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PATHOTYPE VARIATION OF PUCCINIA STRIIFORMIS POPULATION
TRIGGERING STRIPE RUST OF WHEAT IN THE NORTHERN PUN JAB
AND NWFP
SYED NADEEM AFZAL 04-arid-10
Department of Plant Pathology Faculty of Crop and Food Sciences
Pir Mehr Ali Shah Arid Agriculture University Rawalpindi
Pakistan 2009
PATHOTYPE VARIATION OF PUCCINIA STRIIFORMIS POPULATION
TRIGGERING STRIPE RUST OF WHEAT IN THE NORTHERN PUN JAB
AND NWFP
By
SYED NADEEM AFZAL 04-arid-10
A thesis submitted in partial fulfillment of the requirement for the degree of
DOCTOR OF PHILOSOPHY
IN
PLANT PATHOLOGY
Department of Plant Pathology Faculty of Crop and Food Sciences
Pir Mehr Ali Shah Arid Agriculture University Rawalpindi
Pakistan 2009
CERTIFICATION
I hereby undertake that this research is an original one and no part of this thesis falls
under plagiarism. If found otherwise, at any stage, I will be responsible for the
consequences.
Student's Name: Syed Nadeem Afzal Signature: ___________________
Registration No. 04-arid-10 Date: _______________________
Certified that the contents and form of thesis entitled “Pathotype variation of
Puccinia striiformis population triggering stripe rust of wheat in the northern
Punjab and NWFP” submitted by “Syed Nadeem Afzal” have been found
satisfactory for the requirement of the degree.
Supervisor: _________________________ (Prof. Dr. Irfan Ul-Haque)
Co-Supervisor: _________________________ (Dr. Iftikhar Ahmad)
Member: _________________________ (Prof. Dr. Ch. Abdul Rauf)
Member: _________________________ (Prof. Dr. Muhammad Munir)
Chairman: ________________________________
Dean: _____________________________________
Director Advanced Studies: ____________________
LIST OF ELLIPSIS
AUP Agricultural University Peshawar
CCRI Cereal Crop Research Institute
CDRP Crop Disease Research Program
IT Infection Type
NARC National Agricultural Research Center
NIFA Nuclear Institute for Food and Agriculture
NILs Near Isogenic Line/s
NWFP North Western Frontier Province
PMAS-AAUR Pir Mehr Ali Shah – Arid Agriculture University Rawalpindi
PRS Pulses Research Station
Pst. Puccinia striiformis f.sp. tritici
R. Value Response Value
T. spelta album Triticum spelta var. album
WSRTN Wheat Stripe Rust Trap Nurseries
WDSN Wheat Disease Screening Nursery
DEDICATED
TO
THE LOVING PARENTS OF MINE
AS THEY ALWAYS SHELTERED ME
IN RAIN OR IN SHINE
TO
SHAYAN ALI, THE NAUGHTY & PLAYFUL SON
WHO SUFFERED MOST IN HIS EDUCATION
DUE TO LACK OF MY ATTENTION
DUIRNG WRITING OF DOCTORAL DISSERTATION
TO
ZAINAB JAFFERY, MY YOUNGEST LOVELY DAUGHTER
WHOSE DAZZLING EYES AND GORGEOUS FACE
ALWAYS DEMANDED TO PICK UP THE PACE
TO
HUSSAIN NADEEM, MY SERIOUS SON
WHO IS A THOROUGH GENTLEMAN
TO
TASHAN FATIMA, THE ELDEST DAUGHTER
INTELLIGENT, HARDWORKING AND NICE WRITER
TO
MY WIFE AND LIFE PARTNER
ALWAYS ENCOURAGED ME WITH HER LOVELY LAUGHTER
CONTENTS
TITLE PAGE LIST OF TABLES iv LIST OF FIGURES vii LIST OF APPENDICES ix ACKNOWLEDGEMENTS x GENERAL ABSTRACT xii
1. GOLBAL WHEAT PRODUCTION AND RUST SCENARIO 1
1.1 GENERAL INTRODUCTION 1
1.1.1 The Wheat World 1 1.1.2 Wheat Rusts 3 1.1.3 Nature of the Rust Fungi 4 1.1.4 Occurrence of Rusts in Pakistan 5 1.1.5 Objectives of the Research 6 2 PATHOTYPE VARIATION OF PUCCINIA STRIIFORMIS 8
2.1 INTRODUCTION 8 2.2 REVIEW OF LITERATURE 12 2.3 MATERIALS AND METHODS 24
2.3.1 Modus Operandi for Field Sampling of Stripe Rust 25 2.3.2 Collection of Diseased Samples from Farmers' Fields 25 2.3.3 Establishment of WSRTNs 26 2.3.3.1 Stripe Rust Monitoring 27 2.3.3.2 Recording of Observations at WSRTN 30 2.3.4 Establishment of WDSN 31 2.3.5 Glasshouse Virulence Analysis 32 2.3.5.1 Inoculum Increase 32 2.3.5.2 Isolated Development 34 2.3.5.3 Stripe Rust Assessment on NILs 35 2.3.6 Virulence Assessment 36 2.3.7 Designation of Pathotype 36 2.3.8 Population Structure 37 2.3.8.1 Virulence Frequency 37 2.3.8.2 Virulence Complexity 38 2.3.8.3 Pathotype Abundance 38 2.3.8.4 Cluster Analysis 38
i
2.4 RESULTS 39
2.4.1 Survey for Disease Sample Collection 39 2.4.2 Race Analysis 39 2.4.2.1 Prevailing Races of Pst 42 2.4.3 Population Structure of Pst in the Northern Punjab
and NWFP 52
2.4.3.1 Diversity of Virulence 52 2.4.3.2 Virulence Frequency 53 2.4.3.3 Virulence Complexity 58 2.4.3.4 Pathogen Abundance 61 2.4.3.5 Cluster Analysis 63 2.4.4 Wheat Stripe Rust Trap Nurseries 65 2.4.5 Field Response of Commercial Wheat Varieties to Pst 68 2.4.6 Field Response of Breeders advance Lines to Pst 72
2.5 DISCUSSION 76 2.5.1 Population Structure of Pst 76 2.5.2 Wheat Varietals Response to Stripe Rust 83 2.5.3 Epidemiological Zone of the Indian Subcontinent 86 2.5.3.1 Migration and Introduction of Stripe Rust Pathogen 86 2.5.3.2 Influence of Environmental Factors on Stripe Rust 90 2.5.4 Pst Distribution in the Northern Punjab and NWFP 92 2.5.4.1 Race Studies in the Northern Punjab and NWFP 95 2.5.4.2 Stripe Rust Occurrence during 2005-06 and 2006-07 99
2.6 CONCLUSIONS 106
3 ESTIMATION OF WHEAT YIELD LOSSES DUE TO STRIPE RUST
111
3.1 INTRODUCTION 111 3.2 REVIEW OF LITERATURE 114 3.3 MATERIALS AND METHODS 121
3.3.1 Soil Analysis 122 3.3.2 Experimental Layout 122 3.3.3 Trials Conducted under the Research Study 123 3.3.4 Observation of Rust Severity 125 3.3.5 Yield Parameters 125 3.3.5.1 Thousand Grain Weight 126 3.3.5.2 Grain Yield (Kg ha-1) 126 3.3.6 Statistical Analysis 126
ii
3.4 RESULTS AND DISCUSSION 127
3.4.1 Effect of Environmental Conditions in Expressing Stripe Rust
127
3.4.1.1 Development of Wheat Stripe Rust in Experimental area during 2005-06
127
3.4.1.2 Expression of wheat Stripe Rust in experimental area during 2006-07
132
3.4.2 Disease Severity Recorded in Wheat Varieties during 2005-06 and 2006-07
135
3.4.3 Performance of Wheat Varieties against Disease severities on 1,000 Grain weight during 2005-06 and 2006-07
138
3.4.4 Expression of Wheat Varieties for 1,000 Grain Weight in Control during 2005-06 and 2006-07
141
3.4.5 Response of Wheat Varieties against Disease Severities in Yield (Kg ha-1) during 2005-06 and 2006-07
145
3.4.6 Yield Response of Wheat Varieties in Control Plots during 2005-06 and 2006-07
148
3.4.7 Correlation between Disease Severity and Yield 154 3.4.7.1 Correlation between Disease Level and Yield during
2005-06 154
3.4.7.2 Correlation between Disease Level and Yield during 2006-07
155
3.4.8 Regression Analysis 156 3.4.8.1 Regression Analysis Studies during 2005-06 156 3.4.8.2 Regression Analysis Studies during 2006-07 158 3.4.9 Changes in Varietal Susceptibility 164 3.4.10 Disease Severity and Yield Loss of Wheat Cultivars
during 2005-06 and 2006-07
164 3.5 CONCLUSIONS 168
LITERATURE CITED 170
iii
LIST OF TABLES
TABLE NO. PAGE
1. Location of Trap Nurseries Established under the Research area 26
2. List of Commercial Varieties included in WSRTN (2005-06 and 2006-07)
28
3. Field Response of Host Plant against Stripe Rust 31
4. Location of Stripe Rust Samples collected (Natural Infection) during 2005-06 and 2006-07
33
5. Differential set for Race Determination and Virulence Analysis of Stripe Rust Pathogen at CDRI, Murree
34
6. Stripe Rust Virulence Frequency in the Northern Punjab and NWFP during 2005-06 and 2006-07
41
7. Prevalence of Physiological Races in the Northern Punjab and NWFP during 2005-06 and 2006-07
43
8. Physiological Races Identified in the Northern Punjab and NWFP during 2005-06 and 2006-07
43
9. Response of Identified Pst Pathotypes* among Differential Genotypes
45
10. Reaction of Thirteen Pathotypes from 150 Isolates on the Corresponding Stripe Rust Resistant Genes Possessed by 29 Wheat Differentials
53
11. Virulence Frequency (%) in NILs and Isolates Patterns of Pst in the Northern Punjab and NWFP
55
12. Virulence Frequency (%) isolates of Pst collected from the Northern Punjab and NWFP during 2005-06 and 2006-07
57
13. Grouping of Pst Pathotypes collected from the Northern Punjab and NWFP during 2005-06 based on the Frequency of Occurrence and Virulence Gene
59
14. Grouping of Pst Pathotypes collected from the Northern Punjab and NWFP during 2006-07 based on the Frequency of Occurrence and Virulence Gene
60
iv
15. Number of Prevailing Pathotypes found in the Northern Punjab and NWFP during 2005-06
62
16. Number of Prevailing Pathotypes found in the Northern Punjab and NWFP during 2006-07
62
17. Percent disagreement among the Pst Pathotypes identified from the Samples Collected during 2005-06 and 2006-07
64
18. Virulence Analysis through Stripe Rust Reactions in NILs with Avocet Background at WSRTN sites of the Northern Punjab and NWFP during 2005-06 and 2006-07
66
19. Stripe Rust Reactions of Commercial Varieties to Pst at Hot spots of the Northern Punjab and NWFP during 2005-06 and 2006-07
70
20. Stripe Rust Reactions at Seedling Stage on Commercial Varieties at CDRP, Sunny Bank, Murree during 2005-06 & 2006-07
72
21. Environmental Conditions Persisted during 2005-06 and 2006-07 in the Northern Punjab and NWFP
93
22. Physical and chemical properties of the experimental site 122
23. ANOVA Expressing Response of wheat Varieties to Incidence of Stripe Rust during 2005-06
135
24. ANOVA Expressing Response of wheat Varieties to Incidence of Stripe Rust during 2006-07
135
25. Comparison of Mean Values for Disease Severity 136
26. ANOVA Exhibiting Effect of Disease Severities on 1,000 Grain Weight of Different Wheat Varieties (2005-06)
139
27. ANOVA Exhibiting Effect of Disease Severities on 1,000 Grain Weight of Different Wheat Varieties (2006-07)
139
28. Comparison of Mean Values for 1,000 Grain Weight in Diseased Plots
140
29. ANOVA Showing Varying Genetic Potential of Wheat Varieties in 1,000 Grain weight in Control during 2005-06
141
30. ANOVA Showing Varying Genetic Potential of Wheat Varieties in 1,000 Grain Weight in Control during 2006-07
142
31. Comparison of Mean Values for 1,000 Grain Weight in Control 142
v
32. ANOVA Showing impact of Stripe Rust on 1,000 Grain weight of Different Wheat Varieties during 2005-06 and 2006-07
144
33. ANOVA Showing Response of Wheat Varieties against Disease Severities in Yield (Kg ha-1) during 2005-06
145
34. ANOVA Showing Response of Wheat Varieties against Disease Severities in Yield (Kg ha-1) during 2006-07
146
35. Comparison of Mean Values for Kg ha-1 in Diseased Plots 146
36. ANOVA Showing Yield Response of Wheat Varieties in Control Plots during 2005-06
148
37. ANOVA Showing Yield Response of Wheat Varieties in Control Plots during 2006-07
149
38. Comparison of Mean Values for Kg ha-1 Yield in Control Plots 149
39. ANOVA Showing Impact of Stripe Rust on Wheat Yield (Kg ha-1) of different Wheat Varieties during 2005-06 and 2006-07
153
40. Regression Analysis showing Dependence of Wheat Yield on Disease Level (2005-06)
156
41. Regression Analysis showing Dependence of Wheat Yield on Disease Level (2006-07)
158
42. Comparison of Mean Values for Stripe Rust Severity and 1,000 Grain Weight during 2005-06 and 2006-07
159
43. Yield Reduction in different Wheat Varieties due to Stripe Rust during 2005-06 and 2006-07
160
44. Comparison of 1,000 Grain Weight in Wheat Varieties due to Stripe Rust during 2005-06 and 2006-07
165
45. Comparison of Yield difference in Wheat Varieties due to Stripe Rust during 2005-06 and 2006-07
165
vi
LIST OF FIGURES
FIGURE NO. PAGE
1. Field Design for Stripe Rust Disease Assessment and Sampling 25
2. The Modified Cobb's Scale 30
3. Adult Plant Host Response of the Stripe Rust 31
4. Scale 0-9 for Stripe Rust Disease Scoring at Seedling Stage 36
5. District of the Northern Punjab and NWFP Prone to Stripe Rust 40
6. Virulence Frequency of Pst in the Northern Punjab and NWFP during 2005-06
56
7. Virulence Frequency of Pst in the Northern Punjab and NWFP during 2006-07
56
8. Complexity (Virulence loci number in each isolate) distribution for the samples pertaining to the Northern Punjab and NFWP during 2005-06
59
9. Complexity (Virulence loci number in each isolate) distribution for the samples pertaining to the Northern Punjab and NFWP during 2006-07
60
10. Tree Diagram for 13 Pathotypes Collected from the Northern Punjab and NWFP during 2005-06 and 2006-07
64
11. Cluster analysis of 188 commercial and breeders lines based on disease resistance traits
73
12. Field Photographs 100
13. Global Location of Experimental site in the PMAS-Arid Agriculture University, Rawalpindi
121
14. Maximum Disease Expression by Commercial Varieties during 2006 (a) and 2007 (b)
128
vii
15. Distribution of Rainfall in Rawalpindi during 2006 and 2007 130
16. Prevalence of Humid Conditions during 2006 and 2007 130
17. Minimum and Maximum Temperature Regime at Rawalpindi during 2006
131
18. Minimum and Maximum Temperature Regime at Rawalpindi during 2007
131
19. Weekly Average Minimum and Maximum Temperatures at Rawalpindi during 2006
152
20. Weekly Average Minimum and Maximum Temperatures at Rawalpindi during 2007
152
21. Graph Showing Relationship between Disease Severity and Wheat Yield 2006
154
22. Graph Showing Relationship between Disease Severity and Wheat Yield 2007
155
23. Comparison of 1,000 Grain Weight of Wheat Varieties in Control and Diseased Plots during 2005-06
157
24. Comparison of 1,000 Grain Weight of Wheat Varieties in Control and Diseased Plots during 2006-07
157
25. Estimated Yield Losses in Response to Different Levels of Stripe Rust 2006
161
26. Estimated Yield Losses in Response to Different Levels of Stripe Rust 2007
161
27. Shriveling of Wheat Grains due to Stripe Rust Intensities: Wheat Grains in a. Control; b. up to 20%; c. more than 20 but less than 40%; d. more than 40%
