential for foreign pathotype
pathotype assays, and the confounding influence of incursions. Notable examples include:
additional uncharacterized adult plant resistances, (i) Barley stripe rust (P. striiformis f. sp. hordei) in
resulted in contentious results that could not easily Central America in 1975 (Dubin and Stubbs 1986) and
be reconciled with seedling tests. The Avocet S NILs subsequently North America in 1991 (Marshall
overcame some of these constraints and provided and Sutton 1995); it was considered an introduction
helpful, and in some instances unique, insights into the from Europe.
nature of the Pst population across a broad range of (ii) Wheat stripe rust in Australia in 1979 was
locations. The value of this information can be assessed attributed to a single pathotype introduced from
according to the following criteria. Europe (Wellings 2007).
(iii) Wheat stripe rust in South Africa was first
reported in 1996 (Pretorius et al. 1997).
Predicting disease response
(iv) P. striiformis isolates from barley grasses
Efforts aimed at monitoring pathogenicity
(Hordeum spp.) in 1998 were shown to be highly
characteristics in the Pst population in major regions
avirulent on Pst differentials and concluded to be a new
vulnerable to epidemics are able to assist in the
introduction to Australia (Wellings et al. 2000b).
development of control strategies through predicting
(v) New pathotypes of Pst in North America from
and recommending cultivars with resistances known to
2000 (Chen 2005); origins remain unknown.
be effective against current Pst pathotypes. For example,
(vi) The occurrence of Pst in Western Australia for
Inqualab 91 (Yr27) was predicted on the basis of the data
the first time in 2002 was concluded to be due to a
from this project to protect against the advancing Yr9-
foreign pathotype incursion, and suspected to have
virulent pathotype in Pakistan, although it subsequently
originated from the Pst population introduced to North
became ineffective with the occurrence of Yr27 virulence
America in 2000 (Wellings et al. 2003).
in 2002 (Duvellier et al. 2007).
86 The development and application of near-isogenic lines for monitoring cereal rust pathogens
Riley R, Chapman V, Johnson R (1968) Introduction of Wellings CR, McIntosh RA (1990) Puccinia striiformis f.sp.
yellow rust resistance of Aegilops comosa into wheat tritici in Australasia: pathogenic changes during the
by genetically induced homoeologous recombination. first ten years. Plant Pathol 39:316-325
Nature 217:383-384 Wellings CR, McIntosh RA, Hussain M (1988) A new source
Saari EE, Prescott JM (1985) World distribution in relation of resistance to Puccinia striiformis f.sp. tritici in spring
to economic losses. In: Roelfs AP, Bushnell WR (eds) wheats (Triticum aestivum). Plant Breeding 100:88-96
The cereal rusts Vol II, Academic Press Inc. Orlando, pp Wellings CR, Singh RP, Yahyaoui A, McIntosh RA (2000a)
259-298 The assessment and significance of pathogenic
Singh RP, William HM, Huerta-Espino J, Rosewarne G variability in Puccinia striiformis in breeding for
(2004) Wheat rust in Asia: meeting the challenges with resistance to stripe rust: Australian and international
old and new technologies. ‘New directions for a diverse studies. Proc 11th Regional Wheat Workshop, CIMMYT,
plant’, Proc 4th Int Crop Sci Cong, Brisbane Australia. Addis Ababa, Ethiopia, pp 134-143
www.cropscience.org.au Wellings CR, Burdon JJ, McIntosh RA, Wallwork H et al
Stubbs RW (1985) Stripe rust. In: Roelfs AP, Bushnell (2000b) A new variant in Puccinia striiformis causing
WR (eds) The cereal rusts Vol II, Academic Press Inc. stripe rust on barley and wild Hordeum in Australia.
Orlando, pp 61-101 Plant Pathol 49:803
Wan A, Zhao Z, Chen XM, He Z et al (2004) Wheat stripe Wellings CR, Wright DG, Keiper F, Loughman R (2003) First
rust epidemic and virulence of Puccinia striiformis f.sp. detection of wheat stripe rust in Western Australia:
tritici in China in 2002. Plant Dis 88:896-904 evidence for a foreign incursion. Aust Plant Pathol
Wellings CR (1992) Resistance to stripe (yellow) rust in 32:321-322
selected spring wheats. Vort fur Pflan 24:273-275 Yahyaoui AH, Hakim MS, El Naimi M, Rbeiz N (2002)
Wellings CR (2007) Puccinia striiformis in Australia: A Evolution of physiologic races and virulence of Puccinia
review of the incursion, evolution and adaptation of striiformis on wheat in Syria and Lebanon. Plant Dis
stripe rust in the period 1979-2006. Aus J Agric Res 86:499-504
58:567-575
Table 1. Isolates, race identities, origin, and avirulence/virulence formulae of Puccinia graminis f. sp. tritici isolates used
in seedling tests
Isolate Race Origin Avirulence/ virulence formula
04KEN156/04 TTKSK Kenya Sr24 36 Tmp/ 5 6 7b 8a 9a 9b 9d 9e 9g 10 11 17 21 30 31 38 McN
01MN89A-1-2 TTTTF USA Sr24 31/ 5 6 7b 8a 9a 9b 9d 9e 9g 10 11 17 21 30 36 38 Tmp McN
06YEM34-1 TRTTF Yemen Sr8a 24 31/ 5 6 7b 9a 9b 9d 9e 9g 10 11 17 21 30 36 38 Tmp McN
06ND76C QFCSC USA Sr6 7b 9b 9e 11 24 30 31 36 38 Tmp/ 5 8a 9a 9d 9g 10 17 21 McN
59KS19 MCCFC USA Sr6 8a 9a 9b 9d 9e 11 21 24 30 31 36 38/ 5 7b 9g 10 17 Tmp McN
1
USDA-ARS, Cereal Disease Laboratory, and 2Department of Plant Pathology,
University of Minnesota, St. Paul, MN 55108, USA
E-mail: yue.jin@ars.usda.gov
resistant to all five races. Based on the reactions to the have Sr36. Accession CI 11802, which was regarded as the
five races, there appeared to be novel resistance to TTKSK standard for Sr37 (McIntosh et al. 1985), produced IT ;1 to
in T. monococcum. Additional studies are needed to TTKSK, and 3 and 4 to TRTTF and TTTTF, respectively. Five
confirm gene postulations and to determine the number accessions of T. timopheevii ssp. timopheevii produced
and allelic relationships of the potentially new genes. a similar pattern, with ITs ranging from ;1 to ;12+. Three
accessions of T. timopheevii ssp. armeniacum showed
Aegilops tauschii A total of 456 accessions of resistance to TTKSK, with ITs ;1 and ;12. The relationship
Ae. tauschii from the USDA NSGC and Wheat Genetic between resistance in this germplasm and Sr40 is
and Genomic Resources Center (Manhattan, KS) were uncertain at this time.
evaluated for resistance to races TTKSK, TRTTF, and
TTTTF. Sixty-five accessions (14.7%) were resistant T. turgidum ssp. dicoccoides A collection of 157
to TTKSK with ITs ranging from ; to 2+, and eight accessions from the USDA NSGC was evaluated for
accessions (1.8%) were resistant to all races. Based resistance to races TTKSK, TRTTF and TTTTF. There was a
upon the different infection types to various races, we low frequency of resistance to the three races: 16.6% to
postulated that one or more novel genes for resistance TTKSK, 8.9% to TRTTF, and 11.5% to TTTTF. Low ITs were
to race TTKSK is present in this species. Selected variable, ranging from 0; to X with the majority being
accessions are being backcrossed to hexaploid wheat 2 to 2+. Three accessions were resistant to both TTKSK
to introgress the resistances to race TTKSK. and TRTTF, one accession was resistant to TTKSK and
TTTTF, and two accessions were resistant to all three
Ae. speltoides Ninety-two accessions from races.
the USDA NSGC were evaluated and all accessions
were resistant to race TTKSK. One plant from a X Triticosecale A collection of 567 triticale
heterogeneous accession was susceptible with IT 3. accessions from the USDA NSGC was evaluated for
Most of the accessions exhibited ITs from 0 to ;1. The resistance to races TTKSK, TRTTF and TTTTF. There
frequencies for resistance to races TTTTF (96%) and was a high frequency of resistance to race TTKSK; 440
TRTTF (98%) also were high. Although this species accessions (77.7%) exhibited low ITs ranging from 0; to
offers a high frequency of TTKSK resistance, it will be 2+ (Table 2). Based on the ITs we postulated a number
difficult to differentiate novel resistance from resistance of resistance genes, viz. Sr27 (IT ;2= to ;12), SrSatu (IT
genes already transferred to common wheat, such as ;CN), SrVen (IT ;13) or SrNin (IT 22-) (McIntosh et al. 1985).
Sr32, Sr39 and Sr47. ITs different from those conferred by known genes
were also observed, suggesting the presence of novel
T. timopheevii A collection of 298 accessions from resistance genes or unique combinations of known
the USDA NSGC were evaluated, and 15% were resistant genes. Screening of the resistant lines with P. graminis f.
to race TTKSK with ITs ranging from 0 to ;12+. Resistance sp. tritici races virulent on known triticale genes available
to races TTTTF and TRTTF was not detected among these in Australia and South Africa will help to facilitate the
accessions. Accessions resistant to TTKSK were mostly identification of any new resistance genes. Resistance
from T. timopheevii ssp. timopheevii, with infection types to the three races was highly associated as the majority
predominantly 0 to 0;. The low ITs (0 or 0;) to race TTKSK (87%) of the accessions, that were resistant to TTKSK,
in combination with susceptibility to races TRTTF and were also resistant to races TTTTF and TRTTF.
TTTTF suggested that the majority of resistant accessions
90 Progress and prospects in discovery and use of novel sources of stem rust resistance
References
Jin Y, Singh RP, Ward RW, Wanyera R et al (2007)
Characterization of seedling infection types and adult
plant infection responses of monogenic Sr gene lines
to race TTKS of Puccinia graminis f. sp. tritici. Plant Dis
91:1096-1099
McIntosh RA, Wellings CR, Park RF (1995) Wheat rusts: An
atlas of resistance genes. CSIRO, Melbourne, Australia,
200pp
F1 ph1b ph1b
Select ph1b ph1b Sr sr
plants by testing with
Elite line (stem rust susceptible)
stem rust and Ph1 locus
DNA markers
BC1F1
Screen 100s-1000s of progeny with stem rust and translocation-specific DNA markers to identify recombinants
Our goal is to make useful to variety development progeny with smaller translocations. After reducing the
programs, effective stem rust resistance genes derived amount of alien chromatin, the Sr genes are transferred
from wild species. Numerous stem rust resistance genes to elite wheat germplasm adapted to Africa and/or Asia.
(Sr) from wheat relatives, such as Aegilops speltoides The following summaries document progress in our
Tausch, Thinopyrum ponticum (Podp.) Barkworth & D.R. ongoing chromosome engineering efforts on each alien-
Dewey, Triticum timopheevii (Zhuk.) Zhuk., and Secale derived stem rust resistance gene.
cereale L., have been incorporated into wheat genomes
in the form of chromosome translocations. More than Approaches to induce homoeologous recombination
a dozen of these genes are effective against race TTKSK The preferred method for reducing the size
(Ug99) and related derivatives (Singh et al. 2006; Jin et al. of alien segments is to employ the ph1b mutant
2007). Unfortunately, most of them are associated with of hexaploid wheat. In this approach, either F2 or
deleterious linkage drag. Reducing the amount of alien BC1F1 populations are firstly produced from crosses
chromatin increases the likelihood of a translocation between the translocation lines and ph1b mutant
having commercial value. Their manipulation and use stocks. DNA markers that detect the ph1b mutation,
is important given the overarching goal of long-lasting and stem rust phenotypic screening, are applied to
rust protection in wheat crops worldwide, particularly progeny to identify individual plants homozygous
when two or more broadly effective genes are ph1b ph1b and hemizygous for the target translocation
pyramided and/or combined with minor-gene resistance and homoeologous wheat chromosomes (Fig. 1).
in a single cultivar. Populations developed from the selected plants are
The specific gene targets of this project are Sr32, then screened for stem rust resistance and genotyped
Sr37, Sr39, Sr40, Sr43, Sr44, Sr47, SrAeg5, SrAsp5, SrAse3, for translocation-specific marker alleles to identify
SrHv6, SrTt3, and Sr2S#1. Traditional hybridization and putative recombinant progeny. Fluorescent genomic
chromosome manipulation methods are coupled with in situ hybridization (GISH) is then applied on putative
DNA marker development, stem rust phenotyping, and recombinants to confirm shorter translocation lines
genotyping of large populations to identify recombinant carrying each Sr gene.
was developed by crossing TA5599 and TA5602. The results, resistant progeny had 5SL, whereas susceptible
F3 families were evaluated for stem rust resistance progeny lacked 5SL. Putative Robertsonian events
by inoculation with race RKQQ at the two-leaf stage. were identified as progeny lacking 5SS marker alleles
Transmission of T5MgS·5MgL-5DL (23 homozygous and having 5SL marker alleles. F3 families of putative
resistant: 80 segregating: 44 homozygous susceptible; Robertsonian progeny were screened by C-banding.
chi21:2:1 = 7.12; P<0.05) was significantly reduced, but Family U5909-2-166 was identified as a Robertsonian
SrAge5 appeared to segregate as a single gene because translocation T5DS•5S#3L; the others were telosomic
DNA markers tagging 5MgL co-segregated with lines. U5909-2-166 was crossed to CS ph1b to develop
resistance. populations for reducing the size of this translocation.
SrAsp5. TA7693, a disomic addition line with SrHv6. TA7682, a disomic addition line with
chromosome 5S of Ae. speltoides (21”+1” [5S#3]) in CS chromosome 6V of Haynaldia villosa in CS background
background, is resistant to North American races and (21”+1” [6V#3]), is resistant to North American stem rust
race TTKSK. TA7693 was crossed with CS M5D. Selected races and TTKSK. Addition lines involving the other six
double monosomic F1 plants were self-pollinated and chromosomes from the same donor were susceptible.
~250 F2 progeny were screened with race RKQQ and A population was produced from the cross CS M6A
characterized by molecular markers to identify putative (20” + 6A’) / DA 6V in an effort to derive Robertsonian
Robertsonian translocation progeny. Based on marker translocation chromosomes involving the 6V short and
102 Cloned rust resistance genes and gene based molecular markers in wheat: Current status and future prospects.
16. Molecular-genetic dissection of stem rust resistance gene that had been effective for
30 years (Pretorius et al. 2000; Singh et al. 2006; 2007;
rice nonhost resistance to wheat Stokstad 2007). Subsequent mutation of this isolate
has led to the breakdown of further resistance genes
stem rust (e.g. Sr24, Sr36) making many of the world’s commercial
cultivars vulnerable to stem rust epidemics (Singh et al.
Michael Ayliffe1, Yue Jin2, Brian Steffenson3, Zhensheng 2006; 2007; Stockstad 2007).
Kang4, Shiping Wang5, Hei Leung6 In contrast, rice, which is an equally important
cereal in terms of food production, is apparently
Abstract immune to all known rust diseases. This species is
Rust diseases remain a significant threat to the unique compared with all other cereals and cultivated
production of most cereals including wheat. New grasses, which are parasitized by at least one rust
sources of resistance are continually sought by breeders pathogen (e.g. wheat, barley, rye, triticale, maize,
to combat the emergence of new pathogen races. Rice sorghum, millet, oats, sugarcane). The immunity of rice
is atypical in that it is an intensively grown cereal with to rust disease is presumably mediated by nonhost
no known rust pathogen. The resistance of rice to cereal resistance (NHR), a resistance mechanism that has
rust diseases is referred to as nonhost resistance (NHR), recently become tractable using molecular-genetic
a resistance mechanism that has only recently become approaches.
genetically tractable. In this report, the mechanisms Given the apparent durability of NHR, an attractive
of rice NHR to wheat stem rust and other cereal rust proposition is to transfer this rust resistance from rice
diseases are explored and the potential for transferring into wheat and other agricultural cereals. This paper
this durable disease resistance to wheat is considered. describes approaches being undertaken to characterize
Approaches being undertaken for the molecular-genetic the nonhost resistance of rice to cereal rusts at a
dissection of rice NHR to rust are described. molecular genetic level and explores the possibility of
transferring this resistance to other cereals.
Keywords:
Puccinia, Oryza, Triticum, effector, immunity Host and nonhost resistance – mechanistically distinct
or overlapping processes?
Introduction The current model of plant disease resistance (Jones
Rust diseases caused by fungal pathogens in and Dangl 2006) proposes that microbes produce an
the Puccinia genus remain a constant threat to cereal unknown number of conserved molecules that plant
production. Wheat, one of the world’s most important cells can recognize with membrane spanning receptor
agricultural crops, accounting for 30% of global calorific kinases (reviewed by Zipfel 2008). These microbial
intake, is subject to three rust diseases. These diseases molecules, called MAMPs or PAMPs (microbe or
are wheat leaf rust, wheat stripe rust and wheat stem pathogen associated molecular patterns) include diverse
rust caused by the fungal pathogens Puccinia triticina, molecules such as flagellin, chitin, lipopolysacharide
Puccinia striiformis f. sp tritici and Puccinia graminis f. sp. and translation elongation factors. Upon recognition
tritici, respectively. of these molecules a basal defense response, or PAMP-
New sources of resistance to these three wheat triggered immunity, is activated that prevents further
pathogens are continually sought by breeders due to microbial colonization (Zipfel 2008). Pathogens of a
the ability of the pathogens to overcome host resistance given plant species have the capacity to circumvent
by a combination of mutation, parasexuality and this basal defense response by introducing a suite
sexual recombination. Effective, durable resistance to of molecules termed effectors into plant cells which
rust diseases has been coveted by wheat agricultural suppress host cell defenses by interacting with specific
scientists (Ayliffe et al. 2008). A case in point is the host target molecules (reviewed by Hogenhout et al.
emergence of a new stem rust isolate from Uganda (race 2009). In turn plants have evolved a large number of
Ug99) in 1999 that can overcome Sr31, a highly effective genes that encode resistance proteins (R proteins), each
of which recognizes a specific pathogen effector. Upon
1
CSIRO Plant Industry, Box 1600, Canberra, ACT, 2601, Australia;. 2USDA Cereal
Rust Laboratory and 3Department of Plant Pathology, University of Minnesota,, R protein recognition of an effector, previously known
St. Paul, MN 55108, USA; 4College of Plant Protection, Northwest Agriculture as an avirulence product, a resistance response, or
and Forestry University, Yangling, Shaanxi 712100, China; 5National Key
Laboratory of Crop Genetic Improvement, Huazhong Agricultural University, effector-triggered immunity, is activated that frequently
Wuhan 430070, China; 6International Rice Research Institute, DAPO Box 7777, involves hypersensitive cell death. This effector triggered
Manila, The Philippines
E-Mail: michael.ayliffe@csiro.au
Fig. 1 Plant defense mechanisms that must be circumvented for successful infection by a plant pathogen
en
en
og
og
en
n
th
th
ge
og
pa
pa
ho
th
n
ge
d
ed
pa
at
te
ho
pt
tp
ap
nt
da
at
en
le
np
na
na
iru
l
ru
no
no
no
av
vi
Basic compatibility
physical and chemical
Inappropriate signals
signals
Induced basal
Inappropriate
effectors
defenses
PAMP triggered immunity
R protein R protein-mediated
INFECTION recognised
effector
defense
Effector triggered immunity
Fig. 1 Comparative field response (R-MR, up to 20% disease severity with small uredinia; MR-MS, up to 40% disease
severity with medium uredinia; MS-S, 50-100% disease severity with medium to large uredinia) of germplasm screened
at KARI during 2006-2008
90
80
70
60
Percent entries
50 R-MR
40 MR-MS
30 MS
20
10
0
2006 2007 2007 Total
Year
3 2
118 Developing and optimizing markers for stem rust resistance in wheat
for source, markers available, current research activities, markers to improve the resolution of the relationships.
and prioritized for this project (http://rustopedia.get- DNA of each line was also sent to Triticarte, Australia
traction.com/traction). Haplotyping was initiated for (http://www.triticarte.com.au) for DArT analysis. The
stem rust resistance genes, including Sr1A1R, Sr2, Sr9a, identified DArT markers will be used for haplotyping and
Sr13, Sr15, Sr17, Sr19, Sr22, Sr24, Sr25, Sr26, Sr31, Sr32, association analysis in combination with the phenotypic
Sr33, Sr35, Sr36, Sr38, Sr40, Sr45, Sr46, and SrR. Table 1 lists data. Our goal is to be able to predict the stem rust
the major stem rust resistance genes and linked markers, resistance genotype.
including SSR, STS, BARC, and DArT markers. Progress in pyramiding stem rust resistance: Our
Genetic resources: We started with a diverse pyramiding work is a collaboraton with Dr. Gina Brown-
collection of wheat accessions selected by CIMMYT and Guidera, director of the USDA genotyping laboratory in
ICARDA wheat breeders. This collection consists of more Raleigh, NC and Dr. Michael Pumphrey, ARS-Manhattan,
than 300 lines from CYMMIT and ICARDA programs in KS. As the initial targets for pyramiding, we made F1
Africa, China, Turkey and Mexico. In the present study, hybrids by intercrossing wheat lines with combinations
we will report on 260 wheat lines of diverse origins of Sr22, 24, 32, 36 and Amigo, and also between the
including 115 lines from CIMMYT, 43 lines from China, sources of these genes and local lines with high yield and
and 102 lines of miscellaneous origins. To estimate their quality in fall 2008. More crosses will be made between
genetic relationships to known stem rust resistance these sources of major genes and CIMMYT, Chinese and
gene sources, we also included wheat lines with known African wheat germplasm. Our goal is to develop wheat
Sr genes when available. germplasm with durable resistance to Ug99 for the high
Marker validation: To evaluate the functionality risk wheat growing regions, especially Africa.
and polymorphism for the available markers, we first
screened 58 markers associated with 21 stem rust Acknowledgements
resistance genes among 16 randomly selected wheat We acknowledge the germplasm and information
lines. Using DNA from leaf tissue, we tested the primers provided by our co-authors, as well as collaborators
for each marker (Table 1) and analyzed the polymerase Drs. Gina Brown-Guidera, Michael Pumphrey, Alexi
chain reaction (PCR) products using agarose gels. Forty Morgounov, Zhonghu He, Steven Xu, and Yue Jin, in the
six (80%) of the markers amplified clear fragments and, preparation of this report.
of those, 35 (75%) showed polymorphism.
Haplotyping diverse wheat germplasm: We then Section 2. Developing and optimizing
extracted DNA from the other 260 wheat lines and markers for Sr13 and pyramiding Sr2, Sr13
analyzed the haplotypes by comparison with known and Sr25 in tetraploid what (Contributed by
sources of stem rust resistance genes. PCR amplification Zewdie Abate and Jorge Dubcovsky)
was carried out using primers of molecular markers
associated with major stem rust resistance genes as Specific objectives assigned to the UC Davis program
shown in Table 1. PCR products were analyzed using During the March 12, 2008 meeting of the stem
both PAGE and ABI 3730. To date, 20 markers associated rust marker group at UC Davis, it was decided that the
with major genes Sr1A1R, Sr2, Sr9a, Sr13, Sr17, Sr19, UC Davis program would focus on stem rust resistance
Sr22, Sr24, Sr25, Sr31, Sr32, Sr35, Sr36, Sr40 and Sr44 were genes from diploid and tetraploid wheats. It was also
analyzed. The sizes of PCR amplicons were recorded. agreed that the pyramiding activities at UCD would
To analyze the distribution of alleles linked with the incorporate multiple stem rust resistance genes effective
stem rust resistance QTLs, we grouped PCR amplicons against Ug99 into high yielding tetraploid backgrounds
based on their fragment sizes (in bp) among wheat lines that already have been targeted for the incorporation
analyzed. PCR amplicons of the same size were grouped of stripe rust resistance genes. These tetraploid lines
together and color coded to help interpretation of will be excellent parental lines to deliver multiple Sr
haplotype structure. Fig. 1 shows haplotype groups for and Yr genes in simple crosses with germplasm in
10 loci in our panel of accessions. Of those with known the SEWANA durum production regions. This activity
reactions to stem rust, a group of susceptible and will also include the transfer of resistance genes from
moderately susceptible lines were grouped together hexaploid to tetraploid wheat. In the area of mapping,
as a susceptible haplotype (blue color). Genotypes the UC Davis group will focus on the precise mapping of
containing the same gene such as Sr25 or with genes Sr13 and Sr25 and on the discovery of new sources
resistant phenotypes were sorted together as resistant of resistance against Ug99 in tetraploid wheat and T.
haplotypes (red color). These preliminary results were monococcum mapping populations.
from a small number of markers and we will use more
120 Developing and optimizing markers for stem rust resistance in wheat
Fig. 1 Haplotype analysis for stem rust resistance for wheat lines of known or unknown resistance. PCR amplicon
fragment sizes are shown for each accession along with the stem rust reaction or gene if known. R, resistant; S,
susceptible; MR and MS, moderately resistant and susceptible, respectively. Color codes: pink for resistance (bold),
blue for susceptible (bold italic) and yellow for unknown phenotypes (underlined)
BC4F1 BC4F1
For public release by 07/10
Gene selection: We completed a survey of available Y which is desirable in durum wheat (T. turgidum var.
stem rust resistance genes in tetraploid wheat. We durum) (Zhang et al. 2005).
prioritized genes that conferred resistance to Ug99, Pyramiding of stem rust resistance genes in tetraploid
were mapped, and for which molecular markers wheat
were available. Based on these criteria we selected Recurrent parent: Breeding line UC1113 was
resistance genes Sr2, Sr13, and Sr25 as the initial selected as a recurrent parent for initial pyramiding. This
targets for pyramiding. line was selected for several reasons. Firstly, UC1113 is
Sr13 originated from T. turgidum var. dicoccum cv. a high yielding semi-dwarf durum variety with a very
Khapli and was transferred to several wheat cultivars broad adaptability in California. Secondly, we have
grown worldwide (McIntosh et al.1995). already introgressed in the high-grain protein content
Sr2 was introgressed from T. turgidum var. gene Gpc-B1 from wild wheat (Uauy et al. 2006) and
dicoccum cv. Yaroslav and transferred into several wheat the non-race-specific stripe rust resistance gene Yr36
backgrounds. It provides partial resistance to Ug99 and (Uauy et al. 2005) in this line. The incorporation of the
this resistance has been effective for the last 80 years Gpc-B1 gene increased grain protein, iron and zinc by
(McIntosh et al. 1995; Singh et al. 2006). The resistance approximately 10%. Finally, UC1113 is a public breeding
conferred by Sr2 alone is not sufficient and therefore, it is line that can be distributed without intellectual property
necessary to combine this gene with other Sr genes for constraints. The UC1113 line with Gpc-B1 and Yr36 genes
full protection (Ayliffe et al. 2008). has been already deposited in the USDA-ARS National
Sr25 is present in a distal 7EL translocation from Small Grain Collection (PI 638741) (Chicaiza et al. 2006).