167
viii
LIST OF APPENDICES
APPENDIX # APPENDIX TITLE PAGE
I Field response of differential set and NILs to Pst at hot spots in the Northern Punjab and NWFP during 2005-06 & 2006-07
202
II Pathotype determination of Pst through seedling infection types produced on world and European differentials during 2005-06 & 2006-07 (Sheet 1 to 8)
203
III Infection types of single pustule isolates of Pst on stripe rust differentials based on seedling reaction during 2005-06 (Sheet 1 to 9)
211
IV Infection types of single pustule isolates of Pst on stripe rust differentials (NILs) based on seedling reactions during 2006-07 (Sheet 1 to 10)
220
V Stripe rust reactions of commercial varieties to Pst at different locations of Northern Punjab & NWFP during 2005-06 & 2006-07
230
VI Screening results of wheat germplasm based on Coefficient of Infection, Relative Resistance Index recorded during the year 2005-06 and 2006-07
231
VII Wheat Stripe Rust Development Influenced by Environmental Factors Prevailed at Rawalpindi during 2006 (A) and 2007 (B)
234
VIII Precipitation received at Rawalpindi during 2006 and 2007 235
IX Temperature range in Rawalpindi during 2006 (A) & 2007 (B) 236
X Humidity percentage in Rawalpindi during experimentation years 238
XI Disease Severity Percentage using Cobb’s scale during 2005-06 239
XII Disease Severity Percentage using Cobb’s scale during 2006-07 239
XIII 1,000 grain weight of wheat in control plots during 2005-06 239
XIV 1,000 grain weight of wheat in diseased plots during 2005-06 239
XV 1,000 grain weight of wheat in control plots during 2006-07 239
XVI 1,000 grain weight of wheat in diseased plots during 2006-07 240
XVII Wheat yield (kg per hectare) in control plots during 2005-06 240
XVIII Wheat yield (kg per hectare) in diseased plots during 2005-06 240
XIX Wheat yield (kg per hectare) in control plots during 2006-07 240
XX Wheat yield (kg per hectare) in diseased plots during 2006-07 240
ix
ACKNOWLEDGEMENTS
All praises are for his supreme majesty ALLAH , who holds the life of all lives and is
the lord of all lords. I am immensely indebted to Almighty ALLAH , the auspicious,
the compassionate and superior whose exalt and magnificence, thrived my thoughts
and flourished my aspirations, blessed me splendid professors, empathetic parents,
exquisite family and inimitable friends. Cordial esteem for HOLY PROPHET
MUHAMMAD (S.A.W.) who enlightened our scruples with the spirit of fidelity in
ALLAH and for IMAM ALI (A.S.) who is an icon of valor and gallantry for us.
The accomplished research work was documented under the kindhearted approach,
paternal behavior, vigilant tracking, erudite analysis and liberal supervision of Prof.
Dr. Infan Ul-Haque, Chairman, Department of Plant Pathology, PMAS-AAUR. His
meticulous scrutiny and precise appraisal not only enhanced the worth of this
dissertation but also my understanding of the Phyto-pathology. I am in great debt of
thanks to his ever instigating assistance, zealous concern, academic interpretation
and practical ideas through out my studies.
I reckon it my extreme gratification in extending gratitude to my Co-Supervisor, Dr.
Iftikhar Ahmad, Director General, NARC, Islamabad, my committee members
Prof. Dr. Muhammad Munir, Dean, Faculty of Crop and Food Sciences and Prof.
Dr. Abdul Rauf, Department of Plant Pathology for imparting me with technical
solutions as and when requested. Their professional veracity and precious
propositions will always be with me right through the course of life. I extend deep
emotions of appreciation, gratitude and indebtedness for their valuable guidance.
Cordial and virtuous appreciation to all my friends especially Dr. Shoaib
Ahmadani, Syeda Siddiqua Firdous, Dr. A. R. Rattu and Mr. M. Fayyaz who
stood beside me and shared my work load with open heart. I express my sincere
x
gratitude for all my friends and colleagues at the PMAS-AAUR; NARC, Islamabad;
CDRP, Sunny Bank, Murree; PRS, Sialkot; NIFA, Peshawar; AUP, Peshawar and
CCRI, Pirsabak.
PMAS-AAUR has always been my favourite institution and the name was not new
for me as I am among those who graduated from this institution when it was known
as Barani Agricultural College. I must pay homage and tribute to this institution that
has broadened my vision and brought me where I stand today with honor and
dignity.
I have no words to show gratitude to my caring and amorous parents and gorgeous
family whose palms always stretched with wet eyes in prayers for me and especially
my father Syed Afzal Hussain Jaffery, who at age of 70, pretend before me as a
young man so that I may not worry about my family during accomplishment of my
doctoral studies. I must admit that without the moral support of my father, uncle
Syed Shaukat Kazmi, Syed Jafar Raza and my wife, the present destination would
have hardly been an illusion.
SYED NADEEM AFZAL
xi
GENERAL ABSTRACT
Wheat is one of the leading grain crops of Pakistan and being staple diet of the
inhabitants, it grasps a key position in the agricultural policies. Sustainable productivity
of wheat is of paramount importance in the context of many biotic and abiotic factors
that limit its production. Stripe or yellow rust is one of such biotic factors, caused by an
obligate parasite Puccinia striiformis Westend. f.sp. tritici Eriks., that confines wheat
production throughout the world. Presence of several races of each and ever-changing
nature of the stripe rust pathogen cautions cultivation of susceptible wheat cultivars in
humid, high uplands and cooler regions of the country.
Monitoring of the stripe rust population is imperative to determine pathotype variation
so that new virulences with the potential to overcome resistance genes currently
deployed in the wheat cultivars can be detected. Accordingly, the research was executed
in the Northern Punjab and NWFP to identify prevailing Pst virulence pattern and
pathotype variation and trap the stripe rust pathogen through establishing “Trap
Nurseries” at selected sites of the study area; estimate wheat yield losses due to stripe
rust in field under disease stress conditions; and evaluate breeders material for its
disease reaction under field condition.
Analyses of yield loss data confirmed significant loss where disease level was very high.
The most severely diseased cultivars had the lowest yields. Morocco depicted extremely
low yields whereas Inquilab-91 expressed a loss of only 38.73 kg ha-1 against the disease
severity of 36.25 percent in 2007 as compared to the loss of 143.3 kg ha-1 when the
xii
disease severity was just 8.12 percent during 2006. Stripe rust was also severe against
Bakhtawar and Wafaq-2001 during 2007 at the test sites in Rawalpindi and showed a
dramatic affect on yield. Considerable yield losses were observed in 2007 as compared
to the losses estimated during 2006 attributing to the conducive environmental factor
that persisted for more than four weeks and prolonged the infectious period of the stripe
rust pathogen.
Screening of 188 varieties / advanced breeding lines against stripe rust was also carried
out during 2005-06 and 2006-07. Cluster analysis based on the RRI was performed,
which indicated that the entire cultivars could be distributed into six clusters at 20
percent linkage level. Average Coefficient of Infection (ACI) and Relative Resistance
Index (RRI) values of two year trial showed that out of 188 cultivars 150 had RRI value
>7 ≤9 and were found in the desirable range; 28 cultivars were included among the
acceptable range having RRI value ≥5 <7. However, only 10 cultivars showed RRI
value <5 and fell under undesirable range.
In the present study, pathotype variation of Pst population that occur naturally in the
major wheat growing areas of Pakistan were analyzed. The outcomes were highlighted
in the context of prevailing virulences and identification of the Yr resistance genes that
are still effective. During 2005-06 and 2006-07, 12 previously identified Pst races were
confirmed while one new race was identified. In Pakistan, virulence is present for the
stripe rust resistance genes Yr1, Y6, Yr7, Yr8, Yr9, Yr12, Yr17, Yr18, Yr24, YrSu, YrSk
xiii
and YrA. Virulence for Yr resistance genes Yr2+, Yr3V, Yr3N, Yr5, Yr6,2+, Yr7+,
Yr9,2+, Yr10, Yr11, Yr15, YrSd, YrCv and YrSp was neither observed during the
glasshouse investigations nor prevalent at any of the six WSRTN sites. Although
resistance genes Yr4+, Yr8+, Yr26 and Yr29 (Pavon 76) expressed partial virulence but
still have prospective for exploitation. To deploy the identified Yr resistance genes either
singly or in combination in the upcoming wheat breeding program could play an
effective role to lessen yield losses inflicted by stripe rust.
xiv
1
Chapter I
GLOBAL WHEAT PRODUCTION AND RUST SCENARIO
1.1 GENERAL INTRODUCTION
The World Fact Book envisages that global human population has surpassed the
figure of 6.7 billion (6,706,993,152) (CIA, July 2008 est.). The world cereal
production for 2007-08 remained close to 2,127.1 million tons that include 1078.2,
609.4 and 439.6 million tons of course grains, wheat and rice, respectively. To come
up with the nutritional requirement of the earth's inhabitants for the instant year,
2,122.5 million tons segregating into 1,070.8, 614.9 and 436.8 million tons of course
grains, wheat and rice, respectively has been reported by FAO (2008). The figures
emphasize a production and consumption gap in wheat, which can be bridged by
espousing appropriate conservation tactics. The supplementary food thus attained
through the conservation measures will be adequate to gratify the starving
population, projected as 923 millions (FAO, 2008a).
1.1.1 The Wheat World
Wheat is the most extensively cultivated crop and is a staple food of nearly 35
percent of the world's inhabitants. Because of its increasing demand, the anticipated
global wheat requirement at the end of second decade of 20th century varies from 840
(Rosegrant et al., 1995) to 1,050 million tons (Kronstad, 1998). To accomplish the
projected target, world's wheat production will have to enhance from 1.6 to 2.6
2
percent, annually (Rajaram, 1999). Accordingly, the average global grain yield
would have to enhance from the existing 2.5 t ha-1 to 3.8 t ha-1. Only 18 countries in
the world meet target of producing average wheat grain yields of around 3.8 t ha-1 in
1995 whereby most of which are located in the Northern Europe (CIMMYT, 1996).
Of the worldwide 215 million hectares sown to hexaploid (Triticum aestivum) and
tetraploid (T. turgidum var. durum) wheat, 95 million hectares (44 percent) is in Asia
and out of which, 62 million hectares attributed to wheat are located in China, India,
and Pakistan (Singh et al., 2004b). Despite of such a vast area under the golden
grains production, the food security as well as production stability are of paramount
significance in most of the Asian countries as the majority of farmers are resource
poor.
The global location of Pakistan is in south-west Asia and its neighbouring countries
are Iran, Afghanistan, China and India. The federal capital of Islamabad is a unity
symbol of its four provinces: Sindh, Punjab, North-West Frontier Province (NWFP)
and Baluchistan. Pakistan engages a strategic position between Southern Asia and
the Middle East. Four mountain ranges as the Karakorams, the Pamirs, the
Himalayas and the Hindu Kush join within its jurisdictions (WCMC, 1991). Nature
has blessed Pakistan with a blend of climatic conditions and diversified land
including mountains, forests, deserts and the fertile plains. It experiences the most
extreme temperatures on the globe ranging from 50 °C in the Sindh region during
summers to minus 50°C in the northern mountain ranges in winters.
3
1.1.2 Wheat Rusts
Historically, the wheat rust diseases have emerged as foremost biotic production
constraints not only in Asia but through out the wheat world. The introduction of
semi-dwarf spring wheats during green revolution, engaged most of the South and
West Asian area in the 1960s have brought the stem rust caused by Puccinia
graminis, under control. However, Leaf rust (P. triticinia) and stripe rust (P.
striiformis) kept on posing a serious threat to wheat cultivation over an area of about
60 (63 percent) and 43 (46 percent) million hectares, respectively, in Asia provided
cultivation of the susceptible cultivars.
The combination of resistant cultivars with appropriate fungicidal treatment
depending on time, rate and number of applications (Line et al., 1983; Chen and
Wood, 2004; Padgett, et al., 2004) extends adequate rust control in the inoculum
prevalent region (Powelson and Halsey, 1982). Frequent and higher doses of
fungicides can also be toxic to the plants (Rakotondradona and Line, 1984) besides
being hazardous to environment and their use adds a lot to production costs.
Adoption of resistant cultivars in the rust prone areas is, therefore, the most effective
rust control strategy that does not add any cost to farmers and is environment
friendly. Considerable difference for virulence to particular resistance genes in
populations of extremely specialized rust fungi, mingled with the swift evolution of
new virulence has complicated its management (Singh et al., 2004b). Moreover, the
4
application of fungicides is not a long term economical solution to the rust disease
and the best long term alternate is the adoption of resistant cultivars.
1.1.3 Nature of the Rust Fungi
Rust fungi are the obligate parasites and must require living host to survive. The off-
season survival is either on self-sown (or voluntary) wheat plants or other grass
species. Moreover, high input, irrigated agriculture and cool highlands in various
parts of Asia promote the carryover of inoculum among seasons. Likewise, favorable
disease development conditions of temperature and high humidity that leads to dew
formation prevail during wheat growing season (Singh et al., 2004b).
Wheat rusts comprises of five spore stages where large number of Urediospores
production in the spring and summer, are epidemiologically significant. Wind plays
an important role in their dispersal to other plants, where they produce new
infections and secondary urediospores with a minimum interval of seven days
(Wiese, 1987). The nutrient-independent urediospores germinate as soon as they
make a contact with moisture films and penetrate the germ tubes directly in stomata.
Substomatal vesicles are thus formed and intercellular hyphae having globose or
lobed haustoria establish physiologic contact with host cell membranes thus
accomplishing infection process (Wiese, 1987). Since, mycelium have the potential
to remain viable at even -5 oC, therefore, stripe rust is appears first in the field as
compared to leaf and stem rusts.
5
1.1.4 Occurrence of Rusts in Pakistan
All the three wheat rusts occur in Pakistan with variable intensities under various
agro-ecological zones. Puccinia striiformis f.sp tritici, the wheat stripe rust pathogen,
is the most important among all wheat rusts, favored by cool summers, mild winters
and long, cool, wet springs. Its symptoms on the host include appearance of citron-
yellow uredia (spore masses) in long stripes over the leaf surfaces but rarely present
on stem and heads. As the crop matures, black spores (telia) are produced in stripes,
which are covered by the leaf epidermis (Smiley and Cynthia, 2003).
If a susceptible variety is under cultivation on large scale and weather turns
favourable for the stripe rust pathogen, the pathogen losses can be unbearable (Aqil
and Hussain, 2004). So far, four major stripe rust epidemics with intensity exceeding
20 percent have been reported during 1973 (35%), 1978 (55%), 1995 (37.5%) and
2003 (20%) while the stripe rust intensity has never gone below eight percent in the
country's history since 1950s (Ahmad, 2004). After the dawn of new millennium, the
rust intensity is heading towards an epidemic situation with every passing year.
During these epidemic years, the extensively cultivated resistant wheat cultivars
Pirsabak-85, Pak-81 and Inquilab-91 possessing resistance genes Yr7, Yr9 and Yr27
were brutally attacked by the corresponding stripe rust virulent races that rendered
the varieties susceptible. Stripe rust expressed its severity during 2004-05 in the
upper Punjab and NWFP and posed a serious threat against sustainable wheat
production. Such outbreaks has emphasized to avoid monoculture of a single wheat
6
varieties on large scale besides signified the importance of identifying yellow rust
resistant wheat varieties and their cultivation according to different ecological zones
of the country. Development and adoption of high yielding wheat varieties with
narrow genetic base, also influenced by farmer/consumer preference, has led to
cultivating fewer varieties over a large area that has created genetic vulnerability to
stress.
It is worth mentioning that chemical control of rust diseases is un-economical so
cultivation of resistant varieties is of immense significance, however, the presence of
numerous races of each and the ever-transforming nature of the pathogens obscure
breeding for rust resistance. During the years, many superior wheat cultivars have
been developed with an exceptional degree of resistance to one or more rust diseases
but the disease situation was never remained static which thus emphasis that
breeding of new strains of wheat resistant to rusts is an incessant process (Bariana et
al., 2007).
1.1.5 Objectives of the Research
Monitoring of the stripe rust population to determine pathotype variation was
imperative so that new virulences with the potential to overcome resistance genes
currently deployed in the wheat cultivars can be detected. The anticipated research
encompasses information pertaining to the occurrence, development and distribution
of prevalent stripe rust pathotypes in the Northern Punjab and NWFP. Moreover, an
attempt has also been made to assess the impact of stripe rust on commercially
7
cultivated wheat varieties through their response in terms of yield losses when the
disease occur in an epidemic form. Exploration in quest of resistant sources to stripe
rust has also been made by evaluating the breeders' material under the study.
Accordingly, the research was executed keeping in view the following objectives:
1. Conducting periodical surveys of wheat crop to identify prevailing virulences of
stripe rust pathogen in the areas under research and in-vitro study of pathotype
variation.
2. Ascertaining wheat yield losses due to stripe rust in field under disease stress
conditions.
3. Trapping of Puccinia striiformis f.sp tritici through “Trap Nurseries” planted at
selected sites in the Northern Punjab and NWFP.
4. Evaluation of breeders' material for its disease reaction under field condition.
8
Chapter II
PATHOTYPE VARIATION OF PUCCINIA STRIIFORMIS
2.1 INTRODUCTION
Wheat (Triticum aestivum L.) is one of the leading grain crops of Pakistan and being
staple diet of the inhabitants, it grasps a key position in the agricultural policies.
Wheat cultivation engages a major production area of 8.578 million hectares, which
encompasses approximately 34% of the cultivated area of the country and exhibits
production around 23.295 million tons (GOP, 2008). This enhanced production still
remained insufficient to ensure food security for the ever-escalating population
pressure. Sustainable productivity of wheat is of paramount importance in the
context of many biotic and abiotic factors that limit its production.
Stripe or yellow rust is one of such biotic factors, caused by an obligate parasite
Puccinia striiformis Westend. f. sp. tritici Eriks., that precincts wheat production
throughout the world. This pathogen reportedly infects numerous wheat and barley
cultivars as well as certain grass species (Stubbs, 1985). Stripe rust of wheat has
been reported from more than 60 countries of the world and all the continents except
Antarctica (Chen, 2005). Presence of several races of each and ever-changing nature
of the stripe rust pathogen cautions wheat cultivation in humid, high uplands and
cooler regions of the country. Varieties often appear to lose their resistance due to
cultivation in an area of its non-adaptability or change in virulence, which may either
8
9
be due to appearance of a new race(s) or change in the composition of the existing
races (Kilpatrick, 1975).
Based on the avirulence or virulence to cultivars or wheat genotypes, the P.
striiformis f.sp. tritici is further alienated into races. The race differentiation is based
on the types of infection they produce on a set of selected wheat genotypes or single
gene lines which are termed as differentials. The first report that “specialized
varieties” occur in P. striiformis based on specificity on genera of wheat was made
by Hungerford and Owens (1923), which later on was established by Allison and
Isenbeck (1930). Extensive race studies of P. striiformis f.sp. tritici were performed
by Gassner and Straib (1932) in Germany during 1930s and introduced a set of
differential genotypes of wheat. Fuchs (1960) in cooperation with Zadoks (1961)
used the set for race surveys in Europe, and later for international race surveys by
Stubbs (1985). Johnson et al. (1972) introduced the binary notation for races of P.
striiformis f.sp. tritici.
Development and use of resistant cultivars possessing diverse and well characterized
genes is integral for sustainable wheat production (Kaur et al., 2008). Globally,
virulence probably exists for most numbered Yr genes. So far, 40 stripe rust
resistance genes have been identified and officially designated as Yr1 through Yr40
(McIntosh et al., 2005; Kuraparthy et al., 2007) while 30 yellow rust resistance
genes have been provisionally designated as Yr followed by a letter symbol have
been reported (Chen, 2005). Majority of the catalogued genes are unique due to their
10
presence at different chromosomal location, responses to races and wheat genotypes
(Chen, 2005) while virulence has been identified for several of them. Virulence for
Yr9, which was widely deployed in bread wheat cultivars such as Seri82, Giza64 and
Maxipak, spread rapidly throughout the Middle East region, Yemen and Ethiopia in
the late 1980s (Wellings et al., 2000). After the breakdown of Yr9 resistance gene in
the final decade of the last century, several cultivars were released with the
incorporation of resistance gene Yr27. One such important and very well known
cultivar of the north-western Pakistan is Inquilab-91, which is now highly vulnerable
to the stripe rust disease (Singh et al., 2004a).
Wheat rust situation monitored by Khan and Mumtaz (2004) revealed rust epidemic
appearance during 1995 on wheat varieties Pak 81, Pirsabak 85 and afterwards on
Inquilab-91 during 2003. In another survey conducted during 2003-04, incidence of
the disease in Sindh, plains of Baluchistan and Southern Punjab, ranged between 5-
20MSS on different varieties. In central upper Punjab and in NWFP yellow rust was
observed 10MSS to 60S. Since the 2003 epidemic was mainly restricted to NWFP,
therefore, the major production areas of the Punjab escaped so no major losses were
recorded. Due to prevalence of highly conducive environment for stripe rust
development during April 2007 in Baluchistan, the stripe rust severity ranged from
5S-80S, 60S-80S, 5MS-80S, 5S-60S and traces to 80S in district Quetta, Mustung,
Pishin, Loralai and Qilla Saifullah, respectively (Hussain, Personal communication).
11
The stripe rust epidemics that occurred in the past, cautions that war footing efforts
to breed for stripe rust resistance is imperative in disease prone areas. Severe
epidemics on wide scale could be avoided through use of durable resistance and
diversification of resistance genes (Chen et al., 2002).
Monitoring of the stripe rust population is imperative to determine pathotype
variation so that new virulences with the potential to overcome resistance genes
currently deployed in the wheat cultivars can be detected early. Accordingly, the
present study was executed in the Northern Punjab and NWFP to encompass
following objectives.
1. Surveying of wheat crop to identify prevailing Pst virulence pattern and
pathotype variation.
2. Trapping of Puccinia striiformis f.sp tritici (Pst) through establishing “Trap
Nurseries” at hot spots of the study area.
3. Performance of commercial wheat cultivars to stripe rust at the WSRTNs.
4. Evaluation of breeders' material for its disease reaction under field condition.
12
Chapter II
2.2 REVIEW OF LITERATURE
Virulence factors of the yellow rust, Puccinia striiformis f.sp. tritici populations in
bread wheat have been analysed throughout the world. Virulence against the known
resistance genes as well as response of pathotypes to evaluate the resistance in wheat
lines have been studied by numerous scientists.
In this regard one of the studies was conducted in Turkey during 1991-2001 by Cetin
et al. (2002). They monitored changes in virulence pattern of the yellow rust
populations through field-grown trap nurseries, which consisted of wheat entries
with known resistance genes besides some cultivars from Turkey and around the
region including Haymana and Yenimahalle. The disease was developed well in
most of the years, however, high temperatures during 1992 and 1993 reduced rust
incidence. Stripe rust at Yenimahalle had low disease severity than Haymana.
Chinese 166, Vilmorin 23, Hybrid 46, Nord Desprez, Compair, Carstens V,
Spaldings Prolific and T. spelta album, carrying Yr1, Yr3a+, Yr4+, Yr3b+, Yr8, Cv,
Sp and Yr5 respectively, were resistant at both locations during these years. The
differential cultivars Lee, Heines Kolben and Kalyansona that carries Yr7, Yr6 and
Yr2 as well as the lines Anza, Sonalika and Gaby, remained susceptible during this
period. Other genotypes gave variable reactions to yellow rust. Virulence pattern of
12
13
stripe rust in 2001 undergone significant transformation as virulence for genotype
Moro was recorded for the first time in Haymana.
Variation was also observed in P. striiformis by Umirov et al. (2002) by assessing
resistance of 4,500 entries from ICARDA, CIMMYT and the Vavilov Institute,
Russia (VIR) under natural and artificial infection. Maximum rust resistance was
observed in 14 bread wheat (T. aestivum) lines but the highest rust resistance was
found in 11 durum wheat (T. durum) lines. They also found that the most effective
genes in Galla-Aral for yellow rust resistance were Yr3c, Yr5 and Yr4b.
To study occurrence and virulence of wheat stripe rust Manninger (2002) conducted
extensive survey in Hungry during 1999-2001 and reported the incidence only in
traces during 1999. His stripe rusts survey reports explained the importance of
pathotypes, whose virulence spectra (virulent for Yr2, Yr3, Yr6, Yr7, Yr8, Yr9, Yr17,
YrA+, YrCv and YrSd) were able to map the unknown resistance genes. He noticed
that 29 out of 78 Hungarian and foreign wheat cultivars were robustly infected by
yellow rust. Likewise, variation in the virulence for different isolates of P. striiformis
was assessed in Poland during 1999 to 2001 by Wozniak-Strzembicka (2003) after
testing the seedling of world and European differential set of wheat with the Yr
resistance genes. Virulence frequency for the resistance genes was found very high
to Yr1, Yr2, Yr3, Yr4, YrSd, YrCv, Yr2 and Yr6, Yr3a+Yr4a. It was generally low to
Yr10, YrSu, YrA and Yr6. Frequency of virulence for the Yr9 slowly progressed year
after year but virulence was not detected to the resistance genes Yr6, Yr7, Yr17 and
14
YrA. He also identified seven pathotypes in the yellow rust population. The
differential cultivars were also tested in the field under artificial inoculation with a
mixture of pathogen isolates. In the adult stage, the genes Yr6, Yr7, Yr6+Yr7, Yr17
and YrA provided full protection against the population of Pst.
The study executed by Flath and Bartels (2002) illustrated frequent virulence against
the resistance genes Yr1, Yr2, Yr3, Yr4, Su and Sd during 1998-2002 and observed
that genes Yr9 and Yr17 responded with partial resistance until 1998. Later on, in
1999 and 2000 the epidemics induced a resistance crash of the genes Yr9 and Yr17 as
well as of Yr7 and Yr8, respectively. They concluded that the most efficient
resistance genes left in Austria and Germany were Yr5, Yr10, Yr15, YrSp and YrA.
The most frequent rust pathotype was found in 34 percent of the German and 8
percent of the Austrian yellow rust populations, which contained a combined
virulence against Yr1, Yr2, Yr3, Yr4, Yr9, Yr17, Su and Sd. The pathotype was also
found in UK and Denmark explaining north-south wind transport.