Th. ponticum. The 7EL translocation also carries leaf rust We will incorporate additional recurrent durum parents
resistance gene Lr19 and yellow flour pigment gene from Ethiopia when we receive the seeds.
122 Developing and optimizing markers for stem rust resistance in wheat
Table 2 Reaction of UC1113 and Kofa for 9 stem rust races
Race QTHJ RCRS RKQQ TPMK TTTT TTKSK TTKSK TTKST TTTSK
UC1113 ; ;1N ; 3+;1 4 4 4 4 4
Kofa ; ;N ; ;1 ;1 0;/2/4 2 2 2
Progress on pyramiding Sr2, Sr13 and Sr25 in scored against races TTKSK and TTTTF by Dr. Yue Jin and
durum wheat: The initial objective of the pyramiding a single resistance genes, Sr13, was mapped at the end
work in tetraploid wheat is to combine the stem rust of chromosome 6AL between microsatellite markers
resistance genes Sr2, Sr13 and Sr25 in a common genetic wmc580 and dupw167 (Fig. 3). The complete molecular
background. The presence of multiple resistance map has 269 markers (including 23 SNP) and a total
genes is expected to extend the durability of the length of 2,140 cM.
individual genes. Th. elongatum is the source of Sr25, To refine the mapping position of Sr13 we
Yaroslav the donor of Sr2 and Khapli for Sr13. The Th. are backcrossing the Sr13 region into UC1113 and
elongatum chromosome segment also carries the leaf generating BC2F2 segregating seed for high-density
rust resistance gene Lr19. We have completed the mapping. The F2 plants will be screened to identify
incorporation of the Sr25/Lr19 genes together with the recombination events between Sr13 flanking markers
Yr36/Gpc-B1 genes into UC1113 by six backcrosses. (Fig. 3). Seed will be saved from plants heterozygous
The pyramiding will be carried out in three streams at both flanking markers to generate seeds for a high
(Fig. 2). Stream 1 will be used to combine Sr13 and density genetic map in order to identify a tightly linked
Sr25/Lr19, and Stream 3 to combine Sr2 and Sr25/Lr19. diagnostic marker.
Backcrossing 1 and 3 will converge into Stream 2 to put Finally, two additional populations are being
the three genes together. The initial cross to combine developed for validation of the identity of Sr13 using
the linked Sr25/Lr19 genes in UC1113 background Khapli (CItr4013) as one of the parental lines. The crosses
with Sr2 (Yaroslav) was completed in July 2008. We are Rusty/Khapli and 47-1/Khapli have been completed and
currently growing the F1s to generate the first backcross the F1 plants are currently growing to produce F2 seeds
generation to combine different genes in Stream 3. for validation. Once closer markers become available
For stream 1, parents are currently growing F1s for Sr13 we will check Kofa, Khapli, and Khapstein (the
carrying the Sr25/Lr19 and Sr13 resistance genes (in germplasm used to name Sr13) to confirm if they all
addition to Gpc-B1/Yr36). A line homozygous for the have the same haplotype.
three genes (BC1F2) should be available by mid 2010 Sr2 is located on the short arm of wheat
(assuming 4 months per generation). Although this line chromosome 3B (Hare and Mcintosh 1979) closely linked
would not be fixed for the rest of the genome, it will to microsatellite marker Xgwm533 (Spielmeyer et al.
be a useful parental line, and will be distributed to all 2003). Its recessive inheritance of resistance expressed
interested wheat breeding programs. We will continue at the adult plant stage complicates selection for this
the backcrossing program to obtain more stable lines gene (Knott 1968). Drs. W. Spielmeyer and E. Lagudah
for different gene combinations. developed a high-density genetic map of Sr2 and
designed a diagnostic marker for the gene. They have
Mapping and marker development in provided us access to this unpublished marker, and
tetraploid wheat we have confirmed that it is diagnostic for Sr2 in our
Sr13 is located on wheat chromosome 6AL germplasm. Using this marker we have confirmed the
(Klindworth et al. 2007; McIntosh et al. 1995). presence of Sr2 in our seed source of Yaroslav (Fig. 4),
Unpublished results at USDA-ARS Biosciences Research which was essential for our pyramiding work.
Lab showed that Sr13 is closely linked (≈2cM) to marker Sr25 was mapped on the 7EL chromosome segment
Xwmc580 (Drs. Shiaoman Chao, North Dakota; and Evans from Th. ponticum distal to the Lr19 locus and close to
Lagudah, CSIRO). The UCD group is collaborating with the yellow pigment gene Y (Lr19-Sr25/Y). We developed
Drs. Chao and Lagudah to define precisely the location a tetraploid line carrying Sr25, Lr19, and Y in the UC1113
of Sr13. genetic background (line 1-23, Fig. 5) (Zhang et al.
The UCD group used a segregating population 2005). This 7EL segment does not recombine with wheat
from the cross between tetraploid lines Kofa (resistant chromosomes in the presence of Ph1 so any marker
to TTTTF and TTKSK) and UC1113 (susceptible to TTTTF for the 7EL segment works as a perfect marker for Sr25
and TTKSK) to map Sr13 (Table 2). The 93 SSD lines were (Zhang et al. 2005; Zhang et al. 2008).
To better define the map location of Sr25 within the most distal ESTs (BF483039, BG262960, and BF484041)
7EL segment we sent the recombinant lines to Kenya and we are testing them for polymorphism between the
and based on the resistance scores against Ug99 we 7E and 7J alleles.
mapped Sr25 linked to the Phytoene Synthetase 1 gene
(PSY1) at the distal end of chromosome 7EL (Fig. 5). Mapping and marker development in diploid wheat
Precise mapping of Sr25 is important for the T. monococcum
engineering of a shorter segment of 7EL carrying In collaboration with Dr. Yue Jin we screened T.
Sr25 and also for the engineering of a chromosome monococcum lines DV92 and G3116 (parents of the
segment combining Lr19/Sr25 and barley yellow dwarf mapping population DV92 x G3116 (Dubcovsky et al.
resistance gene Bdv2. Bdv2 was mapped on the distal 1996) with seven different stem rust races including
region of chromosome 7JL from T. intermedium, which Ug99 (Table 3).
is co-linear with the 7EL region where Sr25 is located. We developed a single seed descent mapping
In collaboration with Dr. Adam Lukaszewski (University population from the cross DV92/G3116 (150 lines).
of California, Riverside) we developed several lines Results from the evaluation of these lines with races
carrying recombinant 7EL/7JL chromosomes and we RKQQC and TRTTF (Dr. Yue Jin) suggest segregation for 2
started their characterization with molecular markers. resistance genes (one of them probably Sr21).
We sent those lines to Kenya and to Dr. Yue Jin to test Dr. Yue Jin has completed the screening of a large
their reaction to Ug99. number of T. monococcum accessions and new crosses
We are also developing markers distal to PSY1 to are being made. Once we clarify the genes segregating
better define the position of Sr25. We have selected the in the available populations we will use these new
populations to identify new sources of resistance.
124 Developing and optimizing markers for stem rust resistance in wheat
Fig. 4 The unpublished diagnostic marker for Sr2 among species containing stem rust resistance genes
amplifies a ~400 bp band in Yaroslav (PI 2789) and that have been transferred to bread wheat. Three named
Opata 85, but not in Langdon, confirming the presence genes derived from A. tauschii are Sr33, Sr45, Sr46; the
of Sr2 in Yaroslav (Sr2 source of our pyramiding work). first two genes are located on wheat chromosome
1DS and the third on 2DS. One of the advantages of
bp
genes derived from A. tauschii when compared with
Opata 85
Langdon
Yaroslav
750
other wheat relatives, is the ease of gene transfer by
500 homologous recombination with the corresponding D
400 genome of hexaploid bread wheat. The vast germplasm
300 resource of so-called synthetic hexaploids obtained
from crossing tetraploid wheat with A. tauschii provides
200 a potentially rich source of genetic variability for rust
resistance. However, to ensure that unique resistance
genes can be identified, they need to be differentiated
from the known genes. A variety of methods that can
be used include response of a given genotype to a
Table 3 Reaction of three T. monococcum accessions to range of pathogen isolates, chromosomal location /
different races of stem rust (Yue Jin 06/08, 01/08, 12/08) genetic analysis and the use of tightly linked or
diagnostic markers. Similar infection types produced by
Race G1777 G2528 G3116 DV92
resistance genes present in the tetraploid wheats used
TTTT 3 3- 3 - in creating the synthetic hexaploids can often confound
the identification of new sources of resistance from A.
; 3, 4,
TPMK 4 4 4 tauschii in the absence of discriminating pathotypes.
very LIF
The availability of diagnostic markers for the known
RKQQ 4 4 4 0 or named stem rust resistance genes provides a useful
tool to establish the unique identities of putatively new
TTKSK 1/1 2Z ;12- 22+ 0;
sources of resistance. A diagnostic marker for the Sr46
TTKST ;1 ; 1 2- 22 + 0; gene was recently developed (Lagudah unpublished ).
Efforts to develop markers for Sr33 and Sr45 are
TTTSK ; 1 2- ;1 12 0;
in progress. Sr33 has shown resistance to a wide range
TRTT 33+ 33+ 3+4 0 of stem rust isolates, and often shows an intermediate
response. In tests conducted with Ug99, Sr33 provided
TTKSK= Ug99; TTKST = Ug99 + Sr24 virulence; TTTSK
a moderate level of resistance under field conditions in
= Ug99 + Sr36 virulence; TRTT: race from Yemen with
Kenya (Jin et al. 2007). Stem rust isolates virulent for Sr45
virulence on lines with 1AL.1RS. LIF, Low Infection
have previously been reported. In comparisons between
Frequency; many leaves with IT 0, 1, or 2
Sr33 and Sr45 against isolates where both genes are
effective, the latter often shows a stronger resistance
Acknowledgements response (McIntosh et al. 1995; Sambasivam et al. 2008).
We acknowledge the support and information Seedling tests conducted using Ug99 (TTKS) have shown
provided by our colleagues Dr. Shiaoman Chao, Dr. Yue Sr45 to be effective.
Jin, Dr. Adam Lukaszewski and Dr. Evans Lagudah for the
preparation of this report Genetic stocks and marker development
An important variable in the progress and
Section 3. Developing markers for the success of marker development is the reliability of the
stem rust resistance genes Sr33 and Sr45 phenotypic assessment. To ensure accurate phenotyping
(Contributed by P.K. Sambasivam, U.K. Bansal, for Sr33 and Sr45, we made use of genetic stocks where
H.S. Bariana and E.S. Lagudah) the respective genes were introduced into the wheat
variety Chinese Spring (CS) as a single chromosome
Background substitution line. The parental stocks for Sr33 designated
Diverse species from the Triticeae have been CS(1D5405) and Sr45 designated as CS(1D5406) were
explored in attempts to identify new sources of tested using Puccinia graminis f. sp. tritici (Pgt) pathotype
resistance to stem rust. Resistance sources from the 34-1,2,3,4,5,6,7,11 (Plant Breeding Institute[PBI] culture
diploid D genome progenitor, Aegilops tauschii, are no. 171) at the seedling stage, and pathotype 98-
Y
Xcdo673 Xcdo53 Xpsr129 Xpsr547 Xwg420 STSLr19 Xmwg2062 Xpsr680 Xpsr121 XPSY-1
T#1 +Lr19 +Y
1-7 + Lr19 +Y
1-23 +Lr19 +Y
1-22 +Lr19 +Y
1-49 - Lr19 +Y
1-32 - Lr19+Y
Sr25
1,2,3,5,6 (PBI culture no. 279) was used for adult plant by exploring the co-linearity of ESTs on wheat group
stem rust assessment in Australia. Seedling infection 1 with rice chromosome 5. None of the ESTs mapped
types were scored on a 0 – 4 scale (McIntosh et al. 1995) in the vicinity of Sr33 or Sr45, and all were located in
and adult plant assessments were made on 1 – 9 scale more proximal regions. Thus, the targeted Sr33 and Sr45
(Bariana et al. 2007). CS1D5405 (Sr33) and CS1D5406 regions show negligible co-linearity with the predicted
(Sr45) produced low infection types (IT) IT2= and IT;;1-, syntenic region of rice chromosome 5. Comparative maps
respectively, when tested against Pgt pathotype 34- of wheat and its relatives in the group1S chromosomal
1,2,3,4,5,6,7,11 at the seedling stage. The susceptible region were examined for additional markers; however,
parent Chinese Spring displayed a susceptible response there are very few markers in the Sr33 and Sr45 regions.
(IT3+). At the adult plant stage, resistant parents We have since isolated more wheat ESTs and through
produced a rust response of 4 (CS1D5406) and 5 to 6 RFLP analysis have identified markers closely linked to
(CS1D5405), whereas the susceptible parent produced Sr33 and Sr45. These markers will be tested on the high
a high field response of 8. A low resolution mapping resolution mapping family, and if shown to be tightly
family based on recombinant inbred lines (<100) linked, will provide entry points towards developing
from the crosses CS(1D5405)/CS and CS(1D5406)/CS markers suitable for breeding applications.
were phenotyped at the seedling and adult stages. A
perfect correlation between the seedling and adult rust Fig. 6 Low resolution genetic maps of the Sr33 and Sr45
responses was obtained. Using microsatellite markers regions in wheat chromosome 1DS (Sambasivam et al.
from 1DS, framework maps around the Sr33 and Sr45 2008)
regions were developed (Fig. 6). It was apparent that the
genetic map resolution was very low as several markers
immediately distal to Sr45 could not be separated by
recombination.
126 Developing and optimizing markers for stem rust resistance in wheat
Section 4. Developing diagnostic markers Sixteen wheat ESTs mapped to deletion bin 6AL8-
for Sr25 and Sr26 and pyramiding Sr25 and 0.90-1.00 were selected to design STS markers. None
Sr26 (Contributed by Sixin Liu and James A. of the 16 STS markers were co-dominant between
Anderson) lines with or without Sr26, but six markers specific to
chromosome 6AL amplified no PCR product from Eagle.
Materials and Methods We reasoned that multiplex PCR with the combination
Eight wheat lines including two lines of one 6AL-specific marker and Sr26-specific marker
with gene Sr25, ‘Wheatear’ and CIMMYT line Sr26#43 could be used to distinguish Sr26 homozygotes
C80.1/3*Batavia//2*Weebil, gene Sr26 line Eagle, and from heterozygotes. Because the 6AL-specific marker
five lines without gene Sr25 or Sr26, ‘Cranbrook’, ‘Weebil’, UMN6A-5 (Table 4) consistently worked well and
MN02072-7, MN03130-1-62 and MN03148, were used amplified a 320 bp fragment from lines without Sr26, we
to validate published markers for Sr25 and Sr26 (Table combined equal amounts of primer for marker UMN6A-5
4). The PCR protocols were the same as described by and Sr26#43 to genotype an F2 population segregating
Liu and Anderson (2003) with the exception of 400 nM for Sr26. For the heterozygous plants, the 320 bp allele
instead of 100 nM for each primer. The PCR products was weaker than the 207 bp allele amplified with primer
were separated on 3% agarose gels and visualized with Sr26#43. After doubling the primer concentration for
ethidium bromide under UV light. marker UMN6A-5, the 6A-specific allele and the Sr26-
Wheat ESTs mapped to deletion bin 6AL8- specific allele were amplified with similar intensities. Two
0.90-1.00 (Qi et al. 2004) were used to design STS F2 populations segregating for Sr26 were successfully
(sequence tagged site) markers with software Primer 3. genotyped with this optimized, multiplex PCR.
Chromosome 6AL-specific markers were identified with One of the goals of this project is to combine Sr25
aneuploid analysis. and Sr26 in the genetic background of four high yielding
CIMMYT lines that are well-adapted to Ug99-affected
Results and surrounding regions and already possess at least
Marker Gb (Prins et al. 2001) for gene Sr25 and moderate adult plant resistance to Ug99. In fall 2008, we
Sr26#43 (Mago et al. 2005) for gene Sr26 were validated made F1 hybrids between each of the CIMMYT lines and
with eight wheat lines. As expected, a faint 130 bp lines with Sr25 or Sr26. Complex F1’s will be made this
fragment was amplified with marker Gb in the two lines season and lines containing Sr2, Sr25 and Sr26 will be
with Sr25, Wheatear and C80.1/3*Batavia//2*Weebil. selected using co-dominant markers.
The other six lines without Sr25 did not amplify any
detectable fragment with these primers. Similarly, only Section 5. Mapping of new sources of race-specific
cultivar Eagle was positive for marker Sr26#43 and no resistance in CIMMYT-derived spring wheat
PCR product was observed for the other seven lines. (Contributed by Sridhar Bhavani and Ravi P. Singh)
Since co-dominant markers are needed to distinguish Summary: Genomic regions for three putative,
homozygous plants from heterozygous plants, we novel race-specific resistance genes, temporarily
decided to develop and test co-dominant markers for designated as SrA, SrB and SrC, present in three CIMMYT
genes Sr25 and Sr26. derived spring wheats were determined. Gene SrA
Ayala-Navarrete et al. (2007) developed STS markers mapped on 3DL (linked markers Xgwm52, Xgwm341), SrB
from wheat ESTs mapped to chromosome 7DL that on 3BS (Xgwm566, Wmc231) and SrC on 5DL (Xgwm292,
is homoeologous to the translocated segment of Th. Xgwm212).
elongatum containing Sr25 and Lr19. Among the six
markers tested on the eight wheat lines, BE404744 and Mapping of new sources of race-specific resistance in
BF145935 were co-dominant in marking Sr25. We tested CIMMYT-derived spring wheats
both markers with additional lines used to pyramid About 150 F3 or F4 lines derived from the crosses of
Sr25 and Sr26. Both markers were diagnostic for Sr25. susceptible parent PBW343 with three resistant parents,
However, marker BF145935 consistently worked well viz. ‘Milan/Sha7/3/Thb/CEP7780//Sha4/Lira/4/Sha4/Chil’,
and was easier to score. We successfully genotyped ‘Ning9415/3/Ures/Bow//Opata/4/Ningmai 7’ and ‘Chen/
five F2 populations segregating for Sr25 with marker Ae.Sq//2*Weaver/3/Oasis /5*Borl95’ suspected to carry
BF145935. Therefore, marker BF145935 is a robust co- novel race-specific resistance genes, were characterized
dominant marker for Sr25. for stem rust responses in the field at Njoro during
Dundas et al. (2007) reported that Sr26 is located in 2008 and in the USDA-ARS greenhouse by Dr. Y. Jin.
the extreme distal portion of the 6Ae#1 chromosome. The three resistant parents displayed infection types
2- or 2 in seedling tests and MR to MR-MS (moderately
128 Developing and optimizing markers for stem rust resistance in wheat
Fig. 9 Molecular markers linked to stem rust resistance Hare RA, Mcintosh RA (1979) Genetic and cytogenetic
gene SrC located on chromosome 5DL studies of durable adult-plant resistances in Hope
and related cultivars to wheat rusts. Zeitschrift Fur
Pflanzenzucht 83:350-367
Jin Y, Singh RP, Ward RW et al (2007) Characterisation
of seedling infection types and adult plant infection
responses of monogenic Sr gene lines to race TTKS of
Puccinia graminis f. sp. tritici. Plant Disease 91:1096-
1099
Klindworth DL, Miller JD, Jin Y, Xu SS (2007) Chromosomal
locations of genes for stem rust resistance in
monogenic lines derived from tetraploid wheat
accession ST464. Crop Sci 47:1441-1450
Knott DR (1968) Inheritance of resistance to stem rust
races 56 and 15B-IL (Can) in wheat varieties Hope and
H-44. Can J Genet Cytol 10:311-320
rust at the seedling stage and acceptable to moderate Liu SX, Anderson JA (2003) Marker assisted evaluation of
levels or resistance at the adult stage. The genomic Fusarium head blight resistant wheat germplasm. Crop
regions identified in this study (3DL, 3BS and 5DL) Sci 43:760-766
likely represent new genes and sources of resistance Mago R, Bariana HS, Dundas IS, Spielmeyer W et al (2005)
to Ug99. Selected F3 or F4 lines that were segregating Development of PCR markers for the selection of
for the above genes are being used in developing new wheat stem rust resistance genes Sr24 and Sr26 in
mapping populations in an attempt to remove maturity diverse wheat germplasm. Theor Appl Genet 111:496-
effects that caused difficulties in phenotyping stem rust 504
responses in the field in Kenya. These populations will McIntosh RA, Wellings CR, Park RF (1995) Wheat rusts, an
also be used for identifying tightly linked molecular atlas of resistance genes CSIRO, Melbourne, Australia
markers for use in marker-assisted breeding. McIntosh, RA, Yamazaki Y, Dubcovsky J, Rogers WJ et al
2008. Catalogue of gene symbols for wheat. Gene
References symbols. In: McIntosh RA (ed) http://wheat.pw.usda.
Ayala-Navarrete L, Bariana HS, Singh RP, Gibson JM et al gov/GG2/Triticum/wgc/2008/GeneSymbol.pdf.