Virulence gene frequency in wheat stripe rust populations was investigated by
Zeybek and Yigit (2004) during natural epidemics in the Southern Aegean and
Western Mediterranean regions of Turkey. They collected stripe rust uredinospores
from diseased plants and determined the virulence of pathogens by seedling test. It
was reported that virulence gene frequency on rust population varied depending on
the type of genes and those which showed high virulence frequency were more
stable. In both years, the virulence frequency of Yr1, Yr2, Yr3a, Yr9, Yr17, Sd and Su
15
genes ranged between 7.8 and 100 percent in all populations, while that of Yr6, Yr7,
Yr8 and Yr10 genes was observed in all locations.
Virulence to resistance genes Yr1, Yr2, Yr2+, Yr3V, Yr3Nd, Yr4+, Yr6, Yr6+, Yr7,
Yr7+, Yr9, Yr9+, Yr11, Yr12, Yr18, Yr24, Yr26 and those in the cultivars, Carstens V
(YrCv), Strubes Dickkopf (YrSd), Suwon 92/Omar (YrSu), Spaldings Prolific (YrSp),
Anza (YrA+) and Selkirk (YrSk) was identified in Ecuador between 1973 and 2004
by Ochoa et al. (2007). No virulence was found to Yr5, Yr8, Yr10 and Yr15. It was
also indicated that these cultivars carry single or the resistance factors Yr1, Yr2, Yr3,
Yr6, Yr9 in combinations as well as some other undesignated resistance factors.
Deployment of these resistance genes in the present cultivars of Ecuador has brought
an evolution in the yellow rust pathogen as a single deployed Yr resistance gene is
now not effective to the present stripe rust population in Ecuador.
Seedlings and adult plants reactions to P. striiformis races have also been evaluated
by Yildirim et al. (1995) after assessing a collection of 279 Triticum tauschii
accessions for resistance to yellow rust. They inoculated seedlings with four different
races of P. striiformis and found seventeen percent (44) of the T. tauschii accessions
resistant to all races of stripe rust. In another seedlings reaction study, Ma et al.
(1997) evaluated 75 accessions of Triticum boeoticum, T. monococcum, T. urartu, T.
turgidum L. var. durum (Durum wheats) and amphiploids (genome AAAABB)
involving crosses of three diploid species (AA) with T. turgidum (AABB) in the
greenhouse for their reaction to P. striiformis race 14E14. They also evaluated
16
Durum wheats and amphiploids at two field locations in Mexico with the same race
for their adult plant response. Resistant seedling reactions (infection types: 0-3 on a
0-9 scale) were seen for 10 (13%) accessions of T. boeticum, 19 (8%) accessions of
T. turgidum and 32 (25%) amphiploids. The remaining accessions were either
moderately resistant (ITs 4-6) or susceptible (ITs 7-9). The three amphiploids
derived from the crosses of seedling resistant T. boeoticum and T. turgidum, were
resistant as seedlings. Among the 51 amphiploids involving one resistant parent, 29
were found resistant while remaining 22 displayed intermediate to susceptible
reactions.
Wheat response by employing five pathotypes of yellow rust, 14E176A super(+),
134E142A+, 6E210A+, 4E128A- and 64E146A+ was assessed by Torabi and
Nazari, (1998) at seedling stage under greenhouse conditions at Karaj, Iran and as
adult plants in field conditions at four locations. Their results showed that few
cultivars possess seedling or overall resistance to the pathotypes and some have just
adult plant resistance. Cultivars M-70-4 and MV-17 were resistant to all pathotypes
at seedlings and exhibited desirable adult plant resistance. Later on, about 415 bread
wheat genotypes of the Preliminary Wheat Screening Nursery of Iran were tested for
yellow rust resistance at the seedling stage under controlled conditions as well as
adult plant stage in the field (Miandoab, Moghan and Gharakhil areas) during 1999-
2000 by Malihipour et al. (2002). Tests were carried out with three P. striiformis
pathotypes: 230E62A+ (Miandoab), 134E134A+ (Moghan) and pathotype 38E2A+
(Gharakhil). In the seedling tests, about 296 accessions (71.3% of the entries) were
17
found susceptible to 230E62A+ while those susceptible to 134E134A+ and 38E2A+
comprised 283 (68.2%) and 199 accessions (48.0%), respectively. About 149
accessions (35.9%) were susceptible regardless of the pathotype used. Resistance to
38E2A+, 134E134A+ and 230E62A+ were observed in 91, 49 and 60 accessions
(21.9, 11.8 and 14.5 percent), respectively. Thirty-six accessions (8.7%) were
resistant to 230E62A+ and 134E134A+ while thirty-five accessions (8.4%) recorded
the same reaction to 230E62A+ and 38E2A+. Thirty-two accessions (7.7%) were
resistant to 134E134A+ and 38E2A+. Meanwhile, adult plant tests revealed that most
of the accessions were yellow-rust resistant in one or more locations. Some 273
accessions (65.8%) were resistant at all locations. Fifty-five accessions (~13%) with
infection types of 0-3 (resistant) as seedlings were resistant or moderately resistant to
Miandoab isolate at the adult plant stage. Some accessions with infection types of 4-
6 (intermediate) as seedlings were resistant or moderately resistant at the adult plant
stage, including 49 accessions (11.8%) for Miandoab, 71 accessions (16.7%) for
Moghan, and 118 accessions (28.4%) for Gharkhil isolates. Some accessions with
infection types of 7-9 (susceptible) at the seedling stage were resistant or moderately
resistant at the adult plant stage, including ~52% of the entries for Miandoab, 59%
for Moghan, and 47% for Gharakhil isolates.
Afshari (2008) later on in Iran identified 27 pathotypes during 2003 and 2004 in
greenhouse tests whereby pathotypes 6E6A+, 6E22A+, 6E130A+, 6E134A+,
6E142A+, 6E158A+, 134E130A+ and 134E142A+ remained most common during
experimentation study. He detected virulence on plant/s with gene/s Yr2, Yr6, Yr7,
18
Yr8, Yr9, Yr24, Yr25, YrSD, YrSP, Yr3N, Yr2+, Yr6+, Yr9+, Yr7+, Yr32+ and YrA
under greenhouse conditions and observed that majority of isolates with a high
frequency (more than 88 percent) showed virulence on plant/s with Yr2, Yr6, Yr7,
Yr9, YrA and Yr24 genes. No virulence was detected on plant/s with Yr1, Yr3V, Yr4,
Yr5, Yr10 and YrSU genes.
From ICARDA germplasm, the resistance in 100 advanced wheat lines was
evaluated in seedling and adult plant stages by Alibaba et al. (2002). They inoculated
five seedlings from each entry separately with uredinospores of four yellow rust
races (6E0A-, 17E182A, 134E134A+, 142E150A+) after their first leaves had been
completely expanded. Data was recorded on the basis of Infection type (IT) for each
entry 15 to 17 days after inoculation using the McNeal et al. (1971) scale. They
classified the lines into two groups as 66 percent with major and 34 percent with
minor resistance genes in the seedling stage. Adult plant studies were conducted in
the field in Miyandoab Research Station during 1998 whereby entries were
inoculated with 142E150A+ stripe rust race. After sufficient disease development on
flag leaf of the susceptible cultivar (Bolani), IT and disease severity of each line was
recorded at three-week intervals by calculating Relative Area under Disease Progress
Curve (RAUDPC) and Final Disease Severity (FDS) for each entry. Cluster analysis
using the UPGMA method showed that 62% were susceptible in seedling stage and
resistant in the adult stage (APR), 6% were SS, 35 were RS and 29% were RR. They
observed a significant correlation coefficient between infection type of the first leaf
and flag leaf RAUDPC against 142E150+.
19
Su, et al. (2003) after testing 57 isolates of P. striiformis from western Canada for
their virulence on the 17 world and European differentials as well as 7 supplemental
differentials, identified thirty-nine races. New races detected in Canada since 2000
were virulent on both 'Compair' (Yr8, Yr19) and 'Clement' (Yr9,Cle). Race
33(238E150) was repeatedly detected from 2000 to 2002 and other new races had a
virulence spectrum that was similar to that of race 33. None of the races were
virulent on Triticum spelta var. album (Yr5). 'Chinese 166' (Yr1), 'Moro' (Yr10,Mor),
'Hybrid 46' (Yr3b,Yr4b), T. dicoccoides selection G-25 (Yr15), and 'Spalding Prolific'
(YrSp) were resistant to more than 90 percent of the races detected in western
Canada and all of them were resistant to the new races detected since 2000. They
were of the view that climatic conditions and cultivar resistance played a significant
role in the survival and propagation of these races on the Canadian prairies.
Different trials have been conducted to identify stripe rust resistance genes in wheat
cultivars by using isolates of P. striiformis. One of such trials was conducted in
China where 52 wheat cultivars were studied by using 13 isolates of Pst and 19
known yr genes by Jing, et al. (2005). They detected twelve genes, Yr1, Yr2, Yr6,
Yr7, Yr8, Yr9, Yr11, Yr12, Yr17, Yr18, Yr24 and Avocet in 28 wheat cultivars, either
alone or in combination. Seventeen cultivars were found to have Yr11 gene while 15
contained Yr2, Yr6 and Yr17. Wheat resistance sources such as 92R137 and 92R149
were resistant to all the 13 isolates, including the dominant races of P. striiformis in
China. The commercial wheat cultivars such as Shan 160, Jinan 17, Yan 188, Yan
361 carrying no resistance genes at all, were unable to resist any of the 13 isolates.
20
Afshari and Turabi (2005) also conducted experiment in Iran during 2002-03 to
verify the resistance of nine wheat cultivars, along with the susceptible control
cultivar Bolani, to evaluate stripe rust pathotype 134E134A+ in a field. They found
all the cultivars susceptible to the pathotype at the seedling stage. Contrarily, under
field conditions, the cultivars were either resistant or moderately susceptible to the
pathogen, except for Japateco S and Bolani. Cultivars carrying the Yr29 gene were
resistant, whereas cultivars possessing Lr34 gene were moderately susceptible.
Wellings (2008) confirmed the first Australian pathotype with virulence for the Yr27
resistance from a single sample of stripe rust collected from northeast Victoria. The
resistance gene Yr27 is present in several commercial varieties, which will now be
under a degree of uncertainty until the expected field responses will be determined.
The evidence indicates that the new pathotype arose as a single mutation derivative
from the ‘Jackie’ pathotypes, which is designated 134 E16 A+ J+ Yr27+ and is given
the common name ‘Jackie Yr27’ pathotype.
To evaluate the expression of adult plant resistance in the green house, from seedling
to anthesis, Ma and Singh (1996) used a Mexican race of P. striiformis and also
assessed the effect of such resistance on stripe rust development in the field.
Resistance was measured by infection types (ITs). The cultivars were also assessed
in the field using the same race to determine the disease progress. In all tests,
Morocco was the most susceptible cultivar (IT 7 to 9), followed by Avocet S (IT 6 to
7) and Jupateco 73S (IT 6 to 9). They, however, observed that as the plants grew,
resistance was increased and expressed as lower IT. The change in IT displayed by
21
HD2258 and PBW65 were the greatest as intermediate to low IT (2 to 3) at
completion of tillering which changed to immunity (IT 0) at anthesis. Other cultivars
that expressed adult resistance were Mexico 82, Pavon 76, Jupateco 73R and Chino
79 etc. In the field study, HD2258 and PBW 65 were found highly resistant while
Pavon 76 and Mexico 82 had acceptable resistance level, whereas other cultivars
displayed moderate resistance level.
Experiments to evaluate the level and distribution of wheat stripe rust in Kazakhstan,
Kyrgyzstan and Uzbekistan and to identify the resistance genes in commercial
cultivars or new lines adapted to the region were conducted during 2000 by
Absattarova et al. (2002). They carried out disease surveys during the 2000 spring-
summer seasons in Kazakhstan, Kyrgyzstan and Uzbekistan. They reported that
severe stripe rust was occurred in most regions of Central Asia in 1999 but was less
severe in 2000. Wheat stripe rust survey carried out in their observed areas indicated
the differences between levels of disease development that depended on genotypes
and locations. In their field trials, the winter wheat cultivars from Kazakhstan that
included 25 entries, planted in the demonstration plots were subjected to naturally
occurring yellow rust races. They observed that most of the released commercial
cultivars were susceptible to yellow rust, however, cultivars such as Almaly, Arap,
Aksham and entries 245-99, 2909 and 345-99 were found resistant.
Severe stripe rust infection causes shriveling of grains though the grain is sound
otherwise. Due to the phenomenon physical qualitative characterizes of wheat such
22
as test weight and bulk density are negatively influenced. Besides grains are
sustained which impair market value and quality of grains. To cope with these
problems plant pathologists and the farmers had been trying to sow germplasm from
different sources to find out way to escape losses caused by this disease. Main
emphasis had been to exploit indigenous resources for developing genetically diverse
wheat varieties suitable for varying temperatures, humidity levels, soil
characteristics, slopes and altitudes (Feyissa et al., 2005).
The timely detection of new variants and the identification of resistant material
through strong and vibrant program of monitoring wheat rust virulences have
curtailed any major loss to wheat during the last 35 years as reported by Singh
(2009). The efforts made by his research directorate for managing wheat rusts
involves only the use of available resistance and its intelligent deployment based on
pathotype distribution. The directorate has diversified genetic base of rust resistance
in wheat through pre-breeding activities and has planned to flow the new/unutilized
rust resistance genes Yr5, Yr10, Yr15, Yr(CD), Yr(Mega), Yr(Hobbit), Yr(SP),
Yr(China-84) into the future varieties, which will reduce the danger of rust
epidemics. Chen (2005), Afshari (2008) and Chunmei et al. (2008) has also reported
that virulences to genes Yr5 and Yr15 occur as a rare phenomenon in wheat growing
regions of the globe.
The breeding programs for combating stripe rust disease have been reliant on the
vital genes as well as to respond towards the substantial cut down of resistance
because the fungus has the ability to mutate and overcome the varietal resistance.
23
Combined use of the significant genes that, independently, have previously been
‘matched’ with the pathogen may provide another level of resistance but this too is
potentially susceptible to breakdown and invariably short-lived. Combinations of
partial resistances, which tend to have durability, are now considered to be safer
options. However, understanding and working with such resistance requires a deeper
knowledge of the genetic basis of the resistance. Breeders are now mapping the
major genes involved and identifying varieties with known combinations. This gives
the breeder the opportunity to define the risk associated with any known combination
in varieties under test. In addition the use of molecular markers to tease apart the
contributions of partial resistances now provides an extremely useful tool for
overcoming the threat of this disease. Longer term robust resistance based upon the
‘stacking’ of different sources with a proven level of durability must be the best
solution. The effects of moderate resistances can be additive and combined to
provide near immunity (Angus and Fenwick, 2008).