(2007) Trigenomic chromosomes by recombination Pederson WL, Leath S (1988) Pyramiding major genes
of Thinopyrum intermedium and Th. ponticum for resistance to maintain residual effects. Annu Rev
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Ayliffe M, Singh R, Lagudah E (2008) Durable resistance Prins R, Groenewald JZ, Marais GF, Snape JW, Koebner
to wheat stem rust needed. Curr Opinion in Plant Biol RMD (2001) AFLP and STS tagging of I, a gene
11:187-192 conferring resistance to leaf rust in wheat. Theor Appl
Bariana HS, Miah H, Brown GN, Willey N, Lehmensiek A Genet 103:618-624
(2007) Molecular mapping of durable rust resistance Qi LL, Echalier B, Chao S, Lazo GR et al (2004) A
in wheat and its implication in breeding. In: Buck HT, chromosome bin map of 16,000 expressed sequence
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Chicaiza O, Khan IA, Zhang X, Brevis JC et al (2006) Identification of markers linkedwith stem rust
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Genetic map of diploid wheat, Triticum monococcum Press, Sydney, Australia
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Genetics 143:983-999 Current status, likely migration and strategies to
Dundas IS, Anugrahwati DR, Verlin DC, Park RF et al (2007) mitigate the threat to wheat production from race
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130 Developing and optimizing markers for stem rust resistance in wheat
19. Breeding for minor gene-based causes severe devastation of wheat crops, and is the
most feared disease in many countries where wheat is
adult plant resistance to stem rust in grown. Strong emphases on the identification of sources
of resistance to stem rust and on breeding resistant
wheat varieties were earlier adopted in the USA, Canada,
Australia and parts of Europe. Although the major
R.P. Singh1, J. Huerta-Espino2, S. Bhavani1, D. Singh3, epidemic of 1916 in the USA and Canada had already
P.K. Singh1, S.A. Herrera-Foessel1, P. Njau4, R. Wanyera4,
triggered extensive research on stem rust, efforts in
Y. Jin5
the US, Canada and Australia were intensified further
with epidemics during the following decades. Initially,
Abstract resistances present in some hexaploid wheats were
An attractive strategy for a long-term control of used, but the most successful control was achieved
Ug99 and other races of the wheat stem rust pathogen when H.K. Hayes at the University of Minnesota, and E.S.
is to develop and deploy varieties that have durable, or McFadden at South Dakota State University, transferred
race-nonspecific, adult plant resistance (APR) conferred stem rust resistance from the tetraploid ‘Iumillo’ durum
by multiple minor genes. Screening of spring wheat and ‘Yaroslav’ emmer, respectively, to bread wheat
materials in Kenya under high stem rust pressure, and resulting in the hexaploid varieties ‘Thatcher’ and ‘Hope’
characterization of seedling reactions to infection (Kolmer 2001). Although some race-specific resistance
in the greenhouse, resulted in the identification of genes are present in Hope and Thatcher, the most
a low frequency of high-yielding lines with high to effective components of resistance in both varieties
adequate levels of APR. Slow rusting APR gene Sr2 is were genes conferring adult plant resistance (APR).
present in these materials and is therefore an important Thatcher and Hope (and Hope sib ‘H44-24a’) and other
component of the complex adult-plant resistance, varieties derived from them, such as ‘Selkirk’ and ‘Chris’
commonly known as the “Sr2-complex”. Simple, that combined resistance genes from other sources
single-backcross and top (3-way) crosses followed by including Sr6 found in a single plant selection made by J.
a selected-bulk selection scheme are being used at McMurachy in 1930. ‘Kenya 58’ and other Kenyan varieties
CIMMYT to develop new high-yielding wheat materials carrying Sr6 were also used extensively in Australia by S.L.
with high levels of APR to stem and other rusts, as Macindoe, and in Mexico by N.E. Borlaug in addition to
well as additional necessary traits. We have adopted a Hope and Thatcher derivatives. Efforts to find a solution
shuttle breeding strategy using the Ciudad Obregon and to the stem rust problems facilitated global collaboration
Toluca breeding sites in Mexico and Njoro in Kenya. The amongst wheat scientists via the International Rust
segregating populations are selected at Njoro for two Nurseries to share, grow and evaluate wheat germplasm
generations before making final plant selections at Cd. in the quest of finding different sources of resistance
Obregon. Advanced lines retained in Mexico are then to stem rust. Resistant wheat germplasm developed at
phenotyped for stem rust response at Njoro, and grain Njoro, Kenya, through support from Canadian scientists
yield performance, end-use quality and other traits are in the 1960s and 1970s, contributed substantially to
determined in Mexico. The first products of the breeding international breeding efforts.
scheme are currently (2008-09 crop season) being The APR gene Sr2, transferred to Hope and H44-24a
characterized for yield potential and stem rust resistance from Yaroslav and possibly to ‘Khapstein’ from ‘Khapli’
at Cd. Obregon and Njoro, respectively. emmer by W.L. Waterhouse in Australia, conferred slow
rusting. Combinations of Sr2 with other unknown slow
Keywords rusting resistance genes possibly originating from
Triticum aestivum, Puccinia graminis tritici, Ug99, Thatcher and the Thatcher-derived Chris, commonly
shuttle breeding, durable resistance. known as the “Sr2-complex”, provided the foundation of
durable resistance to stem rust in germplasm from the
Introduction University of Minnesota, in the U.S.A, Sydney University
Stem, or black, rust of wheat, caused by Puccinia in Australia, and spring wheat germplasm developed
graminis Pers. f. sp. tritici Eriks. & E. Henn., periodically by N.E. Borlaug (McIntosh 1988; Rajaram et al. 1988). Sr2
can be detected through its complete linkage with the
1
CIMMYT, Apdo. Postal 6-641, 06600, Mexico, DF, Mexico; 2INIFAP-CEVAMEX, pseudo-black chaff (PBC) phenotype; however, excessive
Apdo. Postal 10, 56230, Chapingo, Mexico; 3CIMMYT, Nairobi, Kenya; 4Kenya
Agricultural Research Institute- Njoro Plant Breeding Research Center (KARI- expression of PBC in certain environments sometimes
NPBRC), P.O. Njoro, Kenya; 5USDA-ARS, Cereal Disease Laboratory, St. Paul 55108, leads to its elimination in breeding programs. Under
MN, USA
E-mail: R.singh@cgiar.org the same environmental conditions, negligible to
132 Breeding for minor gene-based adult plant resistance to stem rust in wheat
Table 1 Stem rust resistance of entries included in 1st, 2nd and 3rd SRRSN (Stem Rust Resistance Screening Nursery)
1st SRRSN 2nd SRRSN 3rd SRRSN
Resistance category
Number % Number % Number %
Adult-plant1:
R (5-10% severity) 4 4 0 0 3 3
R-MR (15-20% severity) 19 18 26 20 33 32
MR (30% severity) 6 6 22 17 21 20
MR-MS (40% severity) 2 2 15 12 0 0
MS (50-60% severity) 0 0 17 13 0 0
S (70-100% severity) 0 0 4 3 0 0
Race-specific:
Sr24 39 38 0 0 0 0
Sr25 17 17 0 0 11 11
Sr36 (+Sr24) 0 0 0 0 4 4
Sr1A.1R (+Sr24) 2 2
SrTmp 0 0 25 20 6 6
SrSynt 4 4 8 6 5 5
SrSha7 9 9 8 6 4 5
SrND643 0 0 0 0 12 12
SrUnknown 1 1 3 2 5 5
Total 103 128 104
1
Adult plant resistance categories include lines that were susceptible in seedling greenhouse tests and with the
highest rating during multiple years/seasons (2005-07) when susceptible entries were dead. Ratings based on the
modified Cobb Scale as described in Roelfs et al. (1992)
Shuttle breeding to develop high-yielding and Kenya halve the number of years required to
wheat with APR to stem rust generate and test advanced breeding lines. The “single-
Because a large proportion of high-yielding spring backcross, selected-bulk” breeding approach (Singh
wheat varieties and germplasms do not carry effective and Trethowan 2007) is being applied for transferring
race-specific stem rust resistances to race Ug99, the multiple minor genes to adapted backgrounds. Simple
availability of genotypes with moderate to high levels and three-way crosses, where one or more parents carry
of APR provide opportunities to reconstitute high levels adult-plant resistance, are being used to breed new
of adult plant resistance in newer hybrid populations. high-yielding, near-immune wheat materials. The flow
In the absence of molecular markers for adult plant of breeding materials in the “Mexico-Kenya Shuttle” is
resistance genes and the absence of Ug99 in Mexico, a described in Table 5.
shuttle breeding scheme between Mexican field sites In the single-backcross approach, we cross the
(Ciudad Obregon in northwestern Mexico during winter, resistance sources with adapted high yielding wheats.
and Toluca or El Batan in the highlands near Mexico City A single backcross is made with the recurrent parent to
during summer) and Njoro, Kenya, was initiated in 2006 obtain 350-400 BC1 seeds. The BC1 plants are selected
to build adult-plant resistances in modern semidwarf for desired agronomic features and resistance to leaf
wheats. Two crop seasons per year in both Mexico rust and stripe rust, and harvested as bulk in Mexico.
134 Breeding for minor gene-based adult plant resistance to stem rust in wheat
Table 3 Some wheat lines with adult plant resistance (APR) to stem rust in the 2nd Stem Rust Resistance Screening
Nursery distributed by CIMMYT during 2007-08 with field ratings to stem rust in three crop seasons at Njoro, Kenya
Stem rust rating1
Entry No. Cross 2007 Off- 2007 Main- 2008 Main-
season season season
6026 Barbet1*2/Kiritati 30 MSS 30 MSS 30 M
6036 Galvez/Weebill1 10 M 15 M 30 M
6044 Kiritati//PBW65/2*Seri.1B 20 M 20 M 40 M
6048 Kiritati//Seri/Rayon 20 M 15 M 30 M
6049 Kiritati/4/Seri.1B*2/3/Kauz*2/Bow//Kauz 30 MSS 10 M 30 M
6057 Pfau/Weaver*2//Chapio 20 M 20 M 30 MSS
6059 Pfau/Weaver*2//Pavon 7S3 20 M 30 MSS 20 MSS
6070 Thelin/2*Waxwing 15 M 10 MS 30 M
6077 Pauraque 20 M 30 MSS 40 M
6085 Grackle 30 M 30 MSS 40 M
60101 Becard 20 M 15 M 30 M
6113 Inqualab 91*2/Kukuna - - 30 M
6114 Kiritati 15 M - 15 M
6118 Pavon F76 10 M - 15 M
6119 PBW343*2/Kukuna - - 30 M
6122 Pfau/Weaver//Kiritati 10 MSS - 30 M
6123 Pvn//Car422/Ana/5/Bow/Crow//Buc/Pvn/3/Yr/4/Trap#1 5 MS - 30 M
6125 Kingbird 5 MSS - 5M
6126 Temporalers M87*2/Chos - - 5M
6130 Tepoca+LR34/2*Borlaug F95 - - 20 MSS
6137 Cacuke (Susceptible Check) 100 S 100 S(N) 100 S(N)
1
The stem rust rating has two components: disease severity based on the modified Cobb Scale and host reaction,
both described in Roelfs et al. (1992)
Performance of high-yielding wheats with and seed was multiplied at El Batan for distribution. The
APR to stem rust and seed multiplication EBWYT trial has space for 30 entries (including checks)
The scheme presented in Table 5 delivers material with 3 replicates arranged in an alpha-lattice design. The
for CIMMYT international yield trials and screening 1stEBWYT was used initially to distribute new lines with
nurseries to co-operators in the 7th year. However, the APR to leaf rust and stripe rust, but 2nd and 3rd EBWYT also
best Ug99 resistant advanced lines were delivered include lines with Ug99 resistance.
to high risk countries during the current and past Eleven entries in the 2nd EBWYT had APR in the
two cropping seasons for growing in the 6th year after following categories (based on at least two seasons
crosses were made, through a nursery called the Elite of data from Njoro and final scores recorded when
Bread Wheat Trial (EBWYT). To achieve this we used the susceptible check was dead (100% severity): R-MR
one year of grain yield data combined with agronomic (15-20% severity) = 4, MR (30% severity) = 4, MR-MS
characteristics, quality and disease resistance information (40% severity) = 3. The resistance of two entries each
was based on Sr25 and SrTmp, a temporary designation Ug99-resistant entries were also chosen by Egypt and
for an undesignated gene derived from ‘Triumph’. Afghanistan. Iran has initiated multiplication of three
Similarly, 21 entries in the 3rd EBWYT had APR in the entries from the 2nd EBWYT on its own initiative. Countries
following categories: R (<10% severity) = 1, R-MR = 15, participating in the seed multiplication program will
MR = 5. Resistance in four entries was based on Sr25, conduct further yield performance evaluations while
two on SrTmp and one entry possesses a resistance simultaneously multiplying the seed. This approach is
gene from ‘HUW234+Lr34’, temporarily designated as likely to result in the release of at least one variety in each
SrHuw234. Using the 2008 stem rust data from Njoro, a of a number of countries that should have sufficient seed
season marked by the most severe stem rust seen since for rapid seed multiplication after official release.
2005, the 21 entries with APR in the 4th EBWYT were
classified: R-MR (30% severity) = 6, MR = 40% severity, Conclusions
and MR-MS (50% severity) = 4. In addition, four entries Significant progress has been made in identifying
carry Sr25 and one possesses SrHuw234. The two best and utilizing APR to stem rust in CIMMYT-derived spring
performing entries from the 2nd and 3rd EBWYT were bread wheats since the launch of Borlaug Global Rust
included as CIMMYT checks in the 4th EBWYT and both Initiative and initiation of a screening program at Njoro,
carry APR to stem rust. Kenya, in 2005. The Mexico-Kenya shuttle breeding
Eleven wheat lines (Table 6) were selected based strategy currently used should result in the accumulation
on grain yield performance data in the 2nd and/or 3rd of adequate APR to stem rust in modern high-yielding
EBWYT trials from various countries, and were included wheat germplasm with higher yield potential than
in a seed production program implemented during the current popular varieties. The initiation of multiplication
2008-09 crop season. Seed was provided from Mexico. and release of Ug99-resistant materials, especially those
Eight of the 11 entries carry APR to stem rust. Additional with APR, in various countries should reduce the Ug99
136 Breeding for minor gene-based adult plant resistance to stem rust in wheat
threat and help to achieve durable resistance. The Acknowledgements
identification of molecular markers, tightly linked to We acknowledge financial resources from the
the minor genes contributing to APR, should improve Durable Rust Resistant Wheat Project led by Cornell
breeding efficiency especially when it is not possible to University and supported by the Bill & Melinda Gates
shuttle segregating materials to field sites exposed to Foundation; ICAR- India; USDA-ARS and USAID, USA;
Ug99 epidemics. GRDC and ACIAR, Australia, Agrovegetal-Spain; the
Northwestern Mexican Farmer Association (Patronato)
and CONFUPRO, Mexico, SDC, Switzerland; and our
institutions.
Table 5 Flow of breeding materials in the Mexico-Kenya shuttle scheme, utilizing two crop seasons per year, for
developing high-yielding wheat germplasm combining adult plant resistance to stem rust with other traits
Year Locations1 Activities
name Cross category1 type Bangla-desh Nepal Pakistan Turkey Afghanistan Egypt Ethiopia
Seri.1B*2/3/Kauz*2/Bow//
Chonte#1 APR: R-MR Normal 300 100 50 25 100
Kauz/4/PBW343*2/Kukuna
Babax/Lr42//Babax*2/3/
Quaiu#1 SrTmp Normal 300 50
Vivitsi
Babax/Lr42//Babax*2/3/
Quaiu#2 SrTmp Normal 100 100 100 100
Vivitsi
Waxwing*2/4/Sni/Trap#1/3/
Breeding for minor gene-based adult plant resistance to stem rust in wheat
Pauraque#1 APR: R-MR Early 100 100
Kauz*2/Trap//Kauz
1
Resistance category is based on Ug99 response data obtained for at least two seasons (2006 and 2007) at Njoro, Kenya. APR (adult-plant resistance) categories
are R-MR = resistant to moderately resistant (15-20% stem rust severity) and MR = moderately resistant (30% disease) when susceptible checks were dead
following 100% severity
References Roelfs AP, Singh RP, Saari EE (1992) Rust diseases of wheat:
Knott DR (1982) Multigenic inheritance of stem rust concepts and methods of disease management.
resistance in wheat. Crop Sci 22:393-99 CIMMYT, Mexico, D.F.
Knott DR (1988) Using polygenic resistance to breed Singh RP, Trethowan R (2007) Breeding spring bread
for stem rust resistance in wheat. In: Simmonds NW, wheat for irrigated and rainfed production systems of
Rajaram S (eds) Breeding strategies for resistance to developing world. In: Kang M, Priyadarshan PM (eds)
the rust of wheat. CIMMYT, Mexico, D.F., pp 39-47 Breeding major food staples. Blackwell Publishing, U.K.,
Kolmer JA (2001) Early research on the genetics of pp 109-140
Puccinia graminis stem rust resistance in wheat in Singh RP, Huerta-Espino J, Rajaram S (2000) Achieving
Canada and the United States. In: Peterson PD (ed) near-immunity to leaf and stripe rusts in wheat
Stem rust of wheat: from ancient enemy to modern by combining slow rusting resistance genes. Acta
foe. APS Press, St. Paul, MN, pp 51-82 Phytopathlogica Hungarica 35:133-139
McIntosh RA (1988) The role of specific genes in breeding Singh RP, William HM, Huerta-Espino J, Rosewarne G
for durable stem rust resistance in wheat and triticale. (2004) Wheat rust in Asia: meeting the challenges with
In: Simmonds NW, Rajaram S (eds) Breeding strategies old and new technologies. In: New directions for a
for resistance to the rusts of wheat. CIMMYT, Mexico, diverse planet: Proc 4th Int Crop Sci Cong, Brisbane,
D.F., pp 1-9 Australia. Available via http://www.cropscience.org.au./
Rajaram S, Singh RP, Torres E (1988) Current CIMMYT icsc2004/symposia/3/7/141_singhrp.htm.
approaches in breeding wheat for rust resistance. In: Accessed 1 Feb 2009
Simmonds NW, Rajaram S (eds) Breeding strategies for
resistance to the rust of wheat. CIMMYT, Mexico, D.F.,
pp 101-118
1
USDA-ARS Plant Science Research Unit, Department of Plant Pathology; North
Carolina State University, Box 7616, Raleigh, NC 27695-7616, USA; 2International
Center for Agricultural Research in the Dry Areas (ICARDA), PO Box 5466,
Aleppo, Syria
DZ 04-1167/Dz-129/Yemen/Cit’s’/Pls’s’/3/Taganrog B.B/4/5/
9 MEGENAGNA (DZ2023) DZARC 2004
Chen’s’/RCHI//Hui’s’/BHA
13 LASTE TOB-2 (Dgo /II /3/ Rutt s// rg s / mexi s) CIMMYT 2002
Cocorit71/3/Gerardo//61-130/Gll ‘s’/4/Boohai/Hora//
14 LELISSO (DZ-1605) DZARC 2002
Gerardo/3/Boohai
18 ROBE (DZ-1640) Hora/ cit s // Jo s/ Gs s /3/ some s /4/ hora Respinegroll DZARC 1998/99
19 ASASA (DZ-2085) CHo S T arus// yav s /3/ Fg s /4/ Fg s /cr s /5/ DZARC 1997
21 QUAMI CD 75533-A (Fg s /Crs/5/ 5/ Fg s/ Dom s/6/ Huj S, CD75533-a) CIMMYT 1996
Fig. 1 Frequencies (%) of durum wheat entries (from international nurseries) in stem rust severity groups
for 2005 - 2007
90
80 2005 (n=700)
2006 (n=803)
70 2007 (n=1220)
60
Frequency (%)
50
40
30
20
10
0
0 1 5 10 - 15 20 - 25 >30
Severity classes
152 Strategies to combat race Ug99 and control other wheat rusts in India
23. Strategies and progress towards slow rusting into Chinese varieties. Three way crosses
(CIMMYT/Sichuan//Sichuan genotypes), and selections
the development of rust resistant made in Mexico from F1 to F4, and continued after F4
in Sichuan, proved to be effective in combining yield
wheat varieties in China potential, slow rusting, and adaptation to the Sichuan
environment. More than 10 varieties were released and
Zhonghu He1,2, Xianchun Xia1 Chuanmai 42, developed from synthetic wheat, is a
leading variety in Sichuan.
Yellow (stripe) rust is a major wheat disease in
Stem rust was basically controlled in China after
northwestern and southwestern China. Resistance based
the 1960s, however, race Ug99 could be a potential
on both major and minor genes is employed at present.
threat to Chinese wheat production. Therefore, a
Understanding resistance genes in current varieties,
joint China-CIMMYT collaborative program has been
and development of molecular markers is crucial for
established. The major activities include pathogen
improving breeding efficiency. Gene postulation and
monitoring, introduction and evaluation of new varieties
adult stage testing indicated that resistance genes Yr2,
with rust resistance, testing Chinese varieties in Kenya,
Yr3a, Yr4a, Yr6, Yr7, Yr9, Yr26, Yr27, and YrSD, either singly
development and application of molecular markers,
or in combination were identified, with Yr9, Yr26 and
breeding of stem rust resistant varieties, and training.
YrZH84 being the most predominant genes. However,
The ten most representative varieties or promising new
only Yr5, Yr10, Yr15, Yr26 (or Yr24), and YrZH84 are still
lines from five provinces (Hebei, Henan, Shandong,
effective and can be used in developing new varieties.
Sichuan, and Heilongjiang) will be used for marker-
Thirty-three varieties or advanced lines showed slow
assisted incorporation of race-specific resistance
rusting resistance and they could be used as crossing
through limited backcrossing and field selection.
parents in breeding programs. Various markers for Yr5,
More than 700 Chinese varieties and lines were tested
Yr10 and Yr26 have been developed, and molecular
in Kenya, and 10 conferred medium or high levels of
mapping of YrCH42 and YrZH84 in Chinese wheat
resistance. They included Jimai 20 from Shandong, six
varieties has also been undertaken. Yr24, Yr26 and
lines (ELT 102, ZL-21, KD-9, XKD-21-1, XKD27-4, and Nei
YrCH42 are the same gene, whereas YrZH84 is new.
2836 from Sichuan, Yunxuan 11-12 and Jingmai 8 from
Pyramiding of YrZH84 and YrCH42, each present in
Yunnan, 656 and 12-1 from Xinjiang, Nongpin 5 from
varieties with outstanding agronomic characters, can
Inner Mongolia, Longfu 02-0667 from Heilongjiang, and
be used to develop new varieties. A shuttle breeding
Ningchun 18 and Ningchun 37 from Ningxia. Jimai 20 is
program has been established between CIMMYT Mexico,
a leading variety in north China, grown on more than 1
and Sichuan province to integrate minor gene-based
million hectares in 2008.
1
Institute of Crop Science, and 2CIMMYT China Office, Chinese Academy of
Agricultural Sciences (CAAS), No 12 Zhongguancun South Street, Beijing
100081, China
E-mail: zhhe@public3.bta.net.cn
M. Esmaeilzadeh Moghaddam1, M.R. Jalal Kamali2, M. Table 1 Harvested small grain cereals areas (000 ha) in
Aghaee1, F. Afshari1, M. Roustaii3 the 2005-2006 cropping season
Wheat Barley Total
Abstract
Wheat is the major field crop in Iran. The total wheat Irrigated 2634 607 3241
area exceeds more than 6.95 million ha from which 2.63
million ha (38%) are irrigated, with an average grain
Rainfed 4317 1052 5369
yield of about 3,870 kg ha-1, and about 4.32 million ha
(62%) are rainfed with an average grain yield of 1,100
Total 6951 1659 8610
kg ha-1. In normal years about 70% of wheat production
is produced under irrigation. Winter, facultative and
Source: Statistics and Information Technology Office,
spring types are grown in different agro-climatic regions.
Ministry of Jihad-e-Agriculture, Iran, 2007
The temperate agro-climatic zone is the most favorable
area for wheat production with the highest grain yields
The total harvested area in Iran has not changed
recorded at Kangavar in Kermanshah province (about
significantly over the period 1992-2007; however,
14 tonnes ha-1) and in Daryoun in Fars Province (>12
it dropped to its lowest level of about 5.0 m ha in
tonnes ha-1). Biotic (e.g. YR, LR, SR, Septoria, FHB) and
1998-1999 due mainly to severe drought conditions
abiotic (drought, heat, cold, salinity) stresses are among
and consequently, there was a large decrease in the
the major limiting factors for wheat production. Yellow
harvested areas of dryland wheat (Fig. 1).
(stripe) rust, leaf rust and stem rust occur in different
parts of the country; however, yellow rust remains
Fig.1 Harvested wheat areas in Iran for the period of
the major wheat disease in more favorable years. In
1992-2007
1993, about 1.5 million tonnes of wheat were lost due
to yellow rust. Fusarium head blight and Septoria leaf 8.0 Harvested Areas
blotch are becoming more serious in the Caspian Sea
7.0
regions as well as in south-west Iran. Leaf rust and stem
rust appear late in the season, but there are no reports 6.0
Million Hectares
on crop losses due to these diseases over the last three 5.0
decades. In 2007, stem rust race Ug99 was reported from
Broujerd and Hamadan in western Iran. However, this 4.0
race was not found in 2008. 3.0
2.0 Irrigated
Keywords
Dryland
Triticum aestivum, Puccinia striiformis, Puccinia. 1.0
Total
graminis, Puccinia. triticina, grain yield, irrigated wheat, 0.0
dryland wheat
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
Introduction
Iran is a vast and diverse country. The total land area (After Jalal Kamali et al. 2007)
of Iran is 164.8 million hectares (m ha), much of which is
mountainous. About 18.5 m ha are used for agriculture The average wheat yield peaked to about 2,500
as follows: kg ha-1 in 2005, but decreased to about 2,150 kg ha-1 in
• 6.0 m ha = 32.4%, Irrigated 2007 due to drought conditions (Fig. 2). The average
• 6.0 m ha = 32.4%, Dryland yield of irrigated wheat was 3,870 kg ha-1 whereas the
• 4.5 m ha = 24.4%, Fallow average grain yield for dryland wheat was 1,100 kg ha-1.
• 2.0 m ha = 10.8%, Horticultural crops Clearly, the average grain yields in the country are more
influenced by the average yield of the irrigated areas,
1
Seed and Plant Improvement Institute (SPII), Karaj, Iran; 2CIMMYT-Iran, SPII, which increased over the period 1992-2007, particularly
Karaj, Iran; 3Dryland Agriculture Research Institute, Maragheh, Iran 2003-2007 (Fig. 2).
E-mail: Mohsen_esma@yahoo.com
2000
head blight and Septoria are becoming increasingly
1500 serious diseases in the Caspian Sea region as well as in
1000 southwest Iran. Leaf rust and stem rust appear late in the
500 season and there are no recent reports on crop losses
0 due to these diseases.
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
The stem rust situation in Iran
(After Jalal Kamali et al. 2007) Stem rust was reported for the first time in Iran by
Esfandiari (1947). Epidemics of stem rust in the Caspian
The total wheat production reached about 15.0 mt Sea region in the northern and southern regions of
in 2007 (Fig. 3). In normal years about 70% of wheat Iran in 1975 and 1976 were reported by Bamdadian
production is produced in irrigated wheat areas. The 2000 and Torabi (1978). In 1976, an epidemic of stem rust in
harvest was the lowest (about 8.0 mt) recorded for the the southern part of Iran caused 100% crop losses in
period 1992-2007 (Fig. 3). Reductions are also evident for landraces (Bamdadian and Torabi 1978). The first stem
1999 and 2001, again reflecting the effects of drought. rust race analysis in Iran was reported by Sharif et al.
Total wheat production increased over the period 2002- (1970). Race analyses and responses of cultivars and
2007 because of more favorable growing conditions and a advanced breeding lines were carried out in 1994 and
steady increase in irrigated wheat yields (Fig. 3). 1995 by Nasrollahi et al. (2001). Stem rust was controlled
since 1976 by growing CIMMYT germplasm.
Fig. 3 Wheat production in Iran in the period of 1992- In 2007, the presence of race Ug99 was officially
2007 reported from Broujerd and Hamadan. Race analyses of
samples collected from Borujerd, Hamedan, Poldokhtar
16.0 Production and Kelardasht in 2007 and a race collected from
Irrigated Borujerd in 1997 were conducted using differentials
14.0 Dryland carrying stem rust (Sr) resistance genes plus several
12.0 Total additional wheat genotypes. Isolates from samples
collected at Borujerd and Hamedan in 2007 produced
Million Tonnes
10.0
high infection types (IT 33+ - 4) on differential lines
8.0 (Nazari et al. 2008). These are facultative and winter
6.0 wheat growing areas. The results were later confirmed
using differential lines. There was no evidence of this
4.0
race in 2008.