24
Chapter II
2.3 MATERIALS AND METHODS
Samples of wheat leaves infected with stripe rust were collected during 2005-06 and
2006-07 from eight districts of wheat growing areas of the Northern Punjab (Sialkot,
Chakwal, Rawalpindi and Attock) and NWFP (Nowshera, Peshawar, Mardan and
Charsadda) as well as from the Wheat Stripe Rust Trap Nurseries (WSRTN)
established at six yellow rust hot spots. The virulence pattern was determined in the
glasshouse at Crop Disease Research Program (CDRP), NARC, Murree by using the
disease inoculum collected during both the years on a set of near isogenic lines
(NILs) with Avocet-S background along with a universal susceptible check,
Morocco. Four commercial varieties (Inquilab-91, Bakhtawar-93, Wafaq-2001 and
MH-97) were also added in the differentials to observe their response against the
prevalent stripe rust virulences. Data on Infection Types (ITs), based on a 0-9 scale
(McNeal et al., 1971) was recorded in the glasshouse while adult plant reaction in
response to severity on the NILs with avocet background was evaluated in WSRTN
on the basis of percent leaf area infected according to the modified Cobb’s scale
(Peterson et al., 1948).
To monitor occurrence, development and distribution of stripe rust in the Northern
Punjab and NWFP, wheat growing areas were frequently surveyed from 2nd week of
February to the 1st week of April during 2006 and 2007. In the Northern Punjab,
24
25
wheat engaged areas of district Sialkot, Chakwal, Rawalpindi and Attock while in
NWFP, the wheat growing areas of district Nowshera, Peshawar, Mardan and
Charsadda were surveyed during this period.
2.3.1 Modus Operandi for Field Sampling of Stripe Rust
Farmers’ fields at various stripe rust prone areas in different districts of Northern
Punjab and NWFP were surveyed for disease sample collection. The survey was
executed with first field after 10 kms on the car-odometer and the subsequent stops
were made for each sample every 30 kms on either side of the road. The stripe rust
sampling was conducted with a general procedure in the diagonal transect for each
field, sample were taken at 3 points along a diagonal (Fig.1) and the incidence was
calculated by using the following formula.
Disease Incidence = Diseased Plants
x
100 Total number of plants observed
Fig. 1 Field design for stripe rust disease assessment and sampling
2.3.2 Collection of Diseased Samples from Farmers’ Fields
Farmers’ fields in the wheat growing areas of Sialkot, Rawalpindi, Chakwal, Attock,
Nowshera, Charsadda, Peshawar and Mardan were surveyed during 2005-06 and
26
2006-07 to monitor the disease occurrence and to collect representative samples
along major roads. In each sampling area, leaves with heavy infections were
collected and then pressed in the folds of newspaper before placing in glycine
envelope for their transportation to CDRP Laboratories, Sunny bank, Murree. These
samples were stored in the refrigerator till further analysis.
2.3.3 Establishment of WSRTNs
The WSRTNs were established each year during 2005-06 and 2006-07 to trap the
virulences present in different wheat growing areas of the Northern Punjab and
NWFP. The WSRTNs were surveyed on regular basis during both the wheat growing
seasons to monitor the occurrence, development and distribution of yellow rust. The
selected six locations (Table 1) for establishment of WSRTN served as hot spots for
development of yellow rust.
Table 1 Location of Trap Nurseries Established under the Research Area
Province Location
Punjab Pulses Research Station (PRS), Sahowali, Sialkot;
PMAS - Arid Agriculture University (PMAS-AAUR), Rawalpindi; and
National Agricultural Research Centre (NARC), Islamabad.
NWFP Cereal Crop Research Institute (CCRI), Pirsabak, Nowshera;
NWFP Agricultural University (AUP), Peshawar; and
Nuclear Institute for Food and Agriculture (NIFA), Tarnab, Peshawar.
The WSRTN included two susceptible wheat cultivars Local White and Morocco,
standard world and European set of differential genotypes, single Yr NILs with
27
avocet background to describe virulence or avirulence patterns of isolates of
Puccinia striiformis f. sp. tritici (Pst) (Boshoff et al., 2002; Hovmøller et al., 2002;
Wellings and Kandel, 2004; Chen, 2005). In addition to these, 57 Pakistani
commercial wheat cultivars were inducted in the WSRTN to observe adult plant
reaction in response to severity of the prevailing virulences. Detail of WSRTN,
which comprised of 34 entries (Appendix I) including 17 world and European
differentials, 17 NILs with avocet background (Table 5) and Morocco as a
susceptible check along with 57 commercial cultivars acquired from CDRP, NARC,
Murree is given in Table 2.
2.3.3.1 Stripe rust monitoring
Trap plots for monitoring wheat stripe rust were planted at stripe rust prone areas,
which serve as the hot spots for the development of yellow rust. Each entry of the
nursery was planted in a single meter row length, 30 cm apart. Two rows of
universally susceptible spreaders consisting of Local White and Morocco were
planted around the nursery. In addition, a row of susceptible check (Morocco) was
also raised at every 20th entry. The material was screened in natural conditions and
no artificial inoculation was made. Incidence and severity of stripe rust were
recorded at all the locations according to Loegering (1959). The observation for
severity and reaction were recorded together with severity first. Data for the Trap
nursery at all locations was recorded thrice during the month of March, but was kept
under regular observation from February till April of 2006 and 2007.
28
Table 2 List of Commercial Varieties included in WSRTN (2005-06 & 2006-07)
Sr. No.
Commercial Varieties
Yr Resistance
Genes
Year of Release Parentage / Pedigree
1 Morocco - NA NA
2 Pavon 76 Yr29 APR
1979
VCM//CNO/7C/3/KAL/BB, CM8399-D-4M-3Y-1M-1Y-1M-0Y-0PAK
3 MH 97 NA 1998 ATTILA, CM 85836-50Y-0M-0Y-3M-0Y
4 Inquilab-91 Yr27 1991 WL711/CROW 'S' PB1954-9A-1A-0A-0PAK(PAK)
5 Kohistan 97 NA 1998 V-1562//CHRC 'S'/HORK/3/KUFRA/4/CARP 'S'/BJY 'S' PB24883-B-1A-0A
6 Punjab 85 NA NA KVZ/TRM//PTM/ANA, CM43903-H-4Y-1M-1Y-3M-3Y-0B-0PAK
7 Bakhtawar 93 Yr9 1995 JUP/BJY 'S'//URES
8 Blue Silver NA 1971 153-388/AN/3/YT54/N10B//LR64/AN//YT54/ N10B/3/LR864/4/B4946. A.4.18.2.1Y-Y53//3/Y50A.4.18.2.1Y53//3/Y50
9 Chakwal 86 NA 1986 F1n/ACS//ANA SWM4578-56M-3Y-3M-0Y-0PAK
10 Sindh-81 NA 1983 NORTENO/MEXIPAK
11 Zarghoon Yr6+ 1979 CC/INIA/3/TOB/CTFN//BB/4/7C CM8237-G-1M-3Y-2M-4Y-0M-OPAK
12 Faisalabad 83 Yr7+2 APR 1983 FURY/KAL/BB CM37138-48Y-1M-5Y-1M-4V-5Y-0A-0PAK
13 Faisalabad 85 Yr9 + Yr4 1985 MAYA/MON//KVZ/TRM CM44083-N-3Y-1M-1Y-1M-1Y-0B
14 Kaghan 93 Yr9 1993 TTR/JUN CM59123-3M-IY-2M-1Y-2M-2Y-0M-0PAK
15 Kirin 95 NA NA NA
16 Kohinoor 83 Yr9 1983 OREF1158/FDL/MFN/2*TIBA63/3/COC CM37987-1-1Y-5M-0Y-0PAK
17 LU-26 Yr6 1976 BLS/KHUSHAL
18 Nowshera 96 NA 1996 NA
19 Parwaz 94 Yr6+Yr7 1995 V.5648/PRL PBB20089-7A-4A-0A
20 Pasban 90 NA 1991 INIA F 66/ A DISTCHUM//INIA66/3/GEN
21 Pirsabak 85 Yr9 1985 KVZ/BUHO//KAL/BB CM33027-F-15M-4Y-4M-2Y-1M-1Y-0M-OPAK
22 Punjab 96 NA 1995 SA42*2/4CC/INIA//BB/3/INIA/HD832 Pb1352-B-4K-36A-0A
23 Sariab-92 Yr6+ 1993 BB/GLL//CARP/3/PVN CM33483-C-7M-1Y-0M-OPAK(PAK)
24 Sarsabz Yr7 1985 M20/79 S89-75-76-RS
25 Shaheen 94 NA NA MLT''S"
26 Shahkar 95 Yr6+ 1995 WL711//F3.71/TRM Pb 20371-20A-4A-0A-0K-0A
27 Soughat 90 Yr6+Yr7 1991 PAVON MUTANT-3
28 Tandojam 83 Yr6+ 1985 TZPP/PL/7C CM2587-J-1Y-2M-2Y-3M-OY
29
29 SH-2002 NA 2002 INQILAB-91/FINK'S' PB.25552-1A-0A-0A-1A-0A.
30 Pak 81 Yr9+Yr7 1981 KVZ//BUHO//KAL/BB CM33027-F-15M-500Y-0M-76B-0Y-0PAK
31 Bahawalpur-97 NA 1998 MLT'S' CM47634-1-2M-2Y-1Y-1M-0Y
32 Kohsar 93 NA 1995 PSN/BOW CM69560-1M-1Y-1M-2Y-0M-0PAK(PAK)
33 Rohtas 90 Yr9 1991 INIA F 66/A.DISTCHUM//INIA66/3/GEN W.8461-R-OPAK(PAK)
34 Suleman 96 NA 1996 F6.74/BUN//SIS/3/VEE#7 CM86141-62-0Y-0M-4Y-0M
35 WL 711 Yr2 1978 S308/CHRIS//KAL
36 Zardana NA NA C/CNO67/8156*Tob66-CN067/Nor66/II12300*LR64-8156/PVN76'S;
37 Abadgar 93 NA 1993 NA
38 Anmol-91 Yr9 1991 KVZ/TRM//PTM/ANA CM43903-H-4Y-1M-1Y-3M-3Y-0B-0PAK
39 Bahawalpur-2000 NA 2001 NA
40 Bahkhar-2002 NA 2003 P20102/PIMA/SKA/3/TTR`S'/BOW`S' ,Pb.23826-D-1a-1a-1t-1t-0t.
41 Fakhr-e-Sarhad NA NA KVZ/BUHO//KAL/BB CM33027-F-15M-500Y-0M087V-0Y
42 Marvi-2000 NA 2001 CMH-77A917/PKV 1600//RL6010/68SKA
43 Mehran-89 Yr9 1988 KVZ/BUHO//KAL/BB CM33027-F-15M-500Y-0M-87V-0Y
44 Soorab-96 (Barley) NA NA NA
45 Tatara Yr3 1999 ATTILA CM85836-50Y-0M-0Y-2M-0Y
46 Takbeer NA 1999 NA
47 AS-2002 NA 2002 KHP/D31708//CN74A370/3/CIAN079/4/RL6043/*4NAC PBD795-23A-1A-0A
48 Iqbal 2000 Yr9 2000 BURGUS/SORT 1213//KAL/BB/3/PAK 81 PB 21912-11A-0A-0A-59A-0A-0A
49 Auqab-2000 Yr9 2000 CROW 'S'/NAC//BOW 'S' PB 222138-3A-0A-0A-231A-0A
50 Chakwal-97 NA 1998 BUC'S'/FCT'S' CM64663-7M-0Y-0M-7Y-OM.
51 Durum-97 NA NA NA
52 Watan 94 NA 1994 Lu26/HD2179
53 Moomal 2002 NA 2002 BUC'S'/4/TZPP//IRN46/CN067/3/PRI-FLAKE-56744 7Y-2Y-1M-1Y-0M
54 Zarlashata Yr9 2001 URES/BOW`S’ CM 78108-1M-02Y-02M-22Y-3B-0Y.
55 GA-2002 NA 2002 DWL 5023/S N B//SNB CM 84986-H-1M-3M-2B-0Y.
56 Wafaq-01 Yr9 2001 OPATA/RAYON//KAUZ CMBW 90Y3180-0TOPM-3Y-010M-010M-010Y-1M-015Y-0Y.
57 Margalla-99 NA 1999 OPATA/BOW'S' CM 83398-2M-0Y-0M-5Y-0M
58 Manthar Yr9 2002 KAUZ//ALTAR84/AOS CM 111633-6M-20Y-10M-10Y-10M-2Y-0M-0B.
NA refers to "Not Available"
30
2.3.3.2 Recording of observations at WSRTN
At all the WSRTNs locations, observations regarding prevalence of stripe rust in
terms of percent of infected plants with response and severity were recorded
according to Loegering (1959). The stripe rust severity was recorded on WSRTN
entries and in commercial wheat fields as percent of the rust infection on the wheat
plants according to the modified Cobb’s scale (Fig. 2) (Peterson et al., 1948) and the
field response scale referred to the infection type as described in Table 3 (Fig. 3)
(Mcintosh et al., 1995; Roelf et al., 1992).
Source: Knott, 1989 Fig. 2 The modified Cobb’s scale:
A: refers to the actual percentage occupied by rust uredinia; and
B: pertains to rust severities of the modified Cobb’s scale after Peterson et al.,
1948.
31
Table 3 Field response of host plant against stripe rust
___________________________________________________________________________________ Reaction Description Observation R. Value
No Disease No visible infection O 0.0
Resistant Visible chlorosis or necrosis, no uredia are present R 0.2
Res. – Mod. Resistant RMR 0.3
Moderately Resistant Small uredia surrounded by chlorotic/necrotic areas MR 0.4
Mod. Res. – Mod. Sus MR-MS 0.6
Moderately Susceptible Uredia medium size with no necrotic margins but MS 0.8
possibly some distinct chlorosis
Mod. Sus.-Susceptible MS-S 0.9
Susceptible Large uredia with no necrosis & little or no chlorosis S 1.0
Fig. 3 Adult Plant Host Response to the Stripe rust
2.3.4 Establishment of WDSN
The Wheat Disease Screening Nursery (WDSN) comprising of 188 entries including
14 commercial varieties, 174 breeders’ lines were raised during wheat growing
season of 2005-06 and 2006-07 for their reaction to stripe rust. The WDSN was
established at the experimental area of PMAS-AAUR.
32
Each entry of the nursery was planted in a 5 meter row length, 30 cm apart. Two
rows of Local White and Morocco, as susceptible check were planted around the
nursery. The material was screened in natural conditions and no artificial inoculation
was carried out.
Observation on response and severity of stripe rust were recorded according to
Loegering (1959). The severity was recorded as percent of rust infection on the
plants according to the modified Cobb’s Scale (Peterson et al., 1948).
2.3.5 Glasshouse Virulence Analysis
Out of 354 stripe rust samples collected during the entire study period from farmer’s
fields and WSRTNs, 189 samples revived on the universal susceptible cultivar
Morocco. Virulence analysis of 150 isolates (73 and 77 samples pertaining to 2005-
06 and 2006-07, respectively) which were randomly selected from 189 revived
samples, was conducted at CDRI labs., Murree (Table 4). The method described by
Line and Qayoum (1992) was adopted to determine prevailing races of Pst through
employing seventeen wheat cultivars as differentials (Table 5).
2.3.5.1 Inoculum increase
Wheat seeds were planted in disposable pots containing a mixture of peat and soil
inside the rust-free glass house. Uredineospores from the already collected samples
were transferred by using smear technique to the seedlings at two-leaf stage (Zadoks
et al., 1974) of the susceptible wheat cv. Morocco to increase spore. Inoculated
plants were kept in a dew chamber containing 98 percent relative humidity
33
conditions at 10°C for 48 hours and then transferred to growth chamber programmed
to change temperature steadily from a minimum of 2 to 5°C during the dark period of
8 hours and from a maximum of 18 to 20°C during the light period of 16 hours. The
natural daylight was supplemented with metal halide lights to extend the photoperiod
up to 16 hours (Chen et al., 2002). Cross contamination was checked by isolating the
inoculated plants with plastic booths.
Table 4 Locations for Stripe Rust Samples Collected (Natural Infections) during
2005-06 and 2006-07
Province District
Sampling Year 2005-06 Sampling Year 2006-07
Samples* Collected
(Nos.)
Inoculum Revived (Nos.)
Samples* Collected
(Nos.)
Inoculum Revived (Nos.)
Punjab Sialkot 21 8 24 12
Rawalpindi 36 19 31 16
Chakwal 12 8 16 9
Attock 18 7 23 13
NWFP Nowshera 31 15 29 16
Charsadda 19 10 15 8
Peshawar 23 14 26 18
Mardan 13 9 17 7
TOTAL 173 90 181 99
* Include samples from farmers' fields as well as WSRTNs.
Sporulating streaks of stripe rust appeared on the inoculated leaves after 15-17 days
of inoculation. Urediniospores were collected separately for each disease sample in #
0 capsule using a separate cyclone collector attached to a vacuum pump (Line and
Qayoum, 1992).
34
Table 5 Differential set for Race Determination and Virulence Analysis of
Stripe Rust pathogen at CDRI, Murree
Sr. #
Differentialsa Resistance
genes Sr.
# Near Isogenic Linesb with
Avocet background Resistance
genes
World set 18 Yr1/6 Avocet Yr1
1 Chinese 166 Yr1 19 Yr5/6 Avocet Yr5
2 Lee Yr7 20 Yr6/6 Avocet Yr6
3 Heines Kolben Yr6,Yr2 21 Yr7/6 Avocet Yr7
4 Vilmorin Yr3V 22 Yr8/6 Avocet Yr8
5 Moro Yr10 23 Yr9/6 Avocet Yr9
6 Strubes Dickkopf YrSd 24 Yr10/6 Avocet Yr10
7 Suwon92 x Omar YrSu 25 Yr11/3 Avocet Yr11
8 Clement Yr9,Yr2+,Cle 26 Yr12/3 Avocet Yr12
9 Triticum spelta Yr5 27 Yr15/6 Avocet Yr15
28 Yr17/6 Avocet Yr17
European set 29 Yr18/3 Avocet Yr18
30 Yr24/3 Avocet Yr24
10 Hybrid 46 Yr4+ 31 Yr26/3 Avocet Yr26
11 Reichersberg42 Yr7+ 32 YrSP/6 Avocet YrSp
12 Heines Peko Yr6,Yr2+ 33 Yr27/3 Avocet YrSk
13 Nord Deprez Yr3N 34 Jupateco R (Yr18) Yr18
14 Compair Yr8,YrAPR 35 Jupateco S -
15 Carstens V Yr32,YrCv 36 Avocet R (YrA) YrA
16 Spaldings prolific YrSp Commercial Varieties
17 Heines VII Yr2+ 37 Morocco -
38 Inquilab-91 Yr27
39 Bakhtawar 93 Yr 9+
40 Wafaq-01 Yr9
41 MH-97
a The wheat genotypes used in differential sets to identify prevailing pathotypes. b NILs used to study prevailing virulences against the known Yr resistance genes.
2.3.5.2 Isolate development
The fresh collected urediniospores of Pst suspended in the mixture of 30:70 mineral
oil : petroleum ether, were used with fine atomizer to inoculate 10 days old seedlings
35
of differential sets that are used to differentiate races of Pst, 19 near isogenic lines
with avocet background and four commercial varieties grown in 15 x 18 cm trays
(Table 5). Morocco was included as a susceptible check. The set was transferred to
growth chamber with programmed conditions as mentioned above where each set
was again covered with polyethylene to avoid cross contamination. Data on infection
types (ITs) as described by Johnson et al. (1972) were recorded after 15 to 17 days of
inoculation when susceptible check developed maximum infection (Fig. 4). An
isolate was considered avirulent on a specific differential cultivar when there were
no symptoms (IT 0) or there were necrotic or chlorotic flecks (IT 1), necrotic or
chlorotic blotches without sporulation (IT 2), or necrotic or chlorotic blotches with
only a trace to slight sporulation (IT 3 to 5). An isolate was considered to be virulent
if it caused moderate to abundant sporulation, with or without chlorosis or necrosis
(IT 6, 7, 8 or 9).
2.3.5.3 Stripe rust assessment on NILS
The infection types for all the isogenic lines were recorded upon appearance of
pustules after 15 to 17 days. The infection types for all the NILs were recorded using
standard scoring scale 0-9 (McNeal et al., 1971) (Fig. 4). The virulence patterns on
differential sets were assessed on the basis of low infection types produced by each
line in response to infection (Infection type 0, 1, 2, 3, 4 and 5 represented avirulent
while 6 to 9 expressed virulent response) (Fig. 4).
36
A 0 VR R MR LM M HM MS S VS
B 0 1 2 3 4 5 6 7 8 9
Fig. 4 Scale 0-9 for Stripe Rust Disease Scoring at Seedling Stage
Letter 'A' referrs to "code symbol" while 'B' indicate an "index value"
2.3.6 Virulence Assessment
Differential wheat genotypes with known resistance genes for seedling were used for
the determination of isolate's virulence spectra. These contain nine entries from the
European and eight entries from the world sets (Johnson et al., 1972) besides NILs
with the Avocet S background (Wellings et al., 2004) (Table 5). The entire
differential sets were provided by Dr. Iftikhar Ahmad from CDRP, NARC, Murree
during 2005-06 and 2006-07. These differential wheat genotypes permitted the
detection of virulences corresponding to the resistance genes Yr1, Yr2+, Yr3V, Yr3N,
Yr4+, Yr5, Yr6, Yr7, Yr8, Yr8+, Yr9, Yr9+, Yr10, Yr11, Yr12, Yr15, Yr17, Yr18,
Yr24, Yr26, YrSd, YrSu, YrCle, YrCv, YrSp, YrSk and YrA.
2.3.7 Designation of Pathotype
The pathotypes were designated on the basis of a binary notation system developed
by Johnson et al. (1972) with some additions made by Wellings et al. (1988) for race
37
nomenclature. This system codes a considerable amount of information in a single
number. In this procedure, the differential hosts have been listed in a permanent
order and allocated fixed decanary numbers that are the successive powers of 2 as
shown in (Appendix II). Rust reactions were classified as resistant with binary score
‘0’ or susceptible having binary score '1'. The sum of all decanery values of only
susceptible cultivars was added to acquire a race number for the isolate and their
reverse translations to the binary code render the virulences spectrum. The second
number was headed by the letter E to indicate the wheat genotypes pertaining to
European series. Each blend of susceptible and resistant reactions on the differential
wheat genotypes results in a unique digit. The wheat differential cultivars currently
used in Europe, Australia and elsewhere consist of the world set and European set.
The wheat differential genotype with resistance genes used in this study has already
been mentioned in Table 5 (Appendix II).
2.3.8 Population Structure
2.3.8.1 Virulence frequency
The virulence frequency articulates the relative amount of virulent isolates per gene.
The grouping system described by Felsenstein and Jaser (2000) was adopted to
classify the effectiveness level of the Yr resistance genes by calculating virulence
gene frequency value. They rated the resistance level as very high and high when the
virulence gene frequency percent is between 0 to 10 and more than 10 but less than
20, respectively. The term moderate and low or no resistance level was used when
38
the virulence gene frequency percent ranged between 20 to 50 and more than 50
percent, respectively.
2.3.8.2 Virulence complexity
The virulence complexity of each pathotype was obtained by the sum of its
virulences to the genes in the differential set (Welz, 1986). The information
regarding isolate's aptitude to overcome the resistance genes of the host plant is thus
generated. Above all, the complexity value interpretation depends on the size and
choice of the set of wheat differential genotypes (Löwer, 1999). Accordingly, in the
present study, 0 to 5, 6 to 12 and 13 to 24 were taken as cutting point to characterize
low, moderate and high virulence, respectively.
2.3.8.3 Pathotype abundance
The term pathotype abundance refers to the percentage of individual pathotypes in a
population.
2.3.8.4 Cluster analysis
To scrutinize the pathotypes similarity between sampling regions, cluster analysis
was performed by exploiting complete linkage technique using Statistica (1999)
Kernel release 5.5 software. A dendrogram was developed for the clustering of
isolates.
39
Chapter II
2.4 RESULTS
2.4.1 Survey for Disease Sample Collection
Survey in quest to explore stripe rust exposed area and intensity was conducted in 28
wheat cultivated locations of the Northern Punjab and NWFP (Fig. 5). The surveyed
locations were kept same, whereby 173 and 181 stripe rust samples were collected
from commercial wheat fields (farmers’ fields) and WSRTN during 2005-06 and
2006-07, respectively. Accordingly, a total of 354 collections of wheat yellow rust
samples from 8 districts under the study area, were made during the study period.
Segregation of the total figures reveals that in the Northern Punjab 20 locations were
surveyed and 87 stripe rust samples were collected while in NWFP 8 locations were
surveyed and 86 stripe rust diseased samples were collected during 2005-06 (Table
6) (Fig. 5). Similarly, in the following wheat season, 94 and 87 diseased samples
were collected from same locations of the Northern Punjab and NWFP, respectively.
2.4.2 Race Analysis
During the 2 years of research, 189 viable urediniospores were recovered out of 354
disease samples. From the recovered isolates, 150 viable isolates of P. striiformis
were tested on the differential set (Table 5). Inoculum of 90 revived samples out of
173 samples collected during 2005-06 wheat year was increased on susceptible
cultivar Morocco under glasshouse conditions. Pst inoculum of 73 isolates out of the
90 samples that expressed virulence were further taken up for virulence analysis and
race determination. Almost similar figures of 77 virulent isolates were used after
reviving 99 samples out of 181 samples collected during 2006-07 (Table 6).
39
40
Fig. 5 Districts of the Northern Punjab and NWFP Prone to Stripe Rust (Pst)
The eight districts were encircled to understand virulence pattern of Pst in
these epidemic areas based on rust occurrence and presence of pathogen's
virulence for wheat.
41
Table 6 Stripe Rust Virulence Frequency in the Northern Punjab and NWFP
during 2005-06 and 2006-07
Sr.
No. Location
Number of Isolates Virulence
Frequency (%)
Revived 05-06
Tested for Pathotype
05-06
Revived 06-07
Tested for Pathotype
06-07 05-06 06-07
Northern Punjab
1. Sialkot 8 6 12 9 38.1 50.0
2. Rawalpindi 19 15 16 13 52.8 51.6
3. Chakwal 8 6 9 7 66.7 56.3
4. Attock 7 6 13 10 38.9 56.5
NWFP
5. Nowshera 15 13 16 14 48.4 55.2
6. Charsadda 10 8 8 6 52.6 53.3
7. Peshawar 14 12 18 12 60.9 69.2
8. Mardan 9 7 7 6 69.2 41.1
Total 90 73 99 77 - -
The remaining 165 samples could not expressed virulence on Morocco either due to
loss of viability or they germinated on the diseased leaves by absorbing moisture
during transportation as the disease samples might not be dried properly before
packing.
A few of the original samples in the initial tests expressed as a mixture of different
races and accordingly, sub cultures were obtained from individual cultures, which
were then exposed to the whole differential sets. Considering both the study seasons,
42
maximum isolates originated from the District Rawalpindi (n = 28), followed by
Nowshera (n = 27), Peshawar (n = 24), Attock (n = 16), Sialkot (n = 15), and
Charsadda (n = 14). For districts Chakwal and Mardan the number of isolates
remained the same (n = 13) (Table 6).
2.4.2.1 Prevailing races of Pst
The race analysis conducted during 2006-07 identified another pathotype as 66E16
from the samples pertaining to district Attock, in addition to the already identified 12
pathotypes during 2005-06. The location with year wise prevalence of stripe rust
races are presented in Table 7 while the province wise differentiation made in
detection of Pst races is offered in Table 8. In the Northern Punjab, five (5)
pathotypes were identified in 2005-06 while seven (7) was found in 2006-07. A total
of seven (7) and ten (10) prevalent races were identified in the Northern Punjab and
NWFP, respectively during the year 2005-06 and 2006-07 (Table 8). Out of the 13
identified Pst races during the study, Pathotypes 66E0, 67E0 and 70E0 have been
reported by Kirmani (1980) while pathotypes 2E0, 6E0, 68E0 and 71E0 were
identified in stripe rust samples collected from the northern areas of Pakistan during
2003 to 2005 (Ahmad and Kazi, 2005). In addition to these seven Pst races,
Pathotypes 70E16, 66E16, 7E16, 6E17, 6E16 and 6E1 have also been identified in
the diseased samples collected from the northern areas by the CDRP, NARC during
2007 (Rattu and Fayyaz, unpublished data).