2.0 International monitoring has implicated the
0.0 progressive migration of race Ug99 from Africa to Iran.
Field evaluations of the responses of Iranian wheat
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
160 Breeding strategies for developing wheat cultivars resistant to rust diseases in Egypt and progress in breeding for resistance to stem rust race UG99
breeding program selected adapted promising lines Mutli-location screening in hotspots locally and
with high yielding ability and resistance to yellow rust. regionally accelerated the development resistant
Three semidwarf cultivars resistant to yellow rust were cultivars, but selection under epidemic situations
released in 1972 and 1973 (Mexipak 69, Super X, and identified sources of resistance to new pathotypes.
Chenab 70). However, these cultivars were not accepted Pathotyping studies and rust surveillance in the last
by farmers because of their susceptibility to leaf rust, 15 years indicated that the most frequent P. striiformis
short stature and low straw yield, and red grain color. pathotypes in Egypt were OEO, OE64, 2EO, 4E2, 4E148,
Breeders crossed resistant lines together to develop new 6E134, 70E20, 70E134, and 230E150. The most effective
cultivars with more genes for resistance and succeeded genes for resistance to yellow rust at the seedling stage
in releasing new cultivars resistant to yellow rust from are Yr1, 4, SP, SD and 9, whereas effective genes for
1976 - 1986, i.e. Sakha 8, Giza 157, Giza 158, Giza 160, resistance at the adult stage are Yr5, 8, 10, 15, 1 and SP.
Giza 162, Giza 163, Giza 164, Sakha 61, and Sakha 69.
The last two cultivars are sister lines and had a high level Breeding for stem rust resistance
of resistance to yellow rust. By 1989 all had become Stem rust is one of the most destructive diseases of
susceptible to yellow rust except Sakha 61, Sakha 69, wheat. During the first half of the 20th century, stem rust
and Giza 164. Wheat cultivar Sakha 69 was widely was the major disease (1930s – 1940s) and most wheat
adapted all over Egypt and occupied 70% of the planted cultivars were susceptible until a source of resistance
wheat area. Giza 164 (Veery 5) was a heat tolerant was introduced from Kenya. The first stem rust resistant
cultivar and was preferred by farmers in southern Egypt, cultivar, released in 1947, was Giza 139 (Hindi 90/ Kenya
while Sakha 61 was limited to the north Delta. The B 256) followed by many others. In the early 1970s,
Wheat Research Program had to release new cultivars semidwarf cultivars and lines introduced from CIMMYT
with new sources of resistance. Screening for resistance had different genes for resistance. The new cultivars
was practiced in the north, east and west Delta as were resistant, and stem rust was successfully controlled
hotspots for yellow rust. In addition, new cultivars and for 50 years due to following reasons:
promising lines were tested for resistance to yellow rust 1. Widespread use of resistant cultivars.
in Ethiopia and Yemen (NVRSRP project). Although a 2. Early maturing cultivars.
group of cultivars resistant to yellow rust was released 3. Pathogen stability and no aggressive races.
from 1991 to 1998, the new Yr9-virulent race infected all 4. Absence of the alternate host.
wheat cultivars in Egypt in 1995, except Sakha 61 (Inia
RL 4220// 7C/ Yr “s”), which showed a very high level of Some semidwarf cultivars became susceptible to
resistance. stem rust and were replaced by resistant cultivars. The
The dramatic epiphytotic of 1995 saw most of the most frequent races in Egypt are 11, 15, 9, 17, 39, 24, 34,
commercial cultivars, especially Giza 163, suffering 19, and 53; and the most effective genes are Sr5, 26, 27,
high losses in the north Delta (El-Daoudi et al. 1996). 36, 9e, Gt+, 29, and 30.
This became a dangerous time in Egypt, and maybe in Singh et al. (2006) stated that the importance of
the world, because it included wheat-growing areas stem rust was declining worldwide with the deployment
in the middle, near and far eastern zones, and led to of various alien resistance genes such as Sr24, 26,
high losses in all areas (Abu-Naga et al. 1997). Infection 31 and 38. Translocations carrying these genes also
was very severe especially in the north and east Delta carried additional genes that conferred resistance to
and losses were up to 50% in some regions. Many other diseases such as leaf rust, yellow rust or powdery
promising lines in the breeding program showed high mildew. However, a new stem rust race virulent to
levels of resistance in 1997 and 1998. Screening under resistance gene Sr31 was detected in Uganda in 1999
high levels of infection resulted in the release of four (Pretorius et al. 2000) and named Ug99. Later, Wanyera
resistant cultivars in 1999. These cultivars were Gemmiza et al. (2006) designated this race as TTKS using the North
7 (CMH74A.630/5X//Seri 82/3/Agent), Gemmiza 9 (Ald American nomenclature system. This race was virulent
“S” Huac “S”//CMH74A.630/ 5X), Giza 168 (Mil/ Buc// Seri) not only to Sr31, but also for most of the genes of wheat
and Sakha 93 (Sakha 92 TR 810328). Many promising origin, as well as Sr38 (Singh et al. 2006). Wanyera et
lines were selected under epidemic situations and al. (2006) reported that during 2003 and 2004, the
released in 2004 (Sakha 94, Gemmiza 10, Sids 12). All majority of current Kenyan cultivars and a large portion
susceptible cultivars were eliminated, especially Sakha of CIMMYT wheat germplasm with gene Sr31 planted
69, which occupied more than 70% of the wheat area, in Kenya were susceptible. They found also that the
and resistant cultivars now dominate the entire region. virulence pattern of the isolate collected in 1999 from
Uganda was identical to those in Kenya. Their results
Table 2 Egyptian materials tested for resistance to stem rust in hotspot locations. B, bread wheat; D; durum wheat
No. Year Country No. lines No. resistant entries
1 June 2005 Ethiopia 96 Escape
2 September 2006 Kenya 149 3 “B”
3 October 2006 Ethiopia 189 10 “D”
4 March 2008 Kenya 72D +156B 25 (BW)
5 March 2008 Ethiopia 180 -
6 October 2008 Kenya 163 -
7 November 2008 Yemen 163
162 Breeding strategies for developing wheat cultivars resistant to rust diseases in Egypt and progress in breeding for resistance to stem rust race UG99
Table 3 Average grain yield of eight genotypes selected over six sites in 2007/08
Grain yield
Entry Origin Pedigree
t/ha
Kenya for three years. The first, the 2nd Elite Bread Wheat yielding in Mexico, Egypt, India and Iran. In addition,
Yield Trial (2nd EBWYT) consisting of 29 genotypes and a these four lines were highly resistant to stem rust in
local check, was sown in a yield trial at four sites (north, Kenya. Three of the lines are under seed multiplication in
west, middle, and south Delta). The growing season in Egypt along with selected lines from the previous season.
2006/07 was characterized by high infection with all
three rusts, enabling selection on the basis of resistance Molecular markers to identify resistance
and grain yield. Five genotypes higher yielding by 500 genes
– 1,290 kg/ha than the local check, and resistant to all Identifying genes with markers is the key to durable
three rusts, were selected. resistance whereby breeders can combine more genes
The second nursery, the 1st Stem Rust Resistance into a single genotype. One of the objectives of the
Screening Nursery (1st SRRSN), contained 108 lines. Wheat Research Program is to identify or validate
Thirteen lines resistant to Ug99 in Kenya and to all three markers in Egyptian wheat populations. A cooperation
rusts in Egypt were higher yielding than the local check protocol between the Agricultural Genetic Engineering
and were selected for further assessment. Research Institute and Wheat Research Program was
In the 2007/08 growing season the selected wheat initiated for that purpose. In addition, two researchers
lines (five from 2nd EBWYT and 13 from 1st SRRSN) were from wheat breeding and pathology worked in ICARDA’s
evaluated with four local cultivars in two yield trails at six biotechnology laboratory to identify stem rust resistance
sites representing different ecological zones. Eight were genes in 30 wheat genotypes from Egypt during
selected for superiority in grain yield of 179 – 1,680 kg/ February, 2009. The materials included nine local bread
ha across the six sites (Table 3). wheat cultivars, 15 promising bread wheat lines and six
Exotic materials 2007/ 2008 Two yield trials, received local durum cultivars. They used ten specific primers
from CIMMYT in 2007/2008, included bread wheat linked to stem rust genes (SSR) and the results indicated
genotypes tested for resistance to Ug99 in Kenya for that all tested materials do not have Sr36, Sr46, Sr25, Sr39,
three years. The 29 entries in the Third Elite Bread Wheat Sr26 or Sr24. All, except five promising lines did not have
Yield Trial (3rd EBWYT) resistant to Ug99 were planted Sr31. The results suggested that Sr22 might be present
in two sites at north and west Delta, together with the in two promising lines and two durum cultivars. Sr2 was
2nd Stem Rust Resistance Screening Nursery (2nd SRRSN) found in all local cultivars, except Gemmiza 9, Sids 12 and
including 137 genotypes. Thirteen genotypes were two promising lines. This work will be continued in the
selected from the 3rd EBWYT for superiority in grain biotechnology laboratory in the Wheat Research Program
yield over the local check. However, the selections were using other specific primers to detect more Sr genes.
based on yielding ability only, because environmental
conditions did not allow development of rusts. Similarly, Doubled Haploids
12 promising lines from the 3rd SRRSN were selected for The biotechnology laboratory in the Wheat
high yielding ability. Selected lines are being evaluated Research Program started anther culture research to
at six sites for yield potential in 2008/09. Results for the produce doubled haploid genotypes resistant to Ug99.
3rd EBWYT were collected by CIMMYT from different Nine crosses between five CIMMYT genotypes resistant
countries and four of these lines proved to be high Ug99 and two Egyptian cultivars were made in 2007/08.
164 Breeding strategies for developing wheat cultivars resistant to rust diseases in Egypt and progress in breeding for resistance to stem rust race UG99
26. Combating stem rust to protect especially stem rust and stripe (yellow) rust whereas
the abiotic constraints include drought stress and soil
wheat crops in Kenya acidity. Over the last six years, new races of the stem
rust pathogen (Puccinia graminis tritici) have been
P. Njau1, R. Wanyera1, M. Gethi1, G. Kamau1, R.P. Singh2, the most serious production problem. These races
D. Singh3, Y. Jin4 include Ug99 (Pretorius et al. 2000), or TTKSK based on
North American nomenclature system, and its Sr24-
Abstract virulent variant TTKST (Jin et al. 2007). They are able to
Wheat is the second most important cereal in Kenya overcome resistances of all the commercial varieties
after maize. Both biotic (stem rust, yellow rust, Septoria grown in the country, and yield losses of up to 90%
tritici blotch) and abiotic (drought, acidic soils) factors occurred on some farms in the Narok district.
are important production constraints faced by wheat Breeding for resistance is cumbersome due to the
producers in Kenya. Stem rust race Ug99 has become rapid evolution, survival and build-up of new variant
number one problem during the last six years. Kenya races. Use of fungicides as a control measure is being
Agricultural Research Institute (KARI) - Njoro is involved practiced to fight the disease. Farmers have been trained
in the screening of international wheat germplasm in during field days, with shows and demonstrations
the search for new sources of resistance. In addition to on how to identify the disease at early stages of an
the international screening, KARI identified three wheat epidemic, and on which fungicides to apply. Farmers
lines that are high yielding and resistant to race Ug99 have also been encouraged to grow varieties such as
and its derivatives. These lines are at the final year of Njoro BW2 which are moderately resistant to Ug99.
testing before official release to farmers. This report Breeding for resistance is the most cost-effective
discusses the performance of Kenyan varieties against strategy for control of stem rust. KARI is involved in
Ug99 and its variants, progress on identification of new the screening of international wheat germplasm in the
sources of resistance, results of national advanced yield search for, and breeding with, new sources of resistance
trials, and proposed breeding strategies for incorporating at Njoro. The center is 200 km west of the Kenyan capital,
resistance genes into commercial varieties. Nairobi, in Nakuru district, of Rift valley province. It is
located 2,185m above sea level at latitude 0020’S and
Keywords longitude 35056’E. The average rainfall is 933 mm with
Triticum aestivum, Puccinia graminis tritici, durable average daily minimum and maximum temperatures
resistance of 100 (night) and 230 (day). The center has a national
mandate for small grain cereal research which includes
Introduction wheat, barley and triticale.
In Kenya, wheat (Triticum aestivum) is the second
most important cereal after maize. It is grown in the Evaluation of Kenyan varieties for
highlands of the Rift Valley and on the slopes of Mount resistance to stem rust
Kenya. Annual production stands at 350,000 tonnes Kenyan varieties display seedling resistance to
compared to an annual consumption of 850,000 tonnes races of the stem rust pathogen found in the USA and
making Kenya a net importer of wheat. The total area probably other parts of the world (Table 1). However,
under wheat is 150,000 hectares (ha) with an average when tested with race TTKSK (Ug99) all the Kenyan
yield of 2.5 t/ha. Wheat is grown by both small-scale varieties showed susceptible responses with the
producers (less than 10 ha) and large-scale farmers exception of Bonny which was highly resistant, and
(50 to 10,000 ha). Large scale farmers use high-input Tama and Gem which showed intermediate responses
improved technologies, whereas small scale farmers (Table 1). Given that the races have been evolving over
apply minimal inputs resulting in low returns. There is time, it is clear that most Kenyan varieties were able to
a large production gap (over 70%) between the returns hold resistance against new races until the emergence
for both groups. of Ug99. This can be explained because Ug99 possesses
Wheat production in Kenya is not constraint- a unique combination of virulences for resistance genes
free and wheat is challenged by both biotic and that were commonly used in breeding, especially Sr31
abiotic constraints. The biotic constraints are diseases which is present in some of the more recent varieties
derived from CIMMYT germplasm (Singh et al. 2006).
1
Kenya Agricultural Research Institute- Njoro Plant Breeding Research Center
(KARI-NPBRC), PO Njoro, Kenya; 2CIMMYT, Apdo. Postal 6-641, 06600, Mexico, This degree of stem rust susceptibility to a single race in
DF, Mexico; 3CIMMYT, Nairobi, Kenya; 4USDA-ARS, Cereal Disease Laboratory, St. Kenyan bread wheat has not been observed previously
Paul, MN 55108, USA
E-mail: njaupnn@yahoo.com
R1112* 0.98 1.73 3.85 1.11 1.58 2.52 1.30 2.02 2.94 46.68
R1130* 0.9 2.43 3.51 1.21 1.70 1.11 1.82 1.80 -8.36 30.58
Njoro BW2 1.08 2.13 2.53 0.88 1.13 2.06 2.11 1.82
KS.
1.46 0.49 0.76 .76 0.32 0.65 1.64 0.72
Mwamba
KS. Simba 0.95 2.12 3.03 0.30 0.94 2.44 2.42 1.96
172 Sources of resistance to stem rust race Ug99 and its variants in Canadian wheat germplasm
Table 2 Summary of responses of Canadian wheat cultivars to stem rust race Ug99 and variants at Njoro, Kenya, from
2005 to 2008
Field
Field
reaction % of 2008
reaction
Class & cultivar of most % in 2008 1
Canadian
of most
widely wheat crop
resistant
grown
Reference
CWRS2
CWAD4
174 Sources of resistance to stem rust race Ug99 and its variants in Canadian wheat germplasm
Table 3 Sources of resistance to Ug99 and variants currently used in Canadian wheat breeding programs and the
generation of the experimental lines resulting from hybridization
Sources of
Market class Gene postulation and / or DNA markers Generations
resistance
CW1 Amber Durum Commander Sr9e, Sr13, and Sr14 and other unknown genes F2 to F10
Napoleon Sr9e, Sr13, and Sr14 and other unknown genes F2 to F10
CW Soft White Spring Peace SrCad linked to Bt10, Lr34, Sr2 F1 to F4, DH
CID394092 SrSha7 F1
CID428593 Sr-synth F1
Canada Prairie
AC Cadillac SrCad linked to Bt10, Lr34, Sr2 F1 to F5
Spring_Red
others Sr2, Sr22, Sr24, Sr29, Sr33, Sr35, Sr36, SrTmp F1 to F8, DH
1
CW Canada Western
resistance to stem rust (including Ug99 and its variants) lines with putative resistance based on marker assisted
in Kenya in 2008. The mean stem rust severities for selection will require bioassays under field conditions in
each of the four genotypic classes (above) show that the presence of Ug99 and variants.
the seedling Sr gene on 6DS in combination with Lr34
confers a high level of stem rust resistance, while the Acknowlegements
6DS gene alone only provides only moderate resistance Financial support from producers supporting
in the absence of Lr34 (Fig. 2). All DH lines that expressed the research levy on wheat that is administered by
the high level of resistance found in the Peace parent the Western Grains Foundation, Matching Investment
carried both the 6DS Sr gene and Lr34. Initiative of Agriculture and Agri-Food Canada,
In durum, AM studies identified four regions on Ducks Unlimited, SeCan, and the National Science
chromosomes 1B, 2A, 6A, and 7A (Table 5; Pozniak et al. and Engineering Research Council is gratefully
2008) significant for both field and seedling data and acknowledged. Technical support staff at Cereal
these chromosomes house known Sr genes. Two regions Research Center, Semiarid Prairie Agricultural Research
were identified on chromosome 7A, one distal to the Center, Lethbridge Research Centre, Crop Development
centromere, and a second at gwm276 (Table 5). Sr22 is Center, University of Saskatchewan, R. Wanyera,
linked to gwm276, and that gene is effective against P. Njau, and technical staff at Kenya Agricultural
Ug99. Sr13, derived from cultivated emmer wheat cultivar Research Institute for managing the plots are gratefully
Khapli, resides on chromosome 6A, is effective against acknowledged for their dedication and diligence. Dr. B.
Ug99. Marker gwm617 was significantly associated with Steffenson, University of Minnesota is thanked for the
Ug99 resistance (Table 5) and is likely marking Sr13 as field stem rust ratings in Kenya, 2008.
they are both located on the distal region of 6AL. Sr14,
which is also derived from Khapli and localized distally to Fig. 2 Mean rust severity of four genotypic classes of DH
the centromere on 1BL, provides intermediate resistance lines from the cross of RL6071/Peace. Number of DH lines
to Ug99 (Jin et al. 2007). Marker cfd48 (Table 5) is likely in each group is shown in brackets
detecting variation at Sr14 (Table 5).
Mean stem rust severity in four genotypic
Conclusions
Although all current major cultivars grown in
classes of DH progeny
Canada are vulnerable to African races such as Ug99 and Sr severity by genotype
its variants, CWRS cultivars AC Cadillac and Peace and
Mean stem rust severity
50
deficiencies, can serve as a stopgap in the short term
should Ug99 virulence appear in North America. Genetic 40
analysis is underway to develop DNA markers which are 30
more tightly linked for resistance in hexaploid spring 20
wheat cultivars AC Cadillac and Peace, and to develop
markers for the resistance in durum wheat cultivars 10
Napoleon and Commander. Newly developed markers 0
will be incorporated into the current marker assisted 6D+Lr34 6D Lr34 neither
selection strategies of enhancing the gene frequencies (47) (30) (39) (51)
of Lr34 and Sr2, which appear to confer resistance to Genotype
Ug99 and related races. However, assessment of inbred
Genotype of DH lines
176 Sources of resistance to stem rust race Ug99 and its variants in Canadian wheat germplasm
Table 4 Response of genotypes to Ug99 and variants in rust nurseries near Njoro, Kenya, postulation of genes based on
parentage, molecular markers, and/or phenotype
Ug99 +variants Bt10 Lr34 Sr2 STM559 Other Sr genes
AC Barrie MS no no yes yes
AC Intrepid MS no no yes yes
Alvena S no no no yes
Helios MS no no yes yes
Katepwa MS no no no yes
Manitou MS no no NT yes
Neepawa MS no no no yes
BW553 MR yes no no yes
BW90 MS no yes yes yes
AC Cadillac R yes yes yes yes
Peace R yes yes yes2 yes
BW711 R yes yes No, but PBC3 yes
AC Taber MR yes NT NT yes
AC Foremost I yes no NT yes
AC Crystal MS yes NT NT yes
AC Karma MS yes no NT yes
5700PR MS yes Hetero4 NT yes
1
NT – Not Tested
2
Based on marker X3B028F08
3
PBC pseudo black chaff symptoms expressed under high humidity
4
hetero – heterogeneous
Table 5 Chromosome regions and significance of markers (p-value of F-tests) associated with Ug99 field severity (Sev),
and infection response (IR) and seedling infection type (IT) determined in greenhouse trials. Marker position is based
on the hexaploid wheat consensus map
178 Sources of resistance to stem rust race Ug99 and its variants in Canadian wheat germplasm
28. Principles for rapid variety of global wheat production, is planted with susceptible
varieties. Furthermore, many wheat varieties grown
release, seed multiplication and in major producing and exporting countries, such as
Australia, Canada and USA, are also susceptible to race
distribution in developing countries Ug 99 and derivatives. It is therefore a serious threat to
to counter the threat of wheat rust global wheat production with potentially catastrophic
consequences which could trigger a global food crisis.
The development of resistant varieties, surveillance
Thomas Osborn1, Zewdie Bishaw2
systems and effective plant protection strategies are
important elements of a wheat rust control strategy
Abstract
that need to be urgently implemented. It is critical that
Stem rust race of ‘Ug99’, first detected in Uganda
current, widely grown, stem rust susceptible varieties
(1999), has already reached Kenya (2002), Ethiopia
are replaced with resistant ones to ensure global food
(2003), Yemen (2006) and Iran (2007). Ug99 is a serious
security. Under the Borlaug Global Rust Initiative,
threat to global wheat production with potentially
CIMMYT, ICARDA and a number of Agricultural Research
serious consequences for global food security. The
Institutes (ARIs) and National Agricultural Research
development of resistant varieties, surveillance systems
Systems (NARS) in developed and developing countries
and effective plant protection strategies are important
have tested thousands of accessions in Kenya and
elements to control Ug99. However, to effectively
Ethiopia. A number of elite lines have been identified
counteract the threat of Ug99, a range of resistant
with adequate resistance against Ug99 and up to 15%
varieties and production practices need to quickly get
yield increase. Some of these materials are part of the
into the hands of farmers in areas at risk. In order to do
Elite Bread Wheat Yield Trial (EBWYT) and Stem Rust
this, national contingency planning for seed production
Resistance Screening Nurseries (SRRSN) distributed by
needs to be implemented with a multi-stakeholders and
CIMMYT and/or ICARDA consisting high-yielding wheat
multidisciplinary approach. It should include immediate
lines with adequate levels of resistance to Ug99.
actions such as fast tracking variety evaluation and
Seed is a means for delivering crop-based
release, accelerated seed multiplication, and distribution
innovation to farmers to realize the impacts of
to farmers, in order to replace existing susceptible wheat
investments in agricultural research. Availability of,
varieties in high risk areas. This global threat to food
and access to, wheat seed of resistant varieties is key to
security requires a coordinated rapid response with
counter the threat of stem rust and ensure global food
international, national, local and donor support.
security. The key elements for contingency planning for
the rapid dissemination of wheat rust resistant varieties
Keywords to farmers include:
Variety release, seed multiplication, contingency 1. Rapid variety evaluation and release
planning, wheat, stem rust 2. Streamlining regulatory and phytosanitary protocols
for movement of seed across international
Introduction boundaries
A virulent race of wheat stem rust known as ‘Ug99’ 3. Creating variety awareness and promotions
was first detected in Uganda (1999) and has spread to 4. Rapid seed multiplication
Kenya (2002), Ethiopia (2003), Yemen (2006) and Iran 5. Strengthening infrastructure for seed delivery
(2007). There is a potential threat for the spread of the 6. Human resource development for seed production.
disease to the most important wheat growing areas
of the developing world with serious consequences Status of the national seed sectors
for global food security. From empirical evidence, it is National seed sectors consist of formal (organized)
certain that Ug99 could spread to South Asia and East system and informal (traditional) systems. The formal
Asia. Preliminary screening of widely grown commercial seed system includes seed production and seed supply
wheat varieties from 18 African and Asian countries mechanisms operated by the public and private sectors
against ‘Ug99’ revealed that about 85% of them are under some measure of supervision and regulation
susceptible to Ug99. Accordingly, an estimated 52% in a commercial or quasi-commercial mode within
of the total wheat area of 74.6 million ha planted with the framework of national seed policy and legislation.
wheat in these countries, collectively representing 40% Improved varieties are the results of formal plant
breeding and variety development that are tested and
1
FAO, Rome, Italy; 2ICARDA, PO Box 5466, Aleppo, Syria
E-Mail: thomas.osborn@fao.org; z.bishaw@cgiar.org released in the country if they are proved superior to the
180 Principles for rapid variety release, seed multiplication and distribution in developing countries to counter the threat of wheat rust
eastern states are planted with three ‘obsolete’ varieties coordinated international nurseries network distributing
released between 1971 and 1986, and one variety EBWYT and SRRSN. It is anticipated that countries will
(released in 1996) alone covers about 7 million ha in be able to share promising Ug99 resistant lines in order
the north western states of the country (Ferrara et al. to accelerate the process of testing and releasing a wide
2007). Lack of wheat varietal diversity and dominance range of resistant wheat varieties. It will be important
by a few varieties over large areas is also a widespread for national varietal testing systems to be strongly linked
phenomenon in other countries and regions (e.g. with international information sources and sharing
Ethiopia, Pakistan) predisposing farmers to vulnerability varietal data and performance under a wide range of
to diseases and risks of food insecurity. There is an agro-ecologies.
urgent need to speed up varietal replacement in order Given the global scope of the threat and the need
to counteract the threats of wheat rust epidemics and for a global response, efforts should be made to consider
ensure the food security. a policy for joint release within regions. Since many NARS
For promising new wheat varieties, there are are evaluating similar breeding materials across regions,
standard procedures in most countries for variety with potential for both wide and specific agro-ecological
testing and release before they can be multiplied and zones of adaptation, opportunities must be explored
used by farmers. Variety release can require simultaneous for joint or regional release of varieties. Harmonized
testing of promising new lines for registration concerning regional variety release schemes should be considered.
Distinctness, Uniformity and Stability (DUS), and Streamlining regulatory and phytosanitary protocols for
performance testing of Value for Cultivation and Use movement of varieties and seeds across international
(VCU). Some countries have well defined compulsory boundaries are also needed as part of regional
variety testing and release procedures (both DUS harmonization of seed rules and regulations. Attention
and VCU) in place (e.g. Egypt, Pakistan, India, Turkey), should be given in diversifying the portfolio of varieties
whereas in others, the release system is purely released across the countries. In the absence of a regional
dependent on performance testing conducted by the release system, national authorities should consider a
agricultural research authorities and approved by ad clause for exemption from compulsory registration for
hoc release committees (e.g. Ethiopia, Kyrgyzstan). wheat rust resistant varieties coming from similar agro-
The established procedures for national variety trials ecological conditions outside of the country.
sometimes require many years. This means that even 2.2. Varietal choice It is anticipated there will be
with countries that share similar agro ecologies, most many wheat rust resistant lines adapted to a wide range
countries require compulsory registration in a national of agro-ecological zones and with farmer preferred traits
varietal catalog before a variety released in another available in the near future from international and/or
country could be authorized for cultivation. national breeding programs. National programs with
Lengthy and slow variety testing and release responsibility to screen wheat rust resistant varieties will
remains a critical bottleneck to speedy release of new be able to source the most promising lines for evaluation
varieties. As part of contingency planning, national on release and to use them in national breeding
authorities should allow NARS to adopt a fast-track programs. New wheat varieties must combine not only
release for wheat rust resistant varieties with superior wheat rust resistance, but also resistance to other major
field performance and acceptable organoleptic diseases and yield superiority compared to existing
characteristics. This fast-track approach could introduce varieties as well as adaptation to the use of wheat by
varieties with known resistance and test them for two farmers, consumers and industries. National breeding
growing seasons in multi-location adaptation trials and programs should aim at developing and releasing a wide
release them promptly for large-scale use. The serious range of varieties with diverse genetic backgrounds
threat of wheat rust means that variety evaluation and for wheat rust resistance to reduce vulnerability and
release may need to be streamlined so that it is efficient risk of disease epidemics. It is important to maintain
and effective, but at the same time carried out in as varietal diversity and overcome varietal dominance
short a timeframe as possible. Variety evaluation and by identifying and releasing those with comparable
release, or even compulsory registration, in the national agronomic performance and preferred traits at national
varietal catalog should not delay the process of getting and/or regional levels.
resistant varieties to farmers. 2.3. Access to public varieties Wheat breeding
2.1. Regional variety release International in many developing countries is dominated by IARC
Agricultural Research Centers (IARCs) in partnership breeding materials distributed to the NARS. In many
with NARS are at the forefront of a breeding program countries public seed companies have sole access to
to develop stem rust resistant varieties through a new varieties. Access to new publicly bred varieties
182 Principles for rapid variety release, seed multiplication and distribution in developing countries to counter the threat of wheat rust
Fig. 1 Accelerated seed multiplication scheme for stem rust resistant wheat varieties
collaboration with NARS, or as joint activities with the irrigation, storage, cold storage) for main or off-season
public/private seed producers and suppliers. seed production. Creating and strengthening such
Despite huge investments in variety development, units would institutionalize early generation seed
most NARS pay limited attention to early generation production on a sustainable basis and enable countries
seed production due to lack of funding and incentives to adequately respond to any future emergency
coupled with absence of functioning seed units due to situations. In many countries, the benefits generated by
insufficient land and facilities (e.g. machinery, irrigation). selling early generation seed go to government treasury
These tasks require specialized field equipment (e.g. instead of being directly used by the institutions that
plot planters, plot harvesters, small cleaners/treaters) produce the seed. This financial arrangement is a
for timely operations and appropriate facilities (e.g. disincentive and could be partly responsible for lack
184 Principles for rapid variety release, seed multiplication and distribution in developing countries to counter the threat of wheat rust
Table 1 Estimated seed multiplication with varying yield levels (3, 4 and 6 tonnes ha-1)
Generation Quantity of seed produced in tonnes with different multiplication factors
1:30 1:40 1:60
Initial seed quantity (t) 0.05 0.05 0.05
1.5 2.0 3.0
First
(0.5 ha) (0.5 ha) (0.5 ha)
45 80 180
Second
(15 ha) (20 ha) (30 ha)
1,350 3,200 10,800
Third
(450 ha) (800 ha) (1800 ha)
40,500 128,000 648,000
Fourth
(13,500 ha) (32,000 ha) (108,000 ha)
1,215,000 5,120,000 38,880,000
Fifth
(405,000 ha) (1,280,000 ha) (6,480,000 ha)
Figures in parentheses indicate the areas required to produce the seed of the concerned generation
wheat rust already have systems for seed multiplication, undertake intensive wheat seed production in order to
modifications may be needed to cope with the urgency reduce the time needed to produce the target quantities
of rapid large-scale multiplication and distribution of of wheat seed.
resistant varieties, especially to the most vulnerable National seed programs in risk-prone areas will
small farmers. need to have contingency plans for maintaining and
Partnerships with private sector seed companies managing carry-over certified seed stocks to overcome
may be the quickest and most cost-effective strategy shortfalls in seed supply due to emergencies or crop
for seed multiplication in some countries. Contingency failures. The main purpose is to ensure a reliable seed
planning will include a well coordinated system for rapid supply to the farming community through the activities
seed multiplication with an effective partnership and of the formal sector. The excellent storability of wheat
high level of coordination in order to be successful. seed provides the option of rapid seed multiplication
Table 1 presents the theoretical basis of seed of rust resistant varieties and establishing a strategic
multiplication assuming an initial 50 kg of nucleus reserve of early generation seed even before rust has
seed of a new variety at a planting rate of 100 kg ha-1 threatened production. With this strategy resistant
and anticipated yields of 3, 4 and 6 tonnes ha-1, i.e. wheat varieties can be quickly released when needed.
multiplication factors of 30, 40 and 60, respectively. The
table provides an idea of the numbers of generations, 3.5. Seed import and distribution Another
areas needed for seed multiplication, and areas the seed potential strategy for the urgent provision of seed of
can cover at various multiplication factors. rust resistant varieties may be the direct import and
The importance of intensive production of seed distribution of seed from neighboring countries in
through optimum crop management practices (e.g. areas where agro-climatic conditions are similar and the
weed control, water) to achieve higher multiplication variety is adapted, tested and released in the importing
factors is highlighted in the differences in the MF and country. There are clear practical examples of importing
the resulting total production after five generations. In seed of improved varieties from neighboring countries
addition, the large areas required for seed multiplication for distribution through public/private seed sector and
provide an idea of the scope of the seed multiplication NGOs both under normal and emergency situations.
needed in order to address the wheat rust threat. In order to anticipate such a situation, an
Table 2 provides an idea of the quantities of effort to undertake regional harmonization of seed
seed that may be needed in each country. A tentative rules and regulations, especially in the area of seed
target of seed to cover 10% of the total area in wheat certification and plant quarantine would facilitate
is included. The figures demonstrate the need to seed trade between countries of the same region.
186 Principles for rapid variety release, seed multiplication and distribution in developing countries to counter the threat of wheat rust
Regional harmonization of seed rules and regulations as tractors, implements, irrigation equipment, and
is underway in several regional areas of Africa. In combines, and seed processing and storage facilities
addition, a seed association has been established in the may be needed for large-scale seed multiplication. A
central Asia and west Asia region to create a forum for review of equipment and facilities for seed production,
dialogs amongst stakeholders concerned with the seed seed conditioning (processing) and seed storage
trade/industry development. This association will also should be undertaken to ensure that this element is
play a crucial role in the establishment of procedures not a constraint to the rapid seed multiplication and
and processes for the harmonization of seed rules delivery of quality seed. Provisions of farm machinery
and regulations aimed at facilitating cross boundary and equipment should be made on a case by case basis
movement of seed. Given the importance of wheat specific to the needs of the individual countries.
in this region this initiative will be a very important 4.1. Seed storage facilities Wheat seed can
strategy to counter the threat of wheat rust. be effectively stored without losing its vigor and
3.6. Quality assurance The production of quality germination if well known precautions are taken.
certified seeds will require critical roles for the national Keeping the seed as dry and cool as possible in clean
seed services and related seed certification agencies to stores is the best management practice because
ensure the inspection, testing and certification of the physiological processes and fungal and insect activities
seed. Some countries may need to strengthen their seed are low. Safe storage of wheat seed is possible as long
certification services to respond to this demand through as the moisture level is below 13%, the humidity is low,
training of additional technical officers to handle ambient storage temperature is not excessive and the
the increased workload and transportation support. infestation with storage insects is minimal. In practice,
In addition, related facilities such as seed testing stored wheat seed should be kept at moisture levels
laboratories may need to be established or upgraded to below 12% and relative humidity below 50-60%. Van
cope with the higher demand for services and possibly Gastel et al. (2003) cited practical guideline in choosing
to implement regionally harmonized seed rules and alternative sites for short and medium term seed
regulations for variety release, phytosanitary standards storage. It is advisable to select a seed storage site,
and the seed trade. which is cool and dry (low relative humidity).
The databases of the National Seed Certification Adequate seed storage facilities should be
Agencies and national variety registries should also be made available particularly for the maintenance and
strengthened to include information on attributes such management of carry-over stock of early generation
as responses to rust races to which wheat varieties are seed and certified seed. Small quantities of seed can be
resistant, and lists of plant quarantine pests to facilitate stored for long periods in cold rooms; whereas properly
exchange of tolerant varieties among countries. designed seed storage facilities might be required for
larger quantities.
4. Strengthening seed multiplication capacity 4.2. Human resource development for seed
Rapid seed multiplication of large quantities of production Specific knowledge and practical
seed will require capacity building beyond the ongoing experience is needed to produce high quality seed
normal seed activities at country or regional levels. This with high standards of varietal purity, and physical
emergency situation brings more work to already limited and physiological health. From the outset, assistance
infrastructures to undertake a huge task of both pre- in capacity development of NARS and national seed
release and large-scale seed multiplication where time is programs is a key to enhance their technical and
an essence of all operations. Apart from efficient use of managerial capacities as well as the implementation
existing resources and facilities additional investments of regionally harmonized seed rules and regulation.
may be necessary for robust response to the crisis. Three levels of training could be envisaged from policy
Seed production is a specialized task both in field makers to farmers ranging from policy issues to practical
operations and post-harvest handling of seeds. Speed experience: (i) workshops to create awareness and
and time are the essence in an emergency situation. inform senior staff and policy makers; (ii) training of
There are urgent needs for special equipment (single trainers’ courses for technical managers and technicians;
ear/bundle threshers, plot planters, plot harvesters, and (iii) practical training for farmer seed producers and
and seed cleaners/treaters) for timely operations growers. Private seed companies will be key partners
and appropriate facilities (e.g. irrigation, storage, etc) in certified seed production and they will benefit
for main or off-season seed multiplication of early from training programs. Training will be required in
generation seed. Additional field machinery such principles and techniques of variety maintenance, seed
188 Principles for rapid variety release, seed multiplication and distribution in developing countries to counter the threat of wheat rust
29. Ethiopia’s experience with rapid For the informal seed system, research stations serve
as the initial source of seed, and farmers, farmers’
seed multiplication and cultivar co-operatives, Ministry of Agriculture and Rural
Development, and non-government organizations
replacement are the main participants in seed multiplication and
dissemination. Countrywide attempts at informal seed
Bedada Girma1, Balcha Yai1, Sintayehu Debebe1, Tezera multiplication and scaling-up of crop technologies
Walabu2, Lijalem Korbu3, Sherif Aliye3
have resulted in small success stories in the late 1990s
and early 2000s. Cereals, pulses, oilseeds and tuber
Availability and access to improved seed are the
crop varieties have gone through participatory seed
constraints for improving productivity and production
multiplication and scaling-up with encouraging success.
of crops in Ethiopia. In the case of wheat, even when
Recent experiences show that rendering research
available, varieties often lose their disease resistance
center-based support to both the informal and formal
before or soon after their adoption by farmers due to
seed systems can improve variety adoption and the
inefficient seed multiplication and delivery systems.
access of farmers to high quality seed. Participatory
The current production of improved seed accounts
seed multiplication at the village level, supported by
for less than 3% of all seeds needed for annual crop
some training and minimum guidance, is regarded as a
production. Over 97% of the seeds come from farmer
useful approach for rapid technology dissemination and
sources through traditional seed exchange or seed
cultivar replacement under Ethiopian conditions. This
marketing. There are two seed supply and delivery
paper describes Ethiopia’s experience in informal rapid
systems in Ethiopia. These include the formal and the
seed multiplication of wheat, malting barley, faba bean,
informal sectors. Research, as a source of breeder and
lentils, and haricot beans.
pre-basic seed, the public seed enterprises, and some
private businesses represent the formal seed sector.
192 ExperiencewithrapidseedmultiplicationandcultivarreplacementtargetingraceUg99resistantwheatvarietiesintheEasternIndo-GangeticPlains
listed in Table 3. In the first year (2005-06), the lines Fig. 2 Decline in percent coverage of wheat cultivar HUW
used were the best short duration lines of the CIMMYT 234 and increase in zero till area for wheat in district
breeding program targeting irrigated environments, but Chandouli, Uttar Pradesh, India (wheat area = 100,000 ha)
carried only mild tolerance to Ug99 as it coincided with
the beginning of the Ug99 resistance breeding program.
Each year PVS followed the “mother baby” approach
(Witcombe et al. 2001) with two standard check
varieties, HUW 234 and HUW 468. In the current 2008-09
season, a set of ten varieties carrying resistance to Ug99
are under evaluation at ten locations.
Results for six lines that yielded 10% or above
HUW234 in the year 2005-06 are given in Table 4. Three
lines with 16.6 to 21.1% higher yields than HUW234
were the earliest maturing among the group with the
same days to heading as new cultivar ‘HUW468’. In the
next two years several new lines proved superior to local
checks in yield performance by over 10% (Table 5). As
shown in Table 3, all these superior varieties carry higher 2008-09, and one line was promoted to the National
levels of resistance to Ug99. Initial Varietal Trial (NIVT) by Banaras Hindu University.
Some of these Ug99-resistant varieties have also been
Inclusion of Ug99 resistant lines in the advanced to national coordinated trials by different
national coordinated trials research centers, i.e. Punjab Agricultural University and
In addition to PVS trials, the superiority of many Directorate of Wheat Research.
of the Ug99 resistant lines, e.g. Munal#1 (Waxwing*2/
Kiritati) and Quaiu#2 (BABAX/LR42//BABAX*2/3/ Seed dissemination in South Asia
VIVITSI), has already been established through the Elite Although a substantial network of organized (both
Bread Wheat Yield Trials (EBWYTs) tested across many public and private sectors) seed production does exist
locations of South Asia in the past three years (DWR for germplasm dissemination and adoption in India, the
Report, 2007; 2008). These lines were included in the seed replacement rate is still less than 20% (Joshi et al.
All India Wheat Coordinated Trials organized by ICAR. 2007). The actual figure is believed to be around 10%
Likewise, based on the results of PVS trials in farmers’ in the eastern part of Indo-Gangetic plains (Joshi et al.
fields, five of the Ug99 resistant varieties were included 2007). It is accepted that for proper dissemination and
in the Indian plant protection screening nursery (IPPSN), adoption of germplasm, both the public and private
Table 1 Stem rust responses of wheat cultivars and advanced breeding lines from South Asia at Njoro, Kenya in 2006
and 2007
No of lines
Country of origin Moderately Moderately sus. & Total
Resistant1
resistant2 susceptible3
Bangladesh 4 8 112 124
India 16 7 79 102
Nepal 1 6 153 160
Pakistan 3 24 184 211
Total 24 45 528 597
1
Disease severities up to 20% based on modified Cobb scale; small to intermediate sized uredinia with necrosis or
chlorosis
2
Disease severities between 15 and 30%; medium to large uredinia with or without chlorosis and necrosis
3
Disease severities >40%, medium to large uredinia without chlorosis and necrosis
sectors need to be strengthened in all developing et al. 2000). Therefore, by the time a variety reaches
countries of South Asia. Considerable effort is already the majority of farmers, it has already lost much of its
underway. However, it is also believed that in view of the potential impact due to reduced, or loss of resistance.
huge wheat area in the eastern Gangetic plains covering Because the Gangetic plains cover a vast area with a
a wide range of socio-economic and environmental complex of socio-economic issues, improvements in
diversity, greater scientist-farmer interaction following the availability of quality seed need to be achieved
a participatory mode could play a crucial supportive through a combination of formal and informal activities.
role to meet this objective (Ortiz-Ferrara et al. 2007; Therefore, strengthening the capacity of farmers to
Joshi et al. 2007). The role of participatory research in undertake quality seed production following the
varietal selection (Ferrara et al. 2002; Witcombe et al. participatory approach assumes high priority (Joshi et
2001, 2003) of different crops is well documented. This al. 2007). Indian research centers already work on this
assumes further importance due to the fact that in model and so far the participatory mode has proven
many locations farmers’ access to new varieties is highly quite successful (Ortiz-Ferrara 2001; Joshi et al. 2007). At
restricted, and therefore good technology takes a very many locations in the eastern Gangetic plains, farmers
long time to disseminate. For example, it is believed have started their own seed businesses. To promote
that a good agricultural technology takes around 10 further seed multiplication and dissemination, ICAR
years to spread in the eastern Indo-Gangetic plains, and in 2003 made it mandatory for all research centers
the average life of a resistant variety to rust pathogens receiving support under the National Seeds Project
(which are still the dominant pathogens in most parts of to actively engage in participatory seed production.
the world, and especially, India) is believed to be around The recent success of the new ICAR seed project “Seed
5-6 years (Roelfs et al. 1992; Rajaram et al. 1998; Singh Production in Agricultural Crops and Fisheries”, launched
194 ExperiencewithrapidseedmultiplicationandcultivarreplacementtargetingraceUg99resistantwheatvarietiesintheEasternIndo-GangeticPlains
Table 3 New CIMMYT lines under PVS in eastern Gangetic plains of India and their reactions to Ug99
Year/
Name Pedigree Main season Off season Main season Reponse
No.
SR Kenya SR Kenya SR-Kenya SR-Kenya SR-Kenya category
2005-06 2006* 2007 2007 2007 2007
1 Baaz ATTILA*2/STAR/4/SNI/TRAP#1/3/KAUZ*2/TRAP//KAUZ 60S - 15MSS 40MSS 60MSS MS
2 Labh UP2338*2/4/SNI/TRAP#1/3/KAUZ*2/TRAP//KAUZ 60S - - - - MS
3 Layak INQALAB 91*2/KUKUNA 80S - - - - S
4 Sundar WBLL4/KUKUNA//WBLL1 30MSS - - - - MS
5 Tej WBLL1*2/KUKUNA 30M 30S 30MSS 50MSS 60MSS MS
6 Takat WBLL1/4/HD2281/TRAP#1/3/KAUZ*2/TRAP//KAUZ/5/KAMB1 40S - - - - S
7 Lahar SERI.1B//KAUZ/HEVO/3/AMAD - - - - -
8 Vishal ATTILA*2/PBW65 70MSS - 5MR 10MSS 30MSS MR
2006-07
1 Jhoola KIRITATI/4/SERI.1B*2/3/KAUZ*2/BOW//KAUZ - 80S 30S 50S 50S MS-S
2 Mahak KIRITATI//PRL/2*PASTOR - 70S 40S 60S 60S MS
3 Swasth KIRITATI//ATTILA*2/PASTOR - 80S 20S 40S 60S MS-S
4 Agrim KIRITATI//HUW234+LR34/PRINIA - 10MR 10RMR 20M 30M MR
5 Sona KIRITATI/WBLL1 - 40MSS 30MSS 70S 70S MS
6 Chandi WEAVER/TSC//WEAVER/3/WEAVER/4/PRL/2*PASTOR - 70MSS 70S 100S 100S S
7 Panchi PFAU/SERI.1B//AMAD/3/WAXWING - 60MSS 40MSS 50MSS 70MSS MS
8 Ufan WAXWING*2/VIVITSI - 30M 5M 20M 20M MR
9 Uthan WAXWING*2/TUKURU - 60MSS 40MSS 50MSS 60MSS MS
10 More WBLL1*2/KIRITATI - 30MSS 15MSS 40MSS 60MSS MS
11 Hans KAMB1*2/BRAMBLING - - -
12 Abhinav KAMB1*2/KIRITATI - - -
2007-08
1 Sarpat KIRITATI/2*WBLL1 - 20MSS 30MS 50MSS 50MSS MR-MS
2 Hans HUW234+LR34/PRINIA//PFAU/WEAVER - 5M 5M 10M 20M R-MR
ELVIRA/5/CNDO/R143//ENTE/MEXI75/3/AE.SQ/4/2*OCI/6/
3 Koyal - 40M 30MSS 40MSS 50MSS MR-MS
VEE/PJN//KAUZ/3/PASTOR
4 Ravi PFAU/WEAVER*2//KIRITATI - 30M - - - MR
5 Guru KIRITATI//SERI/RAYON - 20M 5M 10M 20M R-MR
6 Century WAXWING*2/KIRITATI - 20M 5MR 15M 15M R-MR
7 Umang WAXWING*2/4/SNI/TRAP#1/3/KAUZ*2/TRAP//KAUZ - 20M 5MR 20MSS 20MSS R-MR
Checks
1 HUW 234** HUW 12*/CPAN 1966 (Sparrow)
2 HUW 468** CPAN 1962/TONI//LIRA/PARULA
195
Table 4 Mean grain yields, days to heading, heights and 1000 kernel weights for six CIMMYT derived advanced lines
and two cultivars tested at seven sites1 in eastern Gangetic plains of India during crop season 2005-2006
during the 10th Five Year Plan for 2005-06 and 2006- estimated 25,000 tonnes of seed (including all classes,
07 (http://www.teatronaturale.com/article/12.html) viz. breeder, foundation, certified and truthful seed). This
also includes participatory seed production, further cultivar needs to be reselected for resistance for seed
suggesting that new varieties can be disseminated in a multiplication till other new resistant lines get multiplied
much faster way in South Asia. in sufficient quantity. Another new resistant variety, BL
Newly developed Ug99-resistant lines are now 3063 (FRTL/Chirya 7), developed in Nepal is also being
under seed multiplication in the eastern Gangetic multiplied in farmers fields.
plains of South Asia; these include Picaflor#1 (Kiritati//
Seri/Rayon), Pauraque#1 (Waxwing*2/4/SNI/Trap#1/3/ Conclusion
Kauz*2/ Trap//Kauz), Becard#1 (WBLL1*2/Kiritati), The new stem rust race Ug99 is a serious threat
Munal#1 (Waxwing*2/Kiritati), Quaiu#2 (Babax/LR42// to South Asia and to global wheat production. If
Babax*2/3/Vivitsi), Francolin#1 (Waxwing*2/Vivitsi) and not checked through effective research, Ug99 may
Damphe#1 (Kiritati//2*PBW65/2*Seri.1B). These lines become another cause of food shortage for many
currently occupy around 15 ha under participatory countries, including those in South Asia. Replacement
seed production in the present crop season (2008-09) of currently popular susceptible cultivars in these
with a targeted production of at least 30 tonnes. The areas with high yielding resistant lines is the best
predicted production of these varieties in the next crop strategy to protect wheat from the menace of Ug99.
season (2009-10) is around 500 tonnes. This is based This will require a concerted effort involving scientists,
on the results of the seed production of some of the planners, progressive farmers and extension agencies
lines, viz. Baz, Labh and Lahar (Table 3), introduced from associated with governmental and non-governmental
CIMMYT in 2005-06. These three lines with moderate organizations. CIMMYT, in collaboration with national
susceptibility to race Ug99 each occupy about 100 ha in research centers of South Asia, has already developed
2008-09, with an anticipated seed production of more several high yielding Ug99 resistant varieties. These
than 250 tonnes. With likely official release of these lines varieties are under seed production mainly through
in the near future, it is predicted that seed production in participatory seed production in the eastern Gangetic
the next three years will generate seed to saturate the plains of India. In Pakistan, Bangladesh and Nepal the
eastern Gangetic plains and reduce the threat of Ug99. lines are being multiplied by national research centers.
In addition, the heterogeneous Ug99 resistant variety A more concerted seed production and dissemination
HUW 234 (not reselected for resistance) is also under system is required to safeguard South Asia from the
increase in the eastern Gangetic plains to produce an threat of Ug99.