43
Table 7:Prevalence of Physiological Races in Northern Punjab & NWFP during
2005-06 and 2006-07
Sr. No.
District Physiological Races / Pathotype(s) identified during
2005-06 2006-07
Northern Punjab
1. Sialkot 70E0, 6E0 70E0, 6E0
2. Rawalpindi 70E0, 71E0, 6E0 70E0, 71E0, 6E0
3. Chakwal 2E0, 6E0 2E0, 6E0
4. Attock 70E0, 71E0, 66E0 70E0, 71E0, 70E16, 66E16
NWFP
5. Nowshera 70E0, 7E16, 6E17, 6E16, 6E1, 70E16
70E0, 7E16, 6E17, 6E16, 6E1, 70E16
6. Charsadda 66E0, 67E0, 68E0 66E0, 67E0, 68E0
7. Peshawar 67E0, 71E0, 70E0 67E0, 71E0, 70E0
8. Mardan 66E0, 70E0, 67E0, 71E0 66E0, 70E0, 67E0, 71E0
Table 8: Physiological Races identified in the Northern Punjab & NWFP during
2005-06 and 2006-07
Location Races
Northern Punjab 70E0, 6E0, 71E0, 2E0, 66E0, 70E16, 66E16
NWFP 70E0, 7E16, 6E17, 6E16, 6E1, 70E16, 66E0, 67E0, 68E0, 71E0
The response of Pst pathotypes currently characterized in multi-pathotype tests on
the world and European differential genotypes and four commercial varieties along
with a susceptible cultivar Morocco is presented in Table 9 whereas on NILs with
avocet background in Table 10. All the identified pathotypes expressed avirulent
response to the resistance genes in Vilmorin (3V), Moro (10), Strubes Dickkopf (Sd
44
and 25), Clement (2, 9, 25 and Cle), Triticum aestivum subsp. spelta var. album (5),
Reichersberg 42 (7 and 25), Heines Peko (2, 6, and 25), Nord Desprez (3N), Carstens
V (Cv), Spaldings Prolific (Sp), Heines VII (2, 25, and HVII) pertaining to the world
and European differentials (Table 9). The reduced or no IT on NILs with Yr5, Yr10,
Yr11, Yr15 and YrSp was expressed by the pathotypes identified under the study
(Appendix III).
Table 9 envisages that the response of almost all the physiological races was found
to be virulent to Yr7 (Lee) and Yr2+Yr6 (Heines Kolben) in the world set differential
and Yr6, Yr7, Yr9, Yr17, YrSk, Jupateco S (Yr18) and YrA among the NILs. The four
commercial wheat varieties and a susceptible check-Morocco that were included in
the differentials with the purpose to explore their potential for expression of
resistance or susceptibility in the glasshouse were found susceptible against all the
identified pathotypes (Table 9). The pathotypes 70E16, 66E16, 7E16, 6E17 and
6E16 exhibited virulence for Compair (Yr8,19,APR) while pathotypes 6E17 and 6E1
showed virulence for Hybrid 46 (4+) in the European differential sets (Table 9).
2.4.2.1.1 Pathotype 6E16
P. striiformis samples obtained from Nowshera in the NWFP during 2005-06 and
2006-07 were identified as pathotype 6E16 (Appendix II). The response at seedling
stage for infection type of pathotype 6E16 on the world (1 to 9) and European (10 to
17) differentials (Table 9), and a set of isogenic lines with avocet background (18 to
36) are presented in Table 10. Pathotype 6E16 was characterized due to its
45
avirulence response to the Yr resistance genes in the differential cultivars Chinese
166 (1), Vilmorin 23 (3V), Moro (10), Strubes Dickkopf (Sd and 25), Suwon
92/Omar (4 and Su), Clement (2, 9, 25, and Cle), Triticum aestivum subsp. spelta
var. album (5), Hybrid 46 (4+), Reichersberg 42 (7 and 25), Heines Peko (2, 6, and
25), Nord Desprez (3N), Carstens V (Cv), Spaldings Prolific (Sp), Heines VII (2, 25,
and HVII); and virulence to Lee (7), Heines Kolben (2 and 6) and Compair (8 and
19). Perusal of Table 10 reveals that 6E16 was found virulent to the NILs possessing
Yr resistance genes Yr6, Yr7, Yr8, Yr9, Yr12, Yr17, YrSk, and YrA.
Table 9: Response of Identified Pst Pathotypes* among Differential Genotypes
Differential set Yr genes Pathotypes identified from the samples collected during 2005-06 and 2006-07
70E0 71E0 2E0 6E0 66E0 70E16 66E16 7E16 6E17 6E16 6E1 67E0 68E0
World set Chinese 166 1 - + - - - - - + - - - + - Lee 7 + + + + + + + + + + + + - Heines Kolben 2,6 + + - + - + - + + + + - + Vilmorin 3V - - - - - - - - - - - - - Moro 10 - - - - - - - - - - - - - Strubes Dickkopf Sd,25 - - - - - - - - - - - - - Suwon92/ Omar Su + + - - + + + - - - - + + Clement 2,9,25,Cle - - - - - - - - - - - - - Triticum spelta album 5 - - - - - - - - - - - - -
European set Hybrid 46 4+ - - - - - - - - + - + - - Reichersberg 42 7,25 - - - - - - - - - - - - - Heines Peko 2,6,25 - - - - - - - - - - - - - Nord Desprez 3N - - - - - - - - - - - - - Compair 8, 19 APR - - - - - + + + + + - - - Carstens V Cv - - - - - - - - - - - - - Spaldings Prolific Sp - - - - - - - - - - - - - Heines VII 2,25,HVII - - - - - - - - - - - - -
Commercial Varieties Inquilab-91 27 + + + + + + + + + + + + + Bakhtawar-93 9+ + + + + + + + + + + + + + Wafaq-2001 9 + + + + + + + + + + + + + MH-97 + + + + + + + + + + + + + Susceptible Check Morocco - + + + + + + + + + + + + +
*Seedling response of differential lines against Pst based on ITs as described by McNeal et al. (1971)
- = Avirulent response (IT 0 - 5), + = Virulence response (IT 6-9)
46
2.4.2.1.2 Pathotype 70E0
The Pst isolates collected from Sialkot, Rawalpindi and Attock in the Northern
Punjab as well as Nowshera, Peshawar and Mardan in the NWFP during 2005-06
and 2006-07 were recognized as pathotype 70E0 (Appendix II). The infection type of
70E0 at seedling stage on the world and European differentials (Table 9), and a set of
NILs with avocet background are presented in Table 10. Characterization of
pathotype 70E0 based on the avirulence response to the Yr resistance genes in the
differential genotypes Chinese 166 (1), Vilmorin 23 (3V), Moro (10), Strubes
Dickkopf (Sd and 25), Clement (2, 9, 25, and Cle), T. aestivum spelta album (5),
Hybrid 46 (4+), Reichersberg 42 (7 and 25), Heines Peko (2, 6, and 25), Nord
Desprez (3N), Compare (8 and 19), Carstens V (Cv), Spaldings Prolific (Sp), Heines
VII ( 2, 25, and HVII); and virulence to Lee (7), Heines Kolben (2 and 6) and Suwon
92/Omar (4 and Su). The most widely distributed Pst pathotype 70E0 in the northern
region of the country, encompassed virulence to the NILs possessing Yr resistance
genes Yr6, Yr7, Yr9, Yr17, Yr18, YrSk, and YrA (Table 10).
2.4.2.1.3 Pathotype 6E0
Stripe rust samples pertaining to Sialkot, Rawalpindi and Chakwal in the Northern
Punjab while only Nowshera in the NWFP during 2005-06 and 2006-07 were found
to contain the properties of identified pathotype as 6E0 (Appendix II). The response
at glasshouse for seedling stage infection type of pathotype 6E0 on the world and
European differential lines (Table 9) and NILs with avocet background are presented
47
in Table 10. Characterization of Pst pathotype 6E0 was due to its avirulence
expression to the Yr resistance genes in the differential cultivars Chinese 166 (1),
Vilmorin 23 (3V), Moro (10), Strubes Dickkopf (Sd,25), Suwon 92/Omar (4,Su),
Clement (2,9,25,Cle), T. spelta var. album (5), Hybrid 46 (4+), Reichersberg 42
(7,25), Heines Peko (2,6,25), Nord Desprez (3N), Compare (8,19), Carstens V (Cv),
Spaldings Prolific (Sp), Heines VII (2,25,HVII); and virulence to Lee (7) and Heines
Kolben (2,6).
2.4.2.1.4 Pathotype 71E0
The Pst pathotype was identified among the samples acquired from Rawalpindi,
Attock, Peshawar and Mardan during 2005-06 and 2006-07 (Appendix II). The
infection type of 71E0 at seedling stage on the world and European differentials
(Table 9), and a set of supplemental lines with avocet background are presented in
Table 10. The pathotype 71E0 was epitomized due to its virulence response to
Chinese 166 (1), Lee (7), Heines Kolben (2 and 6) and Suwon 92/Omar (4 and Su).
The Pst pathotype responded with an avirulence reply to the remaining Yr resistance
genes in the world and European differential (Table 9).
2.4.2.1.5 Pathotype 2E0
The stripe rust pathotype 2E0 was found in the diseased leaves samples obtained
from Chakwal during the years 2005-06 and 2006-07 (Appendix II). The response at
seedling stage for infection type of pathotype 2E0 on the world (1 to 9) and
European (10 to 17) differentials (Table 9) and on the NILs with avocet background
48
(18 to 36) is mentioned in Table 10. Pathotype 2E0 was designated on the basis of its
seedling stage avirulence response to the Yr resistance genes in the world set at serial
No.1, 3, 4, 5, 6, 7, 8 and 9 and against all the genotypes in the European differential
set. The pathotype expressed virulence to Lee (Yr7) only.
2.4.2.1.6 Pathotype 70E16
Stripe rust samples collected from Nowshera during 2005-06 and 2006-07 as well as
from Attock during 2006-07 were identified as pathotype 70E16 (Appendix II). The
infection type produced by 70E16 at seedling stage on the differential sets is
mentioned in Table 9. Pathotype 70E16 was depicted due to its avirulence response
to the Yr resistance genes in the differential cultivars Chinese 166 (1), Vilmorin 23
(3V), Moro (10), Strubes Dickkopf (Sd and 25), Clement (2, 9, 25, and Cle), Triticum
aestivum subsp. spelta var. album (5), Hybrid 46 (4+), Reichersberg 42 (7 and 25),
Heines Peko (2, 6, and 25), Nord Desprez (3N), Carstens V (Cv), Spaldings Prolific
(Sp), Heines VII (2, 25, and HVII) of the world set and virulence to Lee (7), Heines
Kolben (2 and 6) and Suwon 92/Omar (4 and Su) and Compair (8 and 19) pertaining
to European set.
2.4.2.1.7 Pathotype 66E0
The Pst isolates acquired from Attock (2005-06), Charsadda and Mardan districts
during 2005-06 and 2006-07 were identified as pathotype 66E0 (Appendix II). The
infection type produced by 66E0 at the seedling stage on the world (1 to 9) and
European (10 to 17) differentials (Table 9) as well as on the set of isogenic lines with
49
avocet background (18 to 36) is presented in Table 10. The pathotype 66E0
portrayed avirulence response to the Yr resistance genes in the differential wheat
genotypes possessing Yr2,6; Yr3V; Yr10; YrSd,25; Yr2,9,25,Cle; Yr5, Yr4+; Yr7,25;
Yr2,6,25; Yr3N; Yr8,19; YrCv; YrSp; Yr2,25,HVII; and virulence reaction to Yr1;
Yr7 and Yr4,Su.
2.4.2.1.8 Pathotype 66E16
The pathotype 66E16 was identified from the disease samples grasped from Attock
during 2006-07 (Appendix II). The seedling stage infection type produced by 66E16
on the differentials sets is presented in Table 9. Pathotype 66E16 was typified and
distinguished from 66E0 due to its avirulence response to the Yr resistance genes in
the differential genotypes encompassing Yr2,6; Yr3V; Yr10; YrSd,25; Yr2,9,25,Cle;
Yr5; Yr4+; Yr7,25; Yr2,6,25; Yr3N; YrCv; YrSp and Yr2,25,HVII; and virulence to
Yr1, Yr7, Yr4,Su and Yr8,19 (Table 9). The Pst pathotype 66E16 enjoyed virulence
to the NILs possessing Yr resistance genes Yr1, Yr6, Yr7, Yr8, Yr9, Yr12, Yr17, Yr18,
Yr24, Yr26, YrSk, and YrA (Table 10).
2.4.2.1.9 Pathotype 7E16
The disease samples collected from Nowshera during both the surveyed years
acknowledged the presence of pathotype 7E16 (Appendix II). In the glasshouse,
seedling stage infection types produced by 7E16 on the world and European
genotype differentials (Table 9) as well as on the NILs with avocet background set
are presented in Table 10. The pathotype 7E16 was described due to its avirulence
50
response to the Yr resistance genes in the differential cultivars Vilmorin 23, Moro,
Strubes Dickkopf, Suwon 92/Omar, Clement, Triticum spelta, Hybrid 46,
Reichersberg 42, Heines Peko, Nord Desprez, Carstens V, Spaldings Prolific, Heines
VII. The Pst 7E16 pathotype showed virulence to Chinese 166 (Yr1), Lee (Yr7),
Heines Kolben (Yr2,6) as well as Compair (Yr8,19).
2.4.2.1.10 Pathotype 6E17
The pathotype 6E17 was identified from the disease samples acquired from
Nowshera during 2005-06 and 2006-07 (Appendix II). The infection type responses
produced at the seedling stage by pathotype 6E17 on the world and European
differentials (Table 9) as well as set of NILs with avocet background are tabulated in
Table 10. The pathotype expressed avirulence response to the Yr resistance genes in
the differential cultivars Chinese 166 (1), Vilmorin 23 (3V), Moro (10), Strubes
Dickkopf (Sd and 25), Suwon 92/Omar (4 and Su), Clement (2, 9, 25, and Cle),
Triticum spelta (5), Reichersberg 42 (7 and 25), Heines Peko (2, 6, and 25), Nord
Desprez (3N), Carstens V (Cv), Spaldings Prolific (Sp), Heines VII (2, 25, and
HVII); and virulence to Lee (7), Heines Kolben (2 and 6) and Hybrid 46 (4+) and
Compair (8 and 19).