196 ExperiencewithrapidseedmultiplicationandcultivarreplacementtargetingraceUg99resistantwheatvarietiesintheEasternIndo-GangeticPlains
Table 5 Mean grain yield of CIMMYT derived advanced lines that displayed 10% or higher superiority over one of two
popular cultivars tested at multiplication sites1 in eastern Gangetic Plains of India during crop season 2006-2007 and
2007-08
% over % over
No. Pedigree Local name2 Mean
HUW 234 HUW 468
2006-07 (9 locations)
1 Kiritati/4/Seri.1b*2/3/Kauz*2/Bow//Kauz Jhula 3.91 11.41 12.06
2 Kiritati//Attila*2/Pastor Swasth 3.94 12.22 12.88
3 Kiritati//Huw234+Lr34/Prinia Agrim 3.89 10.91 11.56
Ufan
4 Waxwing*2/Vivitsi 3.98 13.31 13.97
(Francolin#1)3
5 Waxwing*2/Tukuru Uthan 4.08 16.09 16.77
Attila*2/Star/4/Sni/Trap#1/3/Kauz*2/Trap//
6 BAJ 3.84 9.49 10.12
Kauz
Check CPAN 1962/Toni//Lira/Parula Malviya 468 3.49
Check HUW 12*/CPAN 1966 (Sparrow) Malviya 234 3.51
LSD (P = 0.05) 0.29
2007-08 (8 locations)
1 HUW234+LR34/PRINIA//PFAU/WEAVER Hans 3.28 6.92 10.37
Guru
2 KIRITATI//SERI/RAYON 3.46 12.58 16.21
(Picaflor#1)3
3 KIRITATI//HUW234+LR34/PRINIA Ufan 3.39 10.46 14.02
Century
4 WAXWING*2/KIRITATI 3.14 6.71 10.15
(Munal#1)3
Check CPAN 1962/Toni//Lira/Parula Malviya 468 3.07
Check HUW 12*/CPAN 1966 (Sparrow) Malviya 234 2.97
LSD (P = 0.05) 0.18
1
Nine farmers’ fields located in districts of Varanasi, Mirzapur, Azamgarh and Chandouli in the eastern Gangetic plains
and research station of Banaras Hindu University
2
Local names of varieties given by farmers in the region
3
Name given by CIMMYT, Mexico
198 ExperiencewithrapidseedmultiplicationandcultivarreplacementtargetingraceUg99resistantwheatvarietiesintheEasternIndo-GangeticPlains
31. Field efficacy of fungicides used Both small- and large-scale farmers have been affected.
Yield losses of up to 80% were reported (Expert Panel
against stem rust in Kenya 2005; Wanyera et al. 2006). The current commercial
wheat cultivars are highly susceptible to the new race
R. Wanyera1, J.K. Macharia2, S.M. Kilonzo1 and it is not possible to grow a profitable wheat crop
without the application of a fungicide. Over the past two
Abstract decades, varietal resistance to stem rust has generally
Field experiments were conducted at two locations provided adequate protection without the need for
during the 2006 and 2007 growing seasons to assess fungicides (Expert Panel 2005; Loughman et al. 2005).
the effectiveness of two new foliar fungicides; viz. Therefore, fungicide control regimes may play a role
Nativo 300 SC (trifloxystrobin100g/L + tebuconazole in integrated management of the disease until new
200g/L) and Prosaro 250 EC (prothioconazole 125g/L varieties become available. Limited studies have been
+ tebuconazole 125g/L), in controlling stem rust on conducted to determine the effects of foliar fungicides
wheat cultivar ‘Duma’. AmistarXtra 280 SC (azoxystrobin on stem rust severities and yields elsewhere (Dill-Macky
200g/L + cyproconazole 80g/L) and Folicur 250 EC et al. 2000), but not in Kenya. This paper reports field
(tebuconazole) were used as checks. The treatments experiments conducted under natural infection to
at each site and year included an untreated control determine the effect of two new foliar fungicides, viz.
and two spray applications of the fungicides at growth Nativo 300 SC (trifloxystrobin100g/L + tebuconazole
stages (GS) 55 and 65. Stem rust severities were assessed 200g/L) and Prosaro 250 EC (prothioconazole 125g/L +
using the modified Cobb scale at 14-day intervals after tebuconazole 125g/L) on wheat stem rust, grain yield
application. The data were used to calculate mean and 1000-kernel weight.
rust severity (MRS). Stem rust epidemics were severe
at KARI-Njoro in 2006 and the treatment effects on Materials and methods
stem rust severities, grain yield and 1000-kernel weight Field trials were conducted in 2006 and 2007 at
were significant at both the KARI-Njoro and Mau-Narok Kenya Agricultural Research Institute (KARI)-Njoro and
sites. The fungicide treatments, significantly (P ≤ 0.05) Mau-Narok (Purko Ranch), Kenya. The test cultivar was
reduced stem rust severity, increased grain yield and ‘Duma’, which is popular and recommended for low and
1000-kernel weight of the susceptible wheat cultivar medium elevation growing-areas. The cultivar is highly
‘Duma’ compared to the untreated control. susceptible to stem rust, but is fairly resistant to stripe rust
(caused by P. striiformis Westend. f. sp. tritici). Stem rust
Keywords epidemics occurred naturally at both sites. A randomized
Triticum aestivum, resistance, cultivar, epidemics, complete block design with four replications was used.
crop loss The cultivar ‘Duma’ was planted in 9 m2 plots. Planting was
on May 30 and 16 and September 19 and 29 at KARI-Njoro
Introduction and Mau-Narok in 2006 and 2007, respectively. The plots
Stem rust (Puccinia graminis Pers. f. sp. tritici Eriks. were sown using an experimental seed-drill at a seeding
& Henn.) is one of the wheat rusts that cause severe rate of 100g/plot. A uniform application of Di-ammonium
losses throughout the world. Losses of 50-70-% have phosphate fertilizer (18% N: 46% P: 0% K) was applied
often been reported under field conditions. The actual at planting at the recommended rate of 150Kg/ha. The
amount of loss caused by rust can range from slight to plots were sprayed with Stomp 500E (pendimethalin), a
complete destruction of the crop. Grain from infected pre-emergent herbicide, at the rate of 3L/ha, to control
crops is shriveled and light in weight, and therefore has grass weeds, and Buctril MC (bromoxynil + MCPA) at the
reduced quality (Agrios 1988; Stubbs et al. 1986; Zadoks rate of 1.25L/ha at growth stage GS 24 (Zadoks et al. 1974)
et al. 1974). to control broad leaf weeds. Metasystox 250 EC (oxy-
Currently, all three rusts threaten wheat (Triticum demeton-s-methyl) insecticide was applied at the rate of
aestivum L) production in Kenya. Epidemics occur when 0.5L/ha to control cereal aphids. The fungicide treatments
environmental conditions during the growing season included; two new products, Nativo 300 SC (trifloxystrobin
are favorable. The new virulent strain TTKS (Ug99) in the 100g/L + tebuconazole 200g/L) and Prosaro 250 EC
eastern African region has caused repeated stem rust (prothioconazole 125g/L + tebuconazole 125g/L) each
epidemics since 2002, threatening wheat production. applied at three rates; 0.6, 0.75 and 1.0L/ha. Two standard
fungicides; AmistarXtra 280 SC (azoxystrobin 200g/L +
1
Kenya Agricultural Research Institute (KARI)-Njoro, PO. Private Bag, Njoro cyproconazole 80g/L), and Folicur 250 EC (tebuconazole),
20107, Kenya. 2 Egerton University, P.O Box 536, Egerton, Kenya
E-mail: wanyera@plantptotection.co.ke; wanyera@karinjoro.org each applied at the rate of 1.0L/ha, and an untreated
1000-kernel
Stem rust severity Grain yield 1000-kernel weight Stem rust severity Grain yield
weight
Treatmente
Rate % % % % %
MRSa t/ha g % increase c MRSa t/ha g
L/ha reductionb increasec reductionb increasec increase c
27.9 29.4
Untreated - 52.5 - 0.5 - - 40.8 a - 1.4 c -
c c
-
35.9 41.1
Nativo 300 SC 0.75 24.6 53.1 1.2 58.3 28.7 11.3 bc 72.3 2.6 b 46.2 39.8
a a
33.8 38.2
Nativo 300 SC 1.0 23.3 55.6 1.3 61.5 21.1 10.8 bc 73.5 2.6 b 46.2 29.9
b ab
34.9 41.0
Prosaro 250 EC 0.6 25.4 51.6 1.2 58.3 25.1 9.9 c 75.7 3.3 b 75.6 39.5
ab a
34.7 37.4
Prosaro 250 EC 0.75 24.2 53.9 1.1 54.5 24.4 14.2 bc 65.2 2.5 b 44.0 27.2
ab b
35.6 38.4
Prosaro 250 EC 1.0 21.3 59.4 1.2 58.3 27.6 10.0 bc 75.5 2.4 b 41.2 30.6
a ab
38.4
Folicur 250 EC 1.0 25.4 51.6 1.2 58.3 35.9 28.7 16.7 b 59.1 2.7 b 48.1 30.6
ab
Meand - 24.9 52.7 1.2 57.3 28.5 24.5 12.9 68.4 2.7 49.7 38.9 25.13
Abstract Introduction
This paper is aimed at introducing the work of the
On 1 October 1964, the Food and Agriculture
Joint FAO/IAEA Division of Nuclear Techniques in Food
Organization of the United Nations (FAO) and the
and Agriculture, focusing on mutation induction assisted
International Atomic Energy Agency (IAEA) created the
breeding and providing a brief overview of the roles
Joint FAO/IAEA Division of Nuclear Techniques in Food
of molecular genetics and cellular biology as efficiency
and Agriculture [1] through the first arrangements
enhancing bio- and molecular technologies to broaden
concluded by Directors General of both Organizations.
the genetic base of germplasm available to breeders.
The goal was to combine the talents and resources of
An Interregional Project is proposed to complement
both organizations into assisting their Member States
ongoing international activities on wheat stem (black)
in applying nuclear techniques for providing people
rust (caused by the fungal pathogen Puccinia graminis
with more, better and safer food and other agricultural
f. sp. tritici race Ug99), providing a platform for the
products, while sustaining the natural resources base.
coordination of a network of laboratories (based on
Over four decades, Joint Division activities have
previously established laboratory infrastructures
evolved to respond to the ever-changing landscape of
through the International Atomic Energy Agency’s
agriculture and nuclear technology and the expectations
Technical Cooperation Projects) as a defense line against
of national and international organizations for
Ug99. Implementation is projected in three overlapping
cooperation in nuclear research and technology transfer.
phases: (i) Normalization: adoption and training in the
Throughout this process, the Division has successfully
use of uniform detection and pathotyping protocols,
remained at the forefront of assisting countries in
in order to assure homogeneity of handling (capacity
fostering the uses of nuclear science and technology
building); (ii) Quality control: double blind tests for
where these really add value. Today, the Joint Division
identification and characterization of false positives
strives to mobilize commitment and action to meeting
vs. false negatives (periodic network performance
the World Food Summit and Millennium Development
meetings, quality management and steering); (iii)
Goals of reducing hunger, poverty and environmental
Multilocation trials of mutant germplasm in endemic
degradation through sustainable agriculture and rural
hotspots/screen houses. The minimal network of Ug99
development.
surveillance laboratories shall be comprised of: (i) Front
The International Atomic Energy Agency (IAEA)
line: Kenya, Yemen, Sudan, Egypt, Jordan, Syria, Turkey,
serves as the global focal point for nuclear co-operation,
Ethiopia, Uganda and Iran; (ii) Tunisia, Morocco and
mobilising peaceful applications of nuclear science, and
Algeria; (iii) Pakistan and South Africa. This list is not
technology for critical needs in developing countries,
exhaustive, and notes of interest are welcome. Tentatively,
including fighting hunger, disease, poverty and
operative co-ordination shall include: The IAEA
pollution of the environment, thereby contributing to
(Vienna, Austria: scientific backstopping, Seibersdorf
sustainable development goals of its Member States. The
Laboratories, Austria: quality control/training) in
IAEA currently co-ordinates research networks (CRPs) [2]
close collaboration with the International Center for
and supports human and institutional capacity building
Agricultural Research in the Dry Areas (ICARDA, Syria),
Technical Cooperation Projects (TCPs) [3] for integrating
tentatively proposed as the project coordinator. The
efficiency enhancing bio- and molecular technologies
FAO (Rome) would be entrusted with the normative co-
with mutation induction within the framework of
ordination. The international collaborating laboratories
national plant breeding and conservation programmes
and institutes include: ICARDA, the International Maize
to characterize plant genetic resources and widen
and Wheat Improvement Center (CIMMYT), the United
plant genetic diversity, and to identify and introduce
States Department of Agriculture – Agricultural Research
agronomically and commercially useful traits.
International Atomic Energy Agency, Joint FAO/IAEA Division, Plant Breeding The IAEA Program in Food and Agriculture is
and Genetics Section, A2256, Wagramerstrasse 5, PO BOX 100, A-1400 Vienna, planned, implemented and co-financed with FAO and
Austria
E-mail: p.lagoda@iaea.org is known as the Joint FAO/IAEA Program. As such, its
206 Responding to the transboundary threat of wheat stem rust (race Ug99)
lies as well in their unique sensitivity and specificity as above thematic areas, especially animal and crop pests
markers. They can be used to measure - more accurately and diseases are transboundary in nature and require
than is possible by any conventional method - basic and an area-wide approach to be managed successfully.
yet strategically essential processes which take place Regional collaboration is therefore necessary and
within and between soils, plants, and animals. Third, collaboration between international organizations is
radiation can be effectively applied for sanitary and best positioned to coordinate these activities.
phytosanitary purposes in support of food safety and The Joint FAO/IAEA Program is the only
to facilitate international agricultural trade, as well as international body that can provide technology
for specialized and successful applications such as the development and transfer, capacity building and
sterile insect technique, where leadership lies with the services in this area of nuclear applications in food and
Joint FAO/IAEA Program. agriculture to the Member States, and is in this respect
Through IAEA-Technical Cooperation (TC) funding, it is unique.
the Joint FAO/IAEA Program provides technical support
to more than 250 IAEA-TC projects every year, as well Nuclear applications in food and agriculture
as capacity building and technology transfer (expert What are nuclear techniques?
advice, training, and assisting with the procurement Everything in the Universe, including the soil,
of experts and equipment) to Member States through plants and animals that we use for agriculture, and the
these technical cooperation projects. Over the carbohydrates, proteins and fats in the food we eat
past decade, the Joint Program added each year to is made up of around 100 elements. These elements
capacity building through over 50 training courses and consist of atoms with a nucleus composed of neutrons
workshops, 350 fellowships and scientific visits. and protons surrounded by electrons. However, not all
Through the regular budget, the Joint Program atoms of an element have the same number of neutrons
organizes symposia, conferences, consultants meetings, in their nucleus, i.e. they exist in different isotopic forms
interregional training courses and workshops, - some are heavier than others, some are stable, and
provides normative and policy advice, disseminates others undergo decay and emit energy as radiation.
information through databases, e-learning modules, Applications of nuclear techniques in food and
and web pages. It also assists Member States through agriculture make use of isotopes to measure and track
a network of coordinated research projects (CRP) and with great accuracy and precision, various events
research coordination meetings (RCM) to address occurring in agriculturally important processes and
specific practical problems related to a range of areas: compounds, and to manipulate those processes for
approximately 400-500 institutions and experimental greater productivity. They also make use of sealed
stations in Member Countries cooperate in 30-40 R&D facilities containing radiation- emitting isotopes to
networks (CRPs) per year organized by the Joint FAO/ mimic Nature in changing the genetic make-up of
IAEA Program. plants, insects and micro-organisms in order to produce
The IAEA is the only organization within the UN better crops, sterile insects for controlling pests and
family that has the mandate to promote the peaceful increasing the shelf-life and safety of certain foods.
use of nuclear techniques. In some of the agricultural Nuclear techniques, combined with the application
areas, nuclear techniques are an essential component of modern bio- and molecular technologies, are
and when properly integrated with other conventional essential to providing a more efficient way, both
and modern technologies, provide substantial added for understanding the processes that underpin the
value to national and international efforts to sustainable production and transformation of biophysical resources
agricultural development while at the same time into food and agricultural products and, directly or
creating strong synergies. indirectly, for manipulating these processes to increase
By the use of nuclear techniques, the Joint FAO/ crop and livestock productivity while conserving and
IAEA Program provides unique support not only to FAO sustainably using natural resources and improving food
but also to other international bodies in their efforts quality and safety. The effective transfer of existing
to enhance food quality and safety, protect consumer nuclear techniques to developing countries and the
health and facilitate international trade in foodstuffs. development of new and safe bio- and molecular
All major activities of the Joint Program are within the technologies combined with nuclear techniques
‘public good’ area both in developing and developed can greatly enhance the prospects for sustainably
countries and address urgent needs and requirements improving agricultural productivity, both currently, and
from FAO and IAEA Member States. In addition, many in the future.
constraints to agricultural development related to the
208 Responding to the transboundary threat of wheat stem rust (race Ug99)
The economic value of a new variety can be molecular technologies such as plant tissue culture
assessed in several ways. These include area planted to and molecular markers plays a very important role in
the variety and percentage of the area under the crop in crop improvement. Mutation induction is an integral
a region, increased yield, enhanced quality, or reduced part of the newest technology package in the forefront
use of pesticides and fungicides (e.g. in varieties of modern and efficient methods in reverse genetics
resistant to diseases and insect pests). But to make a and breeding: e.g. TILLING (targeting induced local
long and complicated story short, the socio-economic lesions in genomes), and breeding for modified starch
impact of mutant varieties is assessed in billions of quality traits in hexaploid wheat. Mutation induction is
dollars and millions of hectares cultivated (Ahloowalia producing mutation grids for gene discovery and gene
et al. 2004). function analyses (e.g. Arabidopsis, rice, barley), an
Many mutants have made transnational impacts on invaluable resource for genomics, and both reverse and
increasing yield and quality of several seed propagated forward genetics.
crops. Induced mutations will continue to have an In recent years there has been increased interest in
increasing role in creating crop varieties with traits understanding the genome. This goes in parallel with
such as modified oil content, protein and starch quality, the explosion of fundamental and strategic research
enhanced uptake of specific metals, deeper rooting to understand gene structure and function, especially
systems, and resistances to drought, diseases/pests in crop and model plants. The IAEA Plant Breeding and
and salinity as major components of environmentally Genetics section and laboratory unit are adapting the
sustainable agriculture. Future research on induced TILLING strategy to the peculiarities of tropical orphan
mutations will also be important in the functional crops. In addition to the work on the relatively more
genomics of many food crops. studied crop, rice, the Joint Program has made significant
The Agency has addressed the problems of climate progress in the development of protocols, i.e. simplifying
variability and change, disease resistance (including procedures and exploring low cost options, to facilitate
resurgence, and appearance of transboundary threats the use of TILLING to routinely query genomes of the
due to climate change), drought and salinity stress scantily studied polyploid and vegetatively propagated
tolerance to improve the nutrition provided by the crops that are important to the food security and
plants and their resistance to specific environmental livelihoods of Member States, such as cassava and
and geographical problems. Up to 80% of plant yields bananas, thus creating an invaluable resource for reverse
can be lost because of drought and salinity. Problems genetics and breeding for the global community. The
are particularly severe in developing countries in arid widespread routine adoption of TILLING, for instance, will
and semi-arid regions, with both devastating short- significantly reduce the costs and time invested in the
term effects on the livelihoods of poor people and development of superior crop varieties.
long-term effects on food security. These are likely to The Green Revolution was driven by spontaneous
increase in future as competition for water resources mutations affecting plant height. Two homeoalleles
increases. The integration of mutation induction and of the semi-dwarf gene in wheat, Rht1 and Rht2, and
efficiency enhancing bio-/molecular technologies into Sd1 in rice, produced the semi-dwarf stature that had
plant breeding and adoption of advanced selection implications for lodging resistance. The beneficial effects
methods can lead to the official release and wide uptake of spontaneous mutations continue to be reproduced in
by farming communities of new varieties of basic food more controlled and systematic ways using mutagenic
and industrial crops that are higher yielding, have better agents. In fact, the semi-dwarf characteristic was
quality, are more nutritious and better adapted to artificially induced at about the same time as “mystic”
climate change and variability. semi-dwarfing genes were being discovered, and it
It is noteworthy, that worldwide, more than 60% of would be later shown that the “natural” spontaneous
all mutant varieties were officially released after 1985, and “artificial” induced mutations both affected the same
during the era of biotechnology in plant breeding. The gene and were thus alleles.
integration of mutation techniques and efficiency- The use of induced mutations in crop improvement,
enhancing bio-/molecular techniques that permit rapid started over 80 years ago, continued to be massively
selection of the most beneficial mutants has pushed used by plant breeders until the 1980’s when the advent
the use of mutation induction to new and higher levels of modern techniques of molecular biology and the
of applicability. With the integration of molecular potential to move specific genes from one organism
genetic information and techniques, mutation assisted to another seemed to be more viable alternatives.
breeding is in the mainstream of progress to develop Consequently, research on mutation induction for
novel varieties. Mutation induction combined with bio-/ breeding of crops declined, especially for sexually
210 Responding to the transboundary threat of wheat stem rust (race Ug99)
accept rust samples and to culture them under very training on the genetic bases of changes in virulence
strict conditions (timing/period and method of sending and rust epidemiology, and in addition, should receive
the material). This procedure is time consuming, and some field experience so that they know what they are
several of the samples sent may lose viability by the working on and how to interpret results.
time they are tested. Thus, precious time may be lost
before results are obtained and action in the field is Provision of genetic resources
undertaken. It is accordingly highly important that The genetic improvement of plants is dependent
national and/or regional laboratories within the high risk on the availability of useful and exploitable genetic
regions are established or upgraded and personnel are variation within the genepool accessible for
trained in such a way that rust pathotyping takes place manipulation by the plant breeder. Such variation
immediately in the country and without further risks of arises naturally through spontaneous mutations and
spreading the pathogen to other regions. Thus, testing hybridisations between wild and closely related species.
should be done nationally, and ideally only re-confirmed Mutation induction, artificial changes to the genetic
in the international laboratories on a needs basis. make-up of an organism generating variations in
The Interregional Project should therefore potential parental materials is thus a facilitation through
provide a platform for the coordination of a network artificial means of an otherwise natural phenomenon.
of laboratories as a “defense line” against the disease, Plant breeders engaged in the development of new
implementing the following three phases: superior varieties exploit such variations when they
Phase I: “Normalizing” adoption and training in the are useful. In rare cases, the mutants possess traits
use of uniform protocols, in order to assure of agronomic or economic importance to such an
homogeneity of handling (capacity building) extent that they require little or indeed no further
Phase II: Quality Control: double blind tests manipulation before being released to farmers. Most of
for performance of identification and the time however, the mutants are just “raw materials”
characterization false positives vs. false (pre-breeding material) that must be included in a
negatives (periodic network performance normal varietal development mechanism. This would
meetings, quality management and steering) normally involve controlled crosses with otherwise
Phase III: Multilocation trials of mutant germplasm in well established varieties which lack the desirable trait
endemic hotspots/screen houses identified in the mutant, followed by several cycles of
These phases are not necessarily consecutive, but field evaluation.
overlapping. All the listed countries in the project have Agency
supported mutation assisted breeding programs
Network of Ug99 surveillance laboratories: in wheat and/or barley. Through mutation assisted
1. (Front line): Kenya, Yemen, Sudan, Egypt, Jordan, breeding, these member countries develop directly
Syria, Turkey, Ethiopia and Iran new high yielding cultivars with good agronomic
2. Tunisia, Morocco, Algeria characteristics, well-adapted and high value-added traits
3. Pakistan, South Africa. from any germplasm source including local landraces,
This list of countries is by no means exhaustive, and which is difficult or impossible to attain through
notes of interest are encouraged [7]. The First Coordination conventional plant breeding. This helps to enhance crop
and Steering Meeting is planned for the first week in May, production for food security, increases farmer income,
2009, at IAEA Headquarters in Vienna, Austria. conserves biodiversity and enhances agro-biodiversity,
• Normative Coordination: FAO (Rome) thus directly contributing to the conservation and use of
• Operative Coordination: IAEA (Vienna: Scientific plant genetic resources.
Backstopping, Seibersdorf: Quality Control/ Listed in the FAO/IAEA Database on Mutant
Training) Varieties and Genetic Stocks (MVGS) [5], there already
• Project Coordination: Ideally through ICARDA are 235 officially released mutant wheat and 304
• International Collaborating Laboratories/Institutes: officially released mutant barley varieties. One percent
ICARDA, CIMMYT, USDA-ARS of these officially released mutant varieties were created
The training component of this project could be for rust resistance (2 wheat and 3 barley).
provided by any competent laboratory, but it should be Using mutation induction techniques,
group training to warrant homogeneity of procedures. an abundance of more or less advanced and
However, besides the hands-on training, personnel characterized mutant lines have been created through
who will be undertaking pathotyping in the national different Agency sponsored cooperation projects,
laboratories should have additional, complementary providing breeding material for conventional plant
212 Responding to the transboundary threat of wheat stem rust (race Ug99)
33. The UN-FAO Wheat Rust Disease stress, especially in rainfed regions, the impact of
diseases is also expected to increase, resulting in severe
Global Program yield losses.
A major new threat has already surfaced, and that
Wafa El Khoury1 is the widely virulent stem rust race Ug99, also known
as TTKSK (Roelfs and Martens 1988; Fetch et al. these
Abstract proceedings). It appeared in East Africa in 1999, and by
The FAO Wheat Rust Disease Global Program was late 2007, had reached Iran. Ug99 is highly virulent on
launched to respond to the emergence and spread of almost all wheat varieties currently grown throughout
the virulent strain of wheat stem rust, Ug99, and similar the world, and the risk that it could cause global
virulent strains in the future. It is a part of FAO’s Crisis epidemics is very real. If this happens, wheat production
Management Centre for the Food Chain and it reinforces will suffer devastating yield losses.
and complements activities of the Borlaug Global Rust From previous experience with similar rust strains (a
Initiative. The Program aims at contributing to global recent example is the breakdown of the stripe (yellow)
food security through prevention and management of rust resistance gene Yr9 causing widespread epidemics
emerging wheat rust races and the enhancement of between 1986 and 1998; Singh et al. 2004), and based
wheat productivity. It covers the following components: on weather patterns, it is likely a matter of only a few
1) national preparedness and contingency planning; 2) years before most countries in the Near East, East and
surveillance and early warning; 3) national wheat varietal North Africa, Central and South Asia cultivating around
registration programs; 4) national systems for quick seed 80 million hectares of wheat (FAOSTAT 2007) will be
multiplication and distribution; and 5) improvement of affected by Ug99 (Fig. 1; Wheat Rust Global Program,
wheat rusts field management. As a neutral information ftp://ftp.fao.org/docrep/fao/011/i0378e/i0378e.pdf ).
sharing forum, FAO is well positioned to lead such global Agricultural crises resulting from transboundary
efforts through linkages with national governments, pests and diseases are often the result of unsound
regional bodies, farmers, international research and agriculturally related policies, mismanagement of
development institutions, and the donor community. resources, and poor national and regional preparedness
for prevention and early response measures. For
Keywords too many years, governments have reduced their
Ug99, wheat stem rust, contingency planning, investments in agriculture, especially with respect to
transboundary plant diseases farmer education and extension, support to agricultural
research, rural development through infrastructure, and
Introduction market access.
Wheat is grown on more than 200 million hectares Through its Wheat Rust Disease Global Program
and is a source of food and livelihoods for over a (WRDGP), the Food and Agriculture Organization of
billion people in developing countries. The Near East, the United Nations (FAO) is taking global action to
East and North Africa and Central and South Asia prevent a wheat production crisis, in close collaboration
alone account for some 37 percent of global wheat with national governments, international agricultural
production (FAOSTAT 2007). In most countries in these research centers and other international institutions,
regions, wheat is the staple food crop, providing on to manage the Ug99 threat, and to prevent future
average some 40% of the per capita calorie supply, crises caused by similar wheat rust diseases. The main
and is especially important in the diets of the poorest. objective of the Program is to contribute to global food
Many people in these countries heavily rely on wheat security through the prevention and management of
production for their subsistence and livelihood. This emerging wheat rust diseases and the enhancement of
livelihood has been greatly affected by the rise in grain wheat productivity (Wheat Rust Disease Global Program,
prices which the world has witnessed over recent years. ftp://ftp.fao.org/docrep/fao/011/i0378e/i0378e.pdf ).
The rise in prices, largely the result of several years of FAO is well positioned to lead such international
severe drought, the high cost of fuel and an increased efforts because of its experience with international
demand for grains, is not the only problem they face. dimensions of other transboundary pests, such as
With wheat crops coming under pressure from climatic locusts, its standing as a neutral international forum for
information sharing, its experience in the emergency
response and its linkages with grassroots rural
1
Plant Production and Protection Division, Food and Agriculture Organization communities, national governments, regional bodies,
of the United Nations; Rome, Italy
E-mail: wafa.khoury@fao.org international agriculture research and development
institutions, the private sector and the donor community. was established to respond to the recent increases
Within FAO, the WRDGP is part of the newly in the number of outbreaks of transboundary animal
established Crisis Management Centre for the Food diseases, plant pests and food safety emergencies.
Chain (CMC-FC) and works in close collaboration with Changing agro-ecological conditions, intensifying
the Initiative for the Soaring Food Prices. food production systems and expanding global trade
Globally, the WRDGP works in full partnership increase the likelihood of animal and plant diseases
with FAO’s Member Countries and in full synergy and and pests emerging and spreading farther and
coordination with the Borlaug Global Rust Initiative faster than ever before, and for unsafe food to reach
(BGRI)2. First established in 2005 as the Global Rust numerous consumers in distant markets. With the
Initiative (GRI), this initiative was later expanded to what advent and spread of instant mass communication,
is now known as the BGRI and includes in addition to the news of outbreaks can cause generalized consumer
International Center for Agricultural Research in the Dry panic, market collapse and serious economic damage
Areas (ICARDA) and the International Maize and Wheat in regions well beyond affected areas. The CMC-FC
Improvement Center (CIMMYT), Cornell University and reflects FAO’s determination to address the risks to the
FAO as permanent members and Dr Norman Borlaug human food chain in their assessment, management
as the Chairman of its Executive Committee. The overall and communication dimensions in a comprehensive,
objective of BGRI is to “systematically reduce the world’s systematic, inter-disciplinary, institutions-wide,
vulnerability to stem, yellow and leaf rusts of wheat, collaborative approach. Recent external evaluations of
through advocating and facilitating the evolution of a FAO have highlighted the Organization’s comparative
sustainable international system to contain the threat advantage in this domain.
of wheat rusts and consolidating the enhancements in The CMC-FC acts as a broad-based international
productivity required to withstand future global threats center established to bring together the entire food
to wheat” (Sounding the Alarm on Global Stem Rust, chain and to develop activities that allow for the
2005; www.globalrust.org). forecasting, prevention and management of threats that
go beyond national borders.
FAO’s Crisis Management Center for the The organizational structure of the CMC-FC (Fig.
Food Chain 2) foresees three units reflecting various levels of
The Wheat Rust Disease Global Program falls within activities; these are 1) Intelligence and Coordination
the scope of the newly established Crisis Management unit, 2) Prevention and Early Warning unit, and 3) Rapid
Centre for the Food Chain (CMC-FC). The CMC-FC Response unit. The Prevention and Early Warning unit
is provided by the previously established Emergency
2
Named after its chair, U.S. agronomist Dr Norman Borlaug, Nobel Peace Prize
winner in 1970 and widely acclaimed as the “father of the Green Revolution” Prevention System for Transboundary Animal and Plant
Fig. 2 Structure of FAO’s Crisis Management Center for the Food Chain
1
University of Aarhus, Faculty of Agricultural Sciences, Department of
Integrated Pest Management, Flakkebjerg, 4200 Slagelse, Denmark; 2
International Center for Agricultural Research in the Dry Areas, PO Box 5466,
Aleppo, Syria; 3 The International Maize and Wheat Improvement Center
(CIMMYT), AP 6-641, 06600 Mexico
1
This is proposed as a joint work involving national programs in targeted
countries, ICARDA, Virginia Tech University, CIMMYT and FAO.
P.O. Box: 5466, Aleppo- Syria.
E-mail: k.shideed@cgiar.org
1
CIMMYT, Apdo. Postal 6-641, 06600, Mexico, DF, Mexico; 2ICARDA, PO Box
5466, Aleppo, Syrian Arab Republic
Agricultural Research center, P.O. Box 489, Addis Ababa, Ethiopia; 3Sinana
“horizontal” resistance to attain durable stem rust control. Agricultural Research Center, P.O. Box 208; Addis Ababa, Ethiopia; 4Debre Zeit
Agricultural Research Center, P.O. Box 32, Addis Ababa, Ethiopia; 5Holetta
1
Plant Protection Research Center, Ethiopian Institute of Agricultural Research, Agricultural Research Center, P.O. Box 2003, Addia Ababa, Ethiopia; 6Sirinka
P.O. Box 37, Ambo, Ethiopia;2 Julius Kuehn-Institute, Federal Research Institute Agricultural Research Center, P.O. Box 74, Bahir Dar, Ethiopia; 7Awassa
for Cultivated Plants (JKI), Institute for Resistance Research and Stress Tolerance, Agricultural Research Center, P.O. Box 6, Awassa, Ethiopia; 8Debre Birhan
Erwin-Baur-Str. 27, 06484 Quedlinburg, Germany; 3 Justus-Liebig-University Agricultural Research Center, P.O. Box 112, Debre Birhan, Ethiopia; 9Adet
Giessen, Department of Plant Breeding, Heinrich-Buff-Ring 26-32, 35392 Agricultural Research Center, P.O. Box 8, Bahir Dar, Ethiopia; 10Haramaya
Giessen, Germany University, P.O. Box 138, Dira Dawa, Ethiopia
1
USDA-ARS Wheat Genetics, Quality, Physiology, and Disease Research Unit;
2
Department of Plant Pathology, Washington State University, Pullman, WA
99164-6430, USA.
E-mail: xianming@wsu.edu All-Russian Research Institute of Biological Plant Protection, Krasnodar, Russia
The rusts are major diseases of wheat in Turkey CIMMYT-derived durum wheat (Triticum turgidum
and they can cause significant yield losses in years with var durum) germplasm was highly resistant to leaf rust
suitable conditions. However, rust prevalence changes (caused by Puccinia triticina) to prevalent race BBB/BN
from year to year and from region to region depending in Mexico until 2000. However, a large portion of the
on climatic conditions. This study was conducted to germplasm was susceptible in Chile and North Africa.
monitor the occurrence of rusts in different parts of A new race, detected in northwestern Mexico in 2001,
Turkey in 2008. Survey trips were conducted covering was virulent on more than 80% of the germplasm,
the Marmara, Aegean, Thrace, East Mediterranean, including the most popular cultivar Altar C84. This race
Southeast Anatolia, Central Anatolia, East Anatolia and was designated BBG/BN. Apparently a single gene
Mid-Blacksea regions. Two hundred and forty two wheat mutation towards virulence on Lr11 was observed, but
fields were examined for the presence of stripe rust, leaf virulence to the undesignated gene in Altar indicated
rust and stem rust. The frequencies of infected plants the possibility of an exotic origin. During the same year
were recorded and severities were estimated using the a variant, designated as BCG/BN, was identified with an
Modified Cobb scale. Seventy one fields were infected unnecessary virulence for resistance gene Lr26 present
with rusts. Of these, 60 were infected with stem rust, in the 1B.1R translocation in bread wheat (T. aestivum).
6 with leaf rust, and 9 with stripe rust. In some fields, In 2008 leaf rust was observed on previously resistant
more than one rust was present. In 2008 Turkey suffered durum cultivars Jupare C2001 and Banamichi C2004.
from severe drought which was so severe that some Single pustule isolates indicated the presence of a
fields were not harvested. Stem rust was most prevalent new race designated BBG/BP, which evolved through
in inner parts of Black Sea region. Severities of rust a single mutation in race BBG/BN for virulence to the
diseases were therefore non-significant. However, their complementary resistance genes Lr27+Lr31 present in
occurrences under such dry conditions indicate that Gatcher, Jupare C2001 and Banamichi C2004, and adult
they keep their potential to cause severe losses. plant resistance gene Lr12. A variant isolate of race BBG/
BP, designated as CBG/BP, with additional virulence
This study was conducted as part of the project for Lr3 present in CIMMYT durum ‘Storlom’ was also
‘Determination of Races of Wheat Stem Rust (Puccinia identified. Although virulence to Lr3, Lr12 and Lr27+Lr31
graminis f. sp. tritici) and Resistant Wheat Genotypes is known to occur in P. triticina races predominant on
Against Common Races in Turkey, No:106O331’ financed bread wheat, this is first time that we identified such
by the Scientific and Technical Research Council of virulences in races predominant on durum wheat.
Turkey (TUBITAK). Since the introduction of BBG/BN in Mexico in 2001,
this durum P. triticina race has continued to evolve and
defeat race-specific resistance genes commonly present
in both durum and bread wheat.
1
INIFAP-CEVAMEX, Apdo. Postal 10, 56230, Chapingo, México; 2International
Maize and Wheat Improvement Center (CIMMYT), Apdo. Postal 6-641, 06600
Central Research Institute for Field Crops, P.O.B. 226, Ulus, Ankara, Turkey;
1
México D.F; 3Junta local de Sanidad Vegetal de Huatabampo, Ocampo y
Ankara University, Faculty of Agriculture, Department of Plant Protection,
2
Ferrocarril, S/N Huatabampo Sonora, México; 4INIFAP-CEVY, Apdo. Postal 515,
Dışkapı, Ankara, Turkey 850000, Cd. Obregón, Sonora, México
1
Wheat Rust Laboratory, DWR Regional station, Flowerdale, Shimla; India; 1
INRA – CRRA, PO Box 578, Meknès, Morocco; 2INRA – CRRA, PO Box 589,Settat,
2
Directorate of Wheat Research, Karna, India Morocco; 3ICARDA, Rabat Institutes, PO Box 6299, Rabat, Morocco
1
Department of Plant Pathology and 2USDA-ARS Cereal Disease Laboratory, St.
Paul, MN 55108, U.S.A.; 3USDA-ARS Wheat Genetics, Quality, Physiology, and
INRA Morocco, Ville Nouvelle, BP Meknès, Morocco Disease Research Unit, Pullman, WA 99164-6430, U.S.A.
1
Uzbek Research Institute of Plant Industry, Uzbekistan; 2 Institute of Genetics
and Plant Experimental Biology, Uzbekistan; 3CIMMYT Global Wheat Program,
Turkey; 4ICARDA-Central Asia and the Caucasus Regional Program, Uzbekistan
HS Bariana, UK Bansal, H Miah, AK Toor, Stem rust caused by Puccinia graminis f. sp. tritici
F Hussain, RF Park race TTKS commonly known as “Ug 99” is becoming a
serious threat to wheat production worldwide. To cope
The pathotype ‘Ug99’ of the wheat stem rust up with the rapidly changing stem rust pathogen, new
pathogen was first detected in Uganda in 1999. Since sources of seedling and adult plant resistances might be
its first detection, it has produced variants with added sought from the wild relatives of cultivated tetraploid
virulence for Sr24 and Sr36. A strategic global effort was wheat. A total of 1,524 wild tetraploid wheat accessions
undertaken to tackle this menace through deployment were evaluated against the prevailing Syrian stem rust
of genetic resistance in new wheat cultivars. The population under field conditions at the International
identification and characterisation of diverse sources Center for Agricultural Research in the Dry Areas
of resistance is essential to combat the threat posed (ICARDA), Tel Hadya, Aleppo, Syria.
by new variants of pathogens. We studied genetic Two hundred and thirty eight accessions with adult
variation for stem rust resistance among the AE Watkins plant resistance were selected for further seedling and
collection of hexaploid and tetraploid wheat genotypes. adult plant assessments at the Debre Zeit Research
A specific attempt was made to identify new sources Center, Ethiopia; a reputed ‘hotspot’ site for stem rust
of durable minor gene controlled adult plant stem epidemics on tetraploids. The accessions were exposed
rust resistance. Tests on these genotypes under field to a mixture of isolates comprising Ug99 and a local
conditions, followed by seedling tests with the same bulk of urediniospores collected from hexaploid and
pathotype (s) of the stem rust pathogen, indicated tetraploid wheats. About 37% and 36% of the accessions
the presence of minor (non- hypersensitive) genes for showed resistance to stem rust at the seedling and
resistance in both hexaploid and tetraploid genotypes. adult growth stages, respectively. About 15% exhibited
Genotyping using Sr2-linked molecular markers enabled resistance at both the seedling and adult plant stages,
identification of genotypes that lacked Sr2 and carried as leaving 21% with adult plant resistance only. This
yet uncharacterised adult plant resistance gene(s). These preliminary result indicated that wild tetraploid wheats
putative new sources of resistance were crossed with could be potentially important sources of resistance to
susceptible cultivars to develop mapping populations the prevailing stem rust races including Ug99. Some
for genetic characterisation of the resistance. Bulked accessions have been selected for repeat testing to
segregant analysis will be performed to identify confirm the results. Crosses between these and elite
genomic regions that control adult plant resistance in bread wheat and durum varieties have also been
some selected genotypes. initiated. Further ongoing genetic and genomic studies
using these accessions should identify and characterize
the resistance genes and reveal potentially new stem
rust resistance genes for deployment in both durum and
bread wheat breeding.
1
Aleppo University, Faculty of Agriculture, Field Crops Department, Aleppo,
Syria; 2International Centre for Agricultural Research in the Dry Areas, PO Box
The University of Sydney Plant Breeding Institute-Cobbitty, PMB11, Camden, 5466, Aleppo, Syria; 3Ethiopian Institute of Agricultural Research (EIAR), Box
NSW2570, Australia 2003, Addis Ababa, Ethiopia
Twenty-four leaf rust resistant T. aestivum x T. The threat that race TTKSK (Ug99) poses to
timopheevii hybrid lines were developed using five wheat worldwide is well known and documented.
common wheat cultivars. The resistances were analyzed However, this race also threatens barley throughout the
using microsatellite markers specific for T. aestivum world, including those cultivars carrying the durable
and T. timopheevii. Microsatellite analysis revealed rust resistance gene Rpg1. To identify and map loci
two major areas of introgression of the T. timopheevii conferring resistance to race TTKSK, we are using an
genome: chromosomes of homoeologous groups 2 association mapping approach in both cultivated (Barley
and 5. Translocations were detected in the 2A and 2B Coordinated Agricultural Project or BCAP) and wild
chromosomes in 11 lines. The length of the translocated (Wild Barley Diversity Collection or WBDC) Hordeum
fragment in the 2B chromosome was identical in all germplasm. BCAP accessions were genotyped with
hybrid lines and did not depend on the parental wheat 1,536 SNP markers and WBDC with 3,072 SNP and 558
variety. DArT markers. Marked variation in the germplasm was
The hybrid line 842-2 was used for detailed observed in response to race TTKSK at the seedling
characterization of introgression and mapping of loci stage, with some accessions exhibiting a high level
determining resistance to leaf rust. Molecular analysis of resistance. Association mapping analyses of BCAP
using 350 specific short sequence repeat (SSR) markers germplasm identified resistance QTL on chromosomes
identified genes from the T. timopheevii genome in 1H, 2H, 3H, 5H and 7H (p=2.01E-07 to 8.00E-04, r2=1.4
chromosomes 1A, 2A, 2B, 5A, 5B, and 6B. An F2 mapping to 2.4%). The QTL on chromosome 5H was coincident
population of line 842-2 crossed with common wheat with the previously identified resistance gene complex
cultivar Skala was used for analysis of association of rpg4/Rpg5. In the WBDC germplasm, QTL for resistance
phenotypic and genotypic data. Adult plant leaf rust were identified on all seven chromosomes (p=0.000
resistance was determined by loci in chromosomes 5B to 0.002, r2=2.9 to 7.4%). Several identified QTL on
and 2A. The major locus transferred from T. timopheevii chromosomes 5H and 7H were coincident with those
chromosome 5G mapped to the microsatellite interval found in the same region of the BCAP germplasm.
Xgwm408 – Xgwm1257 and controlled 72% of the Additionally, QTL were found coincident with both Rpg1
phenotypic diversity in leaf rust response. The other, on chromosome 7H and rpg4/Rpg5 on chromosome 5H.
less effective gene was located on chromosome 2A at a This work documents of power of association mapping
distance of 10 cM from Xgwm312, and accounted for 7% for identifying and mapping stem rust resistance loci in
of the trait expression. Microsatellite markers located cultivated and wild Hordeum germplasm.
near these loci may be used for the transfer of these
valuable genes to new lines and cultivars.
1
Institute of Cytology and Genetics, Siberian Branch, Russian Academy of
Sciences, Novosibirsk, 630090 Russia; 2Leibniz Institute of Plant Genetics and Department of Plant Pathology, University of Minnesota, St. Paul, MN 55108,
1
Crop Research, Gatersleben, D-06466 Germany USA; 2USDA-ARS Cereal Disease Laboratory, St. Paul, MN 55108, USA
1
Department of Crop and Soil Science, Washington State University, Pullman,
WA 99164, USA2. Department of Plant Pathology, and 3USDA-ARS Cereal
Disease Laboratory, St. Paul, MN 55108, USA Crop Protection, Directorate of Wheat Research, Karnal (Haryana)-132001, India
1
International Maize and Wheat Improvement Center (CIMMYT), Apdo. Postal
6-641, 06600 México, D.F., México; 2Campo Experimental Valle de México INIFAP, 1
Department of Plant Pathology, University of Minnesota, and 2 USDA-ARS,
Apdo. Postal 10, 56230 Chapingo, Edo de México, México Cereal Disease Laboratory, St. Paul, MN 51108, USA
Wheat leaf rust can be controlled by host resistance. One of the best approaches to alleviate the threat
Relatives and progenitors of wheat have been abundant from Puccinia graminis tritici race Ug99 (TTKSK) is to
sources of leaf rust resistance (Lr) genes. Effective Lr identify and characterize sources of resistance within the
genes were transferred from Aegilops speltoides to available wheat (Triticum aestivum) breeding materials
wheat by J. Dvorak and D. Knott. Subsequently, P. Dyck and commercial cultivars. Genes identified can then be
produced a near-isogenic line (RL6161) carrying this deployed in combinations. Identification of molecular
gene in a Thatcher background. To further characterise markers tightly linked to resistance genes can aid their
the resistance in RL6161, agronomic, quality and pyramiding, and allow selection of plants without the
genetic tests were undertaken. Compared to the need for disease screening. This is especially important
recurrent parent (Thatcher), RL6161 showed no penalty with Ug99 and its derivatives, which are absent in many
in yield or quality that sometimes accompanies alien countries. F3 and F4 populations derived from the crosses
transfers. Monosomic analysis placed the Lr gene of susceptible PBW343 with three resistant parents
on chromosome 1B. A doubled-haploid population with race-specific resistance genes were developed and
from the cross Thatcher / RL6161 was tested with characterized for reaction to TTKSK in the greenhouse
microsatellite markers specific to chromosome 1B and at USDA-ARS CDL, St. Paul, MN; and, during 2008,
the results showed that the Ae. speltoides DNA carrying in the field at Njoro, Kenya, where the Sr24 virulent
the Lr gene was linked to markers on the long arm. Ug99 variant was present. Bulk-segregant analysis was
Preliminary mapping data showed that recombination performed to identify marker trait associations and the
occurred between the Ae. speltoides and wheat DNA. linked markers were used for genotyping lines clearly
Therefore, lines with reduced introgression size can be identified in field trials as homozygous resistant and
identified and used as sources of resistance in breeding homozygous susceptible. Genomic regions with 3
populations. Whereas the uniqueness of the resistance putative new resistance genes, temporarily designated
in RL6161 is not known, it is possible that the resistance as SrA, SrB and SrC were identified. Gene SrA was
gene is Lr51, or an allele, since Lr51 was also transferred mapped on chromosome 3DL (linked markers, Xgwm52,
from Ae. speltoides to wheat chromosome 1BL. Xgwm341) of Milan/Sha7/3/Thb/CEP7780//Sha4/Lira/4/
Experiments to demonstrate the relationship between Sha4/Chil, SrB on chromosome 3BS (Xgwm566, Wmc231)
the two resistance sources are in progress. of Ning9415/3/Ures/Bow//Opata/4/Ningmai 7, and SrC
on chromosome 5DL (Xgwm292, Xgwm212) of Chen/
Ae.Sq//2*Weaver/3/Oasis/5*Borl95. Like several other
characterized stem rust resistance genes, the three new
resistance genes provide moderate levels of resistance
at the seedling and adult stages. Further studies to
confirm the results and development of targeted
mapping populations to identify closely linked markers
are under progress.
1
CIMMYT, Apdo. Postal 6-641, 06600, Mexico, DF, Mexico; 2INIFAP-CEVAMEX,
Apdo. Postal 10, 56230 Chapingo, Mexico, 3USDA-ARS, CDL, St. Paul, MN 55108,
Cereal Research Centre, 195 Dafoe Road, Winnipeg, MN R3T 2M9, Canada USA
GF Marais, L Kotze, A Eksteen Wheat stem rust, caused by Puccinia graminis f. sp.
tritici, has been effectively controlled through the use
The wild relatives of wheat constitute a valuable
of genetic resistance. The recently identified race TTKSK
source of rust resistance genes that can be utilized in
(Ug99) possesses virulence to many resistance genes
breeding. Translocation of desirable genes from wild
that have been used in wheat breeding worldwide. One
species inevitably results in co-transfer of un-needed
strategy to aid breeders in developing resistant varieties
alien chromatin. The Lr59 translocation appears to
is to provide resistance genes transferred from wild
involve the complete long arm of chromosome 1A.
relatives to wheat. Stem rust resistance genes Sr22 and
An attempt was made to replace some of the Aegilops
Sr35, derived from Triticum monococcum are effective
peregrina chromatin with wheat chromatin through
against race TTKSK. In order to identify additional genes
induction of homoeologous chromosome pairing
from this relative of wheat, we screened 1,062 accessions
by deleting Ph1. Resistant testcross F1 plants were
of T. monococcum deposited in the National Small Grains
characterized for the presence of three mapped wheat
Collection against TTKSK and two additional races with
microsatellite loci and a newly discovered SCAR locus
broad virulence. We identified 625 accessions (58.85%)
that maps to the Lr59 translocation. Within the mapped
with resistance to TTKSK with infection types ranging
region primarily single crossovers occurred, as expected
from 0 to 2+. Among these resistant accessions, 90
with homoeologous chromosome pairing. Overall, the
accessions (8.47% of the total) were also resistant to
recombination data were reflective of comparatively
TTTTF and TRTTF. Results from the preliminary screening
regular pairing within a highly homoeologous
suggested that new resistance genes are likely to be
chromosome region. Strong segregation distortion
present in T. monococcum. These resistant accessions are
resulted in the recovery of an abnormally high frequency
being characterized further by testing with additional
of recombinants. Eight of the 160 resistant recombinants
stem rust races. Crosses among selected resistant
had recovered wheat chromatin at each of the four
T. monococcum accessions have been initiated to
marker loci and apparently retained comparatively
determine the number and allelic relationships of stem
short terminal segments of foreign chromatin. The
rust resistance genes.
latter plants were used in a search that identified 12
anonymous AFLP loci that could be used for continued
mapping. The data obtained suggested reduced
homoeology between 1AL and the Lr59 translocation in
the distal chromosome regions, most likely due to the
presence of a paracentric inversion. Up to six or seven
of the eight shortest recombinants may have been
produced through crossing over within an inversion
loop and are thus genetically imbalanced. Development
and field evaluation of near-isogenic lines of five of the
eight recombinants will be necessary to identify those
that retained the shortest balanced translocations.
1
Cereal Research Center, Agriculture and Agri-Food Canada, Winnipeg, MB R3T
2M9, Canada; 2Department of Plant Science, University of Manitoba, Winnipeg, Instituto Nacional Autónomo de Investigaciones Agropecuarias (INIAP), Quito-
MB R3T 2N2, Canada Ecuador, Pan, Sur Km1, Ecuador
1
Department of Genetics, University of Stellenbosch, Private Bag X1, Matieland
7602, South Africa; 2Cereal Research Centre, Agriculture and Agri-food Canada, 1
USDA-ARS, Northern Crop Science Laboratory, Fargo, ND 58105 USA; 2 USDA-
195 Dafoe Road, Winnipeg, MT R3T 2M9, Canada ARS, Cereal Disease Laboratory, University of Minnesota, St. Paul, MN 55108, USA
1
Department of Molecular Biology & Genetic Engineering, 2Department of
Biotechnology, 3Department of Plant Pathology, Sardar Vallabh Bhai Patel
University of Agriculture & Technology, Modipuram, Meerut 250110, India;
4
Leibniz Institute of Plant Genetics and Crop Plant Research (IPK), Corrensstr.
3, 06466 Gatersleben, Germany; 5CIMMYT, Apdo. Postal 6-64106600 Mexico,
Department of Plant Breeding and Genetics, Cornell University, Ithaca, D.F., Mexico; 6CIMMYT, South Asia Regional Office, PO Box 5186, Singha Durbar
NY 14853, USA Plaza, Marg Bhadrakali, Kathmandu, Nepal
1
Department of Crop and Soil Science, 2Department of Botany & Plant
1
USDA-ARS Wheat Genetics, Quality, Physiology and Disease Research Unit, Pathology, Oregon State University, Corvallis, OR 97331, USA; 3USDA-ARS,
2
Department of Plant Pathology, 3Department of Crop and Soil Sciences, Deptartment of Plant Pathology, Washington State University, Pullman,
Washington State University, Pullman, WA 99164-6430, USA, 4College of Plant WA 99164-6430 USA; 4International Maize and Wheat Improvement Center
Protection, Northwest A&F University, Yangling, Shaanxi, China (CIMMYT), Apartado, Postal 6641, Mexico, 06600 DF, Mexico
1
Department of Agroenvironmental Science and Technology (DiSTA),
University of Bologna, Viale G. Fanin 44, 40127 Bologna, Italy; 2Società
Produttori Sementi (PSB), Via Macero 1, 40050 Argelato (BO), Italy; 3 Wheat
Program, International Maize and Wheat Improvement Center (CIMMYT) Apdo
Postal 6-641, 06600, Mexico DF, , Mexico; 4USDA/ARS/Cereal Disease Laboratory,
1
Agriculture and Agri-Food Canada, Cereal Research Center, 195 Dafoe 1551 Lindig Street, Univ. Minnesota, St. Paul, MN 55108, USA; 5Plant Breeding
Road, Winnipeg, MB, Canada R3T 2M9; 2Agriculture and Agri-Food Canada, and Genetics Department, Plant Breeding and Acclimatization Institute,
Semiarid Prairie Agricultural Research Center, P.O. Box 1030, Swift Current, 05-870 Blonie, IHAR Radzikow, Poland; 6Department of Plant Sciences, Tel Aviv
SK Canada S9H 3X2 University, 69978 Tel Aviv, Israel
There are many biotic constraints to wheat Sharon goatgrass (Aegilops sharonensis) is a wild
production in Morocco. While leaf rust and Septoria cereal endemic to the coastal plains of Israel. It is a
tritici leaf blotch were known from early times, yellow diploid species (2n=14) and possesses the Ssh genome,
(stripe) rust appeared in the area near the Atlas which is closely related to the B genome of wheat.
Mountains during the late 1980s. It recently spread Sharon goatgrass exhibits a high frequency and level of
to other cereal-growing areas, probably because of resistance to a number of wheat diseases including leaf
changes in virulence patterns (eg, Yr9 is no longer rust, stripe rust and stem rust. Many accessions of this
effective). Hence, a search for multiple disease species are also resistant to the widely virulent stem rust
resistances in wheat cultivars is a major objective. The race TTKSK (Ug99). Gene transfer from Sharon goatgrass
best lines from international nurseries were screened to is not straightforward due mainly to a lack of homology
widen the genetic base for wheat crop improvement. between the alien and wheat chromosomes, and also
Since diseases are not regularly expressed under field to the presence in the wild species of gametocidal
conditions, testing with local pathogen populations genes that prevent recovery of the pure wheat genetic
under controlled conditions was carried out for some background through backcrossing. We developed a
nurseries. The objective of this study was to identify method which combines the use of the ph1 gene to
wheat lines from international nurseries that carry promote pairing between homoeologous (partially
simultaneously adult-plant resistances to leaf rust, homologous) chromosomes and an anti-gametocidal
yellow rust, and Septoria tritici leaf blotch. Severities mutant gene to overcome the gametocidal effect.
and reaction types for leaf rust and yellow rust, and Production of wheat breeding material with a segment
pycnidial coverage for Septoria under field conditions, carrying the desired TTKSK resistance gene is under
and latent period and severity of Septoria under way. Selection of TTKSK resistant progenies during the
greenhouse conditions, were scored. A high frequency transfer process will be aided by molecular markers
of multiply resistant entries was observed among these linked to the gene.
accessions, reinforcing the importance of international
co-operation.
1
Institute for Cereal Crops Improvement, Tel Aviv University, Tel Aviv,
INRA – CRRA, PO Box 578, Meknès, Morocco; 2INRA – CRRA, PO Box 589,Settat,
1
Israel; 2Department of Plant Pathology, University of Minnesota, St. Paul,
Morocco; 3ICARDA, Rabat Institutes, PO Box 6299, Rabat, Morocco MN 55108,USA
1
Department of Plant Pathology, University of Minnesota, St. Paul, MN 55108,
USA; 2Department. of Plant Pathology, Kansas State University, and 3USDA-ARS
Plant Science & Entomology Research Unit, Manhattan, KS 66506, USA; 4USDA- 1
Department of Plant Pathology, University of Minnesota, and 2 USDA-ARS,
ARS Cereal Disease Laboratory, St. Paul, MN 55108, USA Cereal Disease Laboratory, St. Paul, MN 55108, USA
1
Agriculture and Agri-Food Canada, Lethbridge Research Centre, PO Box 3000,
5430-1st Avenue, South, Lethbridge, Alberta T1J 4B, Canada; 2College of Plant
1
USDA-ARS, Cereal Disease Laboratory, and 2Department of Plant Pathology, Protection and Shaanxi Key Laboratory of Molecular Biology for Agriculture,
University of Minnesota, St. Paul, MN 55108, USA Northwestern A&F University, Yangling, Shaanxi 712100, PR China.
W Liu1,2, A Laroche1, Z-S Kang2, DA Gaudet1 Leaf rust is the most common, and one of the most
important, wheat diseases of the world. Current leaf rust
Wheat stripe rust, caused by Puccinia striiformis f.
control in the U.S. consists of breeding resistant cultivars
sp. tritici, is a destructive disease of wheat worldwide
by using identified Lr genes in the host. Cultivars with
and the development of resistant cultivars is the most
such genes usually become susceptible to infection
economical control method. The Yr10 gene in Moro
due to the tremendous extant genetic diversity of the
wheat, that encodes a cytoplasmic NB-LRR protein
pathogen that allows it to overcome resistant cultivars in
containing nucleotide-binding sites (NBS) and leucine-
2-4 years. Development of alternate methods of control
rich repeats (LRR), imparts seedling resistance to stripe
is limited since little is known about Puccinia genomes
rust. Virus-induced gene silencing (VIGS) is a rapid and
and plant : pathogen interactions. Construction of a
powerful tool to analyze the function of plant genes. We
genome-wide physical map is important in order to
employed the barley stripe mosaic virus (BSMV)-VIGS
fully understand the molecular basis of the infection
system to study the function of different domains of the
mechanism of the pathogen and its interaction with the
Yr10 gene, in the resistance response in Moro wheat. A
host. In an effort to discover more about the genetic
series of DNA fragments based on different domains of
potential of leaf rust in terms of AVR and VIR gene
Yr10 were inserted into BSMV-VIGS vectors. Moro wheat
regulation, and to create future novel plant resistance
infection by P. striiformis following transfection with
breeding strategies, we have proposed a study of the
vectors was examined at the morphological, cytological
pathogen genome by constructing a BIBAC library
and molecular levels. Susceptible responses consisting
and a physical map of the pathogen. The BIBAC library
of pustule formation and symptoms of compatibility
is being constructed from the P. triticina type culture
were observed in Moro leaves transfected with some
PRTUS 3 which has AVR1 (avirulence gene corresponding
of the fragments. We evaluated the expression of
to Lr1) disrupted using T-DNA mutagenesis via particle
the Yr10 gene by probing different domains. The
bombardment. The characterization of AVR1 in the BIBAC
effects of changes in expression of Yr10 on function of
library will serve as a point of reference for cloning
plant responses at the leaf and cellular levels will be
heterologous AVR and VIR genes, and for defining their
presented.
regulation and modes of inheritance and recombination.
1
Agriculture and Agri-Food Canada, Lethbridge Research Centre, PO Box 3000,
5403 1st Avenue, South, Lethbridge, Alberta T1J 4B1 Canada; 2College of Plant
Protection and Shaanxi Key Laboratory of Molecular Biology for Agriculture, Department of Soil and Crop Science, Texas A&M University, College
Northwestern A&F University, Yangling, Shaanxi 712100, PR China Station, TX 77843, U.S.A.
1
Department of Genetics, University of Stellenbosch, Private Bag X1, Matieland
7602, South Africa; 2Department of Plant Sciences, University of the Free State,
PO Box 339, Bloemfontein 9300, South Africa. Cereal Research Centre, 195 Dafoe Road, Winnipeg, MN R3T 2M9, Canada
M Cakir1, F Drake-Brockman2, M Shankar2, H Golzar2, D Wheat (Tricticum aestivum L.) leaf rust (caused
Kollehn1, R McLean2, H Bariana3, R Wilson2, I Barclay2, C
by Puccinia triticina), is a devastating foliar disease
Moore2, H Kuchel4, M Jones1, R Loughman2
in the US Great Plains where short-lived, major gene
resistances are mainly utilized. A hotspot rust screening
Molecular markers make possible the deployment
nursery, established at Castroville, Texas, the forefront
of multiple rust genes in adapted elite lines. In this study
of the Puccinia pathway in the US, is a joint effort
we report a summary of the microsatellite tagging of a
between Texas A&M University (TAMU), Oklahoma State
number of leaf rust, stem rust and stripe rust resistance
University (OSU), and Kansas State University (KSU).
genes from a variety of sources. Segregating Leichardt/
It has grown into a 20 acre (8.3 ha) nursery and now
WAWHT2071 and Sunland/Arrino populations were
involves the participation of almost all wheat breeders
used for mapping Lr13 and Lr28 where Leichardt
from eight Universities and three USDA research centers
and Sunland were the respective sources of the
across the US.
resistance genes. Lines C77.19/3*77W:549-163658 and
The nursery was mainly established to screen wheat
Sr33/2*Shortim//4*3/Jacup resistance lines were used
for reaction to leaf rust, stem rust (caused by Puccinia
as sources of Sr32 and Sr33. F2 and F2:3 populations
graminis f. sp. tritici and stripe rust (Puccinia striiformis),
were used for microsatellite tagging of the genes. Very
as well as oats for reaction to crown rust (Puccinia
closely linked SSR markers were identified for Lr13,
coronata) and stem rust (Puccinia graminis f. sp. avenae).
Lr28, Sr32 and Sr33 on chromosomes 2BS, 4AL, 2BS and
Heavy wheat leaf rust and oat crown rust infections
1DS, respectively. Results from field-based studies of
are an annual event and reliable data are obtained on
various mapping populations for the characterization
advanced experimental lines as well as established
of adult plant rust (APR) resistances from a variety of
wheat and oat varieties. The nursery has also been
sources such as Wyalkatchem, Yitpi and Frame will also
utilized for selection of single plants from segregating
be discussed. Molecular markers for a range of other
bulks. A first look at promising resistant germplasm from
rust resistance genes (Lr9, Lr19/Sr25, Lr24/Sr24, Lr34/
CIMMYT was conducted in collaboration with OSU. This
Yr18, Lr46/Yr29, Lr47, Sr26 and Sr36) are currently being
nursery has provided warnings regarding the weakening
implemented for variety development and germplasm
resistance of key Great Plains wheat cultivars.
enhancement. The likely impact of these applications on
Clearly, the rust screening nursery at Castroville has
wheat improvement will be discussed.
provided a rust screening hotspot for US breeders and
has proven indispensable since its inception in 2000.
1
WA State Agricultural Biotechnology Centre, Murdoch University, Murdoch,
WA 6150, Australia; 2Department of Agriculture and Food, 3 Baron-Hay Court, 1
Texas A&M University, 2474 TAMU, College Station, TX 77843, U.S.A;
South Perth, WA 6151, Australia; 3University of Sydney Plant Breeding Institute 2
Department of Plant and Soil Sciences, 368 Ag Hall, Oklahoma State University,
Cobbitty, PMB 11, NSW 2750, Australia; 4Australian Grain Technologies, Stillwater, OK 74078-6026, USA; 3Department of Agronomy, Kansas State
Roseworthy Campus, Roseworthy, SA 5371, Australia University, Manhatten, KS 66506, USA
Wheat in Hungary is threatened by all three rusts, All three rust diseases of wheat occur in the Czech
viz. leaf rust, stem rust and stripe rust. All three diseases Republic. Leaf rust is most frequent. The last stem rust
are capable of causing substantial economic losses, but epidemics occurred in 1972, and the last significant
their incidence varies due to their diverse ecological yellow (stripe) rust outbreak was in 1999-2001.
requirements. The greatest damage is currently caused Resistance breeding aims at combined resistance to
by leaf rust, which infects wheat fields every year. all three rusts. Combined resistance was present in 11
The most environmentally sound, low cost method of 29 new breeding lines recently tested. The highest
of controlling leaf rust is to breed and grow resistant resistance occurred in breeding line SG-S-469-07,
varieties. Both traditional and molecular breeding followed by BR-05-082 and SG-S-316-06. On average
methods are used to improve the leaf rust resistance of the highest degrees of resistance were to yellow rust.
wheat varieties bred in Martonvásár, Hungary. Of the winter wheat cultivars registered in the Czech
The field reponses of wheat genotypes carrying Republic, the highest combined resistance to all three
designated Lr genes have been assessed for many years rusts was in cultivars possessing the translocation from
in order to determine the effectiveness of major leaf rust Aegilops ventricosa (Yr17, Lr37, Sr38). In addition to rusts,
resistance genes. The ‘Thatcher’-based near-isogenic attention is also given to fusarium head blight, powdery
lines, carrying single genes for resistance are sown each mildew, tan spot, Septoria leaf blotch, Septoria glume
year. Eight NILs remain immune or highly resistant: these blotch and BYDV (barley yellow dwarf virus). Ring tests
include lines with Lr9, Lr19, Lr24, Lr25, Lr28, Lr29, Lr35 are organized at several different locations to screen
and Lr37. The levels of infection on four further lines new breeding lines for resistance to the various diseases.
(Lr23, Lr32, Lr3ka and Lr22a) were also quite low. The line Spreaders are inoculated when natural infection is not
exhibiting the greatest degree of infection was the NIL adequate.
carrying Lr26.
The segregating populations in the breeding
program are tested and selected continuously under
artificially inoculated conditions. A special nursery is 51. Screening Wheat Germplasm for
devoted to testing the leaf rust resistance of advanced
breeding lines, special genetic stocks and potential leaf
Resistance to Stem Rust in Georgia
rust resistance sources. The levels of resistance in the Z Sikharulidze1, D Bedoshvili2, L Mgeladze1,
released and cultivated winter wheat varieties bred K Natsarishvili1, N Chkhutiashvili3
in Martonvásár, namely, ‘Mv Magvas’, ‘Mv Marsall’, ‘Mv
Toborzó’, ‘Mv Béres’, ‘Mv Matyó’, ‘Mv Vekni’, ‘Mv Laura’ and Stem rust was a major threat to wheat production
‘Mv Lucia’ at 0–20MR, are sufficient to negate the need in Georgia before the 1970s. However, promotion
for chemical control in farmers’ fields.
of stem rust resistant varieties, such as Bezostaia 1,
Using marker-assisted selection (MAS), the
reduced its impact on production. Recently, there has
resistance genes Lr9, Lr24, Lr25, Lr29, Lr35 and Lr37
been an increase in stem rust occurrence in some areas
were incorporated into four Martonvásár winter wheat
of Georgia. There is also a possibility that race Ug99
varieties. A marker- assisted backcross program to track
will eventually reach Georgia. Barberry is widespread
the transfer of effective Lr genes has begun. Wheat
in the country. Identification and promotion of rust
varieties susceptible or moderately resistant to leaf rust
resistant germplasm is an important strategy for wheat
were crossed with NILs of ‘Thatcher’ each carrying a
rust control, especially because the use of fungicides
different Lr gene (Lr9, Lr24, Lr25, Lr29 or Lr35) and with
on wheat is not a common practice in Georgia. The
the variety ‘Renan’ (Lr37). Plants in the fifth backcross
generation had agronomic traits resembling the
Research Institute of Cro Production, Drnovská 507, 161 06 Praha 6 - Ruzyne,
recurrent parent. Czech Republic
Agricultural Research Institute of the Hungarian Academy of Sciences, 1
Georgian Institute of Plant Immunity, Kobuleti, Georgia; 2ICARDA-CIMMYT;
Brunszvik u. 2. H-2462 Martonvásár, Hungary 3
I. Lomouri Farming Institute, Mtskheta, Georgia
Wheat is the main staple food crop in Tajikistan Kazakhstan is one of the largest wheat producers in
and it is of increasing importance to develop high central Asia. Wheat rusts are important problems in our
yielding varieties with disease resistance and good country. Stem rust (pathogen, Puccinia graminis f. sp.
bread making quality. International collaboration has tritici) causes considerable damage, especially in wetter
been established, and nurseries are received especially years. In order to combat the menace of rust, screening
from the Turkey-CIMMYT-ICARDA International Winter of various nurseries from national and international
Wheat Improvement Program (IWWIP), located in Turkey, breeding programs was initiated. The aim of the present
but also more recently from Oklahoma State University work was to find sources of stem rust resistance and
in the USA. to develop disease-free germplasm. The material was
Together with tan spot, yellow (stripe) rust is a screened with the predominant races in the region.
major biotic constraint faced by wheat farmers in Cultivar Steklovidnaya 24 was used as a susceptible
Tajikistan. Through a multilocation testing system, check. A total of 55 wheat genotypes were included in
several high yielding and resistant lines were identified the 2008 tests; 33 lines showed high or moderate levels
and are in the process of being released. of resistance in the field. Tests of the material under
In order to test for resistance to race Ug99 in Tajik artificial conditions identified eight entries with stem
wheat germplasm a number of varieties and advanced rust immunity ; viz. 86003/F9Norin10/Steklovidnaya24,
lines were tested in Kenya in collaboration with CIMMYT. 86004 /F7322-MA/118-SI, 86006/F6KSI-21/Arthur, 86007/
The results demonstrated a low level of resistance in F6KSI-21/Arthur, 86018/Lawson/Currawong, 86019/
Tajik germplasm, indicating an urgent need to initiate Moro*2(C90) /More*2//Marcuis)2, 86022/F5Janbash/
breeding activities to reduce the consequences of a Anza, 86023/F4KLDN33/MK-3832, 86024/F4Tilek/KLDN-
possible incursion of Ug99 to Tajikistan. A collaborative 95. Two lines were characterized as moderately resistant;
project has been initiated with the Swedish Agricultural viz. 86005/F6 Progress/T. monococcum and 86008/
University to introgress novel genes for resistance to F6KSI-21/97Sr25. Evaluation for agronomic traits allowed
race Ug99 into Tajik wheat germplasm. selection of 10 advanced lines with high yield potential
This paper evaluates the results of multilocation and resistance to stem rust. Because Ug99 is virulent
trials conducted during 2005, 2006 and 2007 through to the great majority of wheat varieties, we sent our
which high yielding and disease resistant lines were promising material to Kenya for testing. Based on the
identified and recommended for submission to official results we will be able to develop cultivars possessing
variety testing trials. Furthermore, the paper discusses genes, or combinations of genes, efective against this
the future breeding strategy to increase the level of widely virulent race of the pathogen.
resistance to race Ug99 in Tajik wheat germplasm.
1
Tajik Agrarian University, Dushanbe, Tajikistan; 2Research Institute of Farming,
Dushanbe, Tajikistan; 3Sida Project “Support to Seed Industry Development in
the Republic of Tajikistan”, PO Box 195, Dushanbe, 734025 Tajikistan; 4Chilgazi 1
Laboratory of Plant Breeding and Genetics, Institute of Plant Biology and
Farm, Isfara Rayon, Tajikistan; 5Vakhsh Branch of the Farming Institute, Bokhtar, Biotechnology, Almaty, Kazakhstan; 2Laboratory of Plant Immunity, Institute of
Tajikistan; 6International Maize and Wheat Improvement Center, Turkey Problems of Biological Safety, Gvardeysky, Kazakhstan
Resistance to the wheat rusts is improved in level The fight against rusts relied heavily on major
and durability when resistance genes are stacked. genes, but other genes also exist. Our own experience
Selecting gene stacks in breeding populations by has been mostly with BYDV and FHB, both diseases
phenotype can be difficult or impossible and marker- having very complex genetics. Twenty five years of
assisted selection is expensive. Furthermore, when attempts to breed resistance based on major genes gave
stacks are selected the effective size of the population poor results. Then we undertook to seek simultaneous
is reduced thus limiting the available variability from resistances to all diseases present in Eastern Canada.
which to select other characters. We propose using We thus breed against rusts, powdery mildew, BYDV
telocentric chromosomes to fix resistance gene and FHB. Using much more biodiversity and selecting
stacks in breeding populations by selecting double intensively for resistance to all diseases should single
monotelodisomic F1 plants (2n = 40 + t + t) with the pair out plants that resist nearly all diseases. Doing this,
of resistance genes in the hemizygous condition. This more than 99% of the germplasm was destroyed by
method was demonstrated in two wheat populations, diseases. Among 10,000 F1 plants inoculated, one single
each with a different two-gene stack of leaf rust cross combination gave the sought-after result. Within
resistance genes. The presence of critical telocentric one year, we had introgressed in one genotype the FHB
chromosomes in the populations rapidly drove stack resistance of Sumai 3, very good BYDV and powdery
frequencies toward fixation by a combination of mildew resistances, and rust resistance equal to that
selection for euhaploid pollen and zygotic selection for of the most resistant parent. Important lessons follow.
diploid and near-diploid (i.e. no ditelosomics) plants. The value of a gene source cannot be fully judged by
Thus, telocentric chromosomes provide a tool to fix gene its disease reaction because epistatic hidden genes
stacks in a population while maintaining the effective can exist in any line. Making many crosses is the way to
size of the population for selection on other criteria. One get the most out of the hidden genetics. A very severe,
point of consideration is the relatively large size of the complex selection protocol can work. The method
linkage blocks being fixed. gave resistance to all Eastern rust races. BYDV tolerance
correlated with yield and biomass potential. Applying
a multiple-stress system to more rust species is worth
a try. Good outcomes are expected in pyramiding
slow rusting genes, and multiple genes form durable
horizontal resistance. Multiple approaches constitute
the best strategy to address a disease that can ruin part
of the world food basket.
1
CRDSGC, Agriculture and Agri-Food Canada, Québec City, QC Canada;
2
EMBRAPA Clima Temperado, Pelotas, Brazil; 3CRC, Agriculture and Agri-
Food Canada, Winnipeg, MB, Canada; 4ECORC, Agriculture and Agri-Food
Canada, Ottawa, ON Canada; 5CÉROM, Saint-Mathieu-de-Beloeil, QC Canada;
6
CÉROM, Québec City, QC Canada; 7CLRC, Agriculture and Agri-Food Canada,
Charlottetown, PEI Canada; 8LRC, Agriculture and Agri-Food Canada,
Cereal Research Centre, 195 Dafoe Road, Winnipeg, MN R3T 2M9, Canada Lethbridge, Alberta, Canada
1
INRAT-Tunis, 2CIMMYT-Mexico, 3ICARDA-Aleppo
*Corresponding author: gharbi.medsalah@iresa.agrinet.tn
1
Department of Genetics and Plant Breeding, and 2Department of Mycology
and Plant Pathology, Institute of Agricultural Sciences, Banaras Hindu
University, Varanasi 221005, India; 3CIMMYT South Asia Office, Kathmandu,
Nepal; 4Centro Internacional de Mejoramiento de Maíz y Trigo (CIMMYT),
Apdo. Postal 6-641, C.P. 06600, D.F. Mexico
This document was prepared by the BGRI Secretariat, with financial support from the
Durable Rust Resistance in Wheat Project (http://wheatrust.cornell.edu).
The abstracts were edited by Dr. Robert McIntosh, Honorary Associate at the
Plant Breeding Institute of the University of Sydney.
www.globalrust.org www.globalrust.org