2.4.2.1.11 Pathotype 6E1
The pathotype was designated as 6E1 on the basis of the infection type it produced
on the world and European differentials (Appendix II). The diseased leaf samples
that confirmed Pst pathotype as 6E1 were acquired from Nowshera during 2005-06
51
and 2006-07 (Table 9). The response of the identified pathotype at seedling stage
was also investigated in the glasshouse on a set of NILs with avocet background as
presented in Table 10. The pathotype 6E1 was characterized due to its avirulence
response to the Yr resistance genes in the differential cultivars Chinese 166 (1),
Vilmorin 23 (3V), Moro (10), Strubes Dickkopf (Sd and 25), Suwon 92/Omar (4 and
Su), Clement (2, 9, 25, and Cle), Triticum spelta (5), Reichersberg 42 (7 and 25),
Heines Peko (2, 6, and 25), Nord Desprez (3N), Compair (8 and 19), Carstens V
(Cv), Spaldings Prolific (Sp), Heines VII (2, 25, and HVII); and virulence to Lee (7),
Heines Kolben (2 and 6) and Hybrid 46 (4+).
2.4.2.1.12 Pathotype 67E0
The response at seedling stage for infection type produced on the world, European
differentials (Table 9) and a set of NILs with avocet background as presented in
Table 10, distinguished the pathotype as 67E0. The disease samples that expressed
presence of pathotype 67E0 were acquired from Charsadda, Peshawar and Mardan
during both the surveyed years (Appendix II). Pathotype 67E0 was characterized due
to its avirulence response to the Yr resistance genes in the differential cultivars
Heines Kolben (2 and 6), Vilmorin 23 (3V), Moro (10), Strubes Dickkopf (Sd and
25), Clement (2, 9, 25, and Cle), Triticum spelta (5), Hybrid 46 (4+), Reichersberg
42 (7 and 25), Heines Peko (2, 6, and 25), Nord Desprez (3N), Compair (8 and 19),
Carstens V (Cv), Spaldings Prolific (Sp), Heines VII (2, 25, and HVII); and virulence
to Chinese 166 (1), Lee (7) and Suwon 92/Omar (4 and Su).
52
2.4.2.1.13 Pathotype 68E0
The pathotype 68E0 was identified from the disease samples attained from
Charsadda during both the surveyed years (Appendix II). The infection type response
at seedling stage produced by 68E0 on the world and European differentials are
presented in Table 9. The pathotype was also assessed for its virulence reactions
after exposing the same on a set of NILs with avocet background (Table 10). The
pathotype 68E0 was epitomized due to its avirulence response to the Yr resistance
genes in the differential cultivars Yr1; Yr7; Yr3V; Yr10; YrSd,25; Yr2,9,25,Cle; Yr5;
Yr4+; Yr7,25; Yr2,6,25; Yr3N; Yr8,19; YrCv; YrSp; Yr2,25,HVII; and virulence to
Yr2,Yr6 and Yr4,Su.
2.4.3 Population Structure of Pst in the Northern Punjab and NWFP
2.4.3.1 Diversity of virulence
The collected 150 isolates of Pst from eight major wheat growing districts of the
Northern Punjab and NWFP were tested for pathogenecity by using 29 differential
wheat genotypes. The isolates belonged to 13 different pathotypes, which were
virulent on a differential wheat genotype possessing resistance gene Yr6, Yr7, Yr9,
Yr17, YrSk and YrA while avirulent on the 13 genotypes that carries Yr2+, Yr3V,
Yr3N, Yr5, Yr6,2+, Yr7+, Yr9,2+, Yr10, Yr11, Yr15, YrSd, YrCv and YrSp genes
(Table 10). The pathotypes differed from each other by virulence on 19 differential
genotypes carrying resistance genes Yr2, Yr3a, Yr3b, Yr4a, Yr6, Yr6+2, Yr8, Yr9,
Yr9+2, Yr10, Yr17, Yr24, Yr26, Yr27, Yr32, YrCv, YrA, YrSd, and YrSu.
53
Table 10 Reaction of Thirteen Pathotypes from 150 Isolates on the Corresponding Yr Resistance Genes Possessed by
29 Differential Genotypes
Pathotype No of Isolates
α α α α α α α α α α α α α α α α α Complexity Yr1 Yr2+ Yr3V Yr3N Yr4+ Yr5 Yr6 Yr6+2 Yr6,2+ Yr7 Yr7+ Yr8 Yr8+ Yr9 Yr9,2+ Yr10 Yr11 Yr12 Yr15 Yr17 Yr18 Yr24 Yr26 YrSd YrSu YrCv YrSp YrSk YrA
2E0 8 - - - - - - + - - + - - - + - - - - - + - - - - - - - + + 6
6E0 17 - - - - - - + + - + - - - + - - - - - + - - - - - - - + + 7
6E1 4 - - - - + - + + - + - - - + - - - + - + - - - - - - - + + 9
70E0 47 - - - - - - + + - + - - - + - - - - - + + + - - + - - + + 10
66E0 9 + - - - - - + - - + - - - + - - - - - + + + - - + - - + + 10
6E16 4 - - - - - - + + - + - + + + - - - + - + - - - - - - - + + 10
67E0 20 + - - - - - + - - + - - - + - - - - - + + + - - + - - + + 10
71E0 23 + - - - - - + + - + - - - + - - - - - + + + - - + - - + + 11
68E0 2 + - - - - - + + - + - - - + - - - - - + + + - - + - - + + 11
7E16 4 + - - - - - + + - + - + + + - - - + - + + - - - - - - + + 12
6E17 4 - - - - + - + + - + - + + + - - - + - + + - - - - - - + + 12
66E16 2 + - - - - - + - - + - + + + - - - + - + + + + - + - - + + 14
70E16 6 + - - - - - + + - + - + + + - - - + - + + + + - + - - + + 15
+ sign refers to compatible reaction (Infection Type 6-9)
– sign attributed to incompatible reaction (Infection Type 0-5) α reflect Yr resistance genes in NILs with Avocet background
53
54
2.4.3.2 Virulence frequency
An average virulence frequency of YrSk and YrA genes were close to the upper limit
of 91-99%, while that of Yr resistance genes Yr6, Yr7, Yr9, Yr17 and Jupateco S was
100% (Fig. 6, Table 11 and 12) during 2005-06. The virulence frequency of Yr18,
Jupateco R and Yr24 varied between 69 to 73 percent (Table 11). During the same
year, resistance gene Yr1 showed virulence frequency of 47 percent. Differentials
with the resistance genes Yr4+, Yr8, Yr8+, Yr12 and Yr26 showed the lowest
average virulence frequency of 4, 13, 16, 16 and 5 percent, respectively (Table 12).
The virulence frequency average of Yr6, Yr7, Yr9, Yr17, YrSk and Jupateco S gene
was 100% (Fig. 7, Table 11 and 12) and the mean virulence frequency of YrA gene
was in the upper limit as 99 percent during 2006-07. The virulence frequency of
Yr18, Jupateco R and Yr24 varied between 73 to 77 percent (Table 11). An
intermediate virulence frequency of 37 percent was once again expressed by the
resistance gene Yr1 during 2006-07.
The virulence revealed variation among the locations under the research area.
Virulence to resistance gene Yr1 was absent in district Sialkot as well as in the
isolates collected from district Chakwal. The virulence for this resistance gene was
present in isolates pertaining to the other six locations (Table 12) during the year
2005-06 and 2006-07. The population mean remained 42 percent and ranged from 18
percent in district Rawalpindi to 100 percent in district Charsadda. Virulence to
Yr18, Yr24 and YrSu was recorded in all districts except those from Chakwal.
54
55
Table 11 Virulence Frequency (%) in NILs and Isolates Pattern of Pst in the Northern Punjab and NWFP
Yr Resistance Genes
Virulent Isolates (Nos.)
Virulence Frequency (%)
Distribution in districts*
2005-06 2006-07 2005-06 2006-07 2005-06 2006-07
1 42 37 47 37 2, 4, 5, 6, 7, 8 4, 5, 6, 7, 8
5 0 0 0 0
6 90 99 100 100 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
7 90 99 100 100 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
8 19 9 21 6 2, 3, 4, 5, 8 4, 5
9 90 99 100 100 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
10 0 0 0 0
11 0 0 0 0
12 22 15 24 15 5, 6 4, 5
15 0 0 0 0
17 90 99 100 100 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
18 68 76 70 77 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 4, 5, 6, 7, 8
24 62 72 69 73 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 4, 5, 6, 7, 8
26 4 5 4 5 5 4, 5
Sp 0 0 0 0
Sk 85 99 94 100 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
Jupateco R 18 66 74 73 75 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 4, 5, 6, 7, 8
Jupateco S 90 99 100 100 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
A 87 98 97 99 1, 2, 3, 4, 5, 6, 7, 8 1, 2, 3, 4, 5, 6, 7, 8
* Refer to Fig. 5 for encircled four districts of the Northern Punjab comprising of: 1.
Sialkot including tehsils Daska, Sialkot, Pasroor and Sambrial; 2. Islamabad and
Rawalpindi consisting of tehsils Gujjar Khan, Kallar Sayedan, Kotli Satian, Murree
and Taxila; 3. Chakwal encompassing areas of Talagang, Choa Saidan Shah and
Kallar Kahar; 4. Attock with field areas of Hassan Abdal, Jand, Pindi Gheb and
Fatah Jang; and in NWFP covering four districts viz. 5. Nowshera, which also
include Nizampur tehsil; 6. Charsadda comprising of two more tehsils of Shab Qadar
and Tangi; 7. Peshawar; and 8. Mardan with areas of Takht Bhai.
56
0
10
20
30
40
50
60
70
80
90
100
Viru
lenc
e fr
eque
ncy
(%)
Yr6Yr1
7Ju
patec
o S Yr7 Yr9YrAYrS
kYr1
8 R Yr18
Yr24
Yr1Yr1
2Yr8
Yr26
Yr5Yr1
0Yr1
1Yr1
5YrS
p
Yr resistance genes
Fig. 6 Virulence Frequency of Pst in the Northern Punjab and NWFP during
2005-06
0
10
20
30
40
50
60
70
80
90
100
Viru
lenc
e fr
eque
ncy
(%)
Yr6Yr1
7
Jupa
teco
SYr7 Yr9
YrSk
YrAYr1
8
Yr18R
Yr24
Yr1Yr1
2Yr8
Yr26
Yr5Yr1
0Yr1
1Yr1
5YrS
p
Yr resistance genes
Fig. 7 Virulence Frequency of Pst in the Northern Punjab and NWFP during
2006-07
57
Table 12 Virulence Frequencies (%) of 150 Pst Isolates Collected from the Northern Punjab and NWFP during 2005-06 and 2006-07
Differential Genotype
Yr Gene
Districts Total
(n = 150) Sialkot (n = 15)
Rawalpindi (n = 28)
Chakwal (n = 13)
Attock (n = 16)
Nowshera (n = 27)
Charsadda (n = 14)
Peshawar (n = 24)
Mardan (n = 13)
Yr1/6* Avocet S Yr1 0 18 0 56 30 100 67 85 42
Heines VII Yr2+ 0 0 0 0 0 0 0 0 0
Vilmorin Yr3V 0 0 0 0 0 0 0 0 0
Nord Desprez Yr3N 0 0 0 0 0 0 0 0 0
Hybrid 46 Yr4+ 0 0 0 0 30 0 0 0 4
Yr5/6* Avocet S Yr5 0 0 0 0 0 0 0 0 0
Yr6/6* Avocet S Yr6 100 100 100 100 100 100 100 100 100
Heines Kolben Yr6+2 100 100 38 81 100 50 71 31 77
Heines Peko Yr6,2+ 0 0 0 0 0 0 0 0 0
Yr7/6* Avocet S Yr7 100 100 100 100 100 100 100 100 100
Reichersberg 42 Yr7+ 0 0 0 0 0 0 0 0 0
Yr8/6* Avocet S Yr8 0 14 0 25 41 0 0 0 13
Compair Yr8+ 0 0 0 25 59 0 0 0 16
Yr9/6* Avocet S Yr9 100 100 100 100 100 100 100 100 100
Clement Yr9,2+ 0 0 0 0 0 0 0 0 0
Yr10/6* Avocet S Yr10 0 0 0 0 0 0 0 0 0
Yr11/3* Avocet S Yr11 0 0 0 0 0 0 0 0 0
Yr12/3* Avocet S Yr12 0 0 0 25 74 0 0 0 16
Yr15/6* Avocet S Yr15 0 0 0 0 0 0 0 0 0
Yr17/6* Avocet S Yr17 100 100 100 100 100 100 100 100 100
Yr18/3* Avocet S Yr18 67 75 0 100 70 100 100 100 78
Yr24/3* Avocet S Yr24 67 75 0 100 74 100 100 100 72
Yr26/3* Avocet S Yr26 0 0 0 19 15 0 0 0 5
Strubes Dickkopf YrSd 0 0 0 0 0 0 0 0 0
Suwon92/ Omar YrSu 67 75 0 100 41 100 100 100 73
Carstens V YrCv 0 0 0 0 0 0 0 0 0
YrSp/3* Avocet S YrSp 0 0 0 0 0 0 0 0 0
Yr27/3*Avocet (CX 94.19.1.1) YrSk 100 100 100 100 100 100 100 100 100
Avocet + YrA YrA 93 100 100 100 100 100 100 100 99 Key: The ranges from 0 - 100 in white, yellow and orange shades indicate as below
0 - 20 percent 21 - 50 percent 51 - 100 percent
58
Likewise, Yr4+ virulence was not detected in the Northern Punjab and NWFP except
district Nowshera. The frequency of Yr26 (population mean of 5 percent) gene was
comparable with that of Yr4+ (population mean of 4 percent) and varied from 15
percent (Nowshera) to 19 percent (Attock). The frequency of Yr8+ and Yr12 was
equal at 16 percent but higher than Yr8 (population mean of 13 percent) as evident
from Table 12.
2.4.3.3 Virulence complexity
The pathotypes frequency and corresponding isolates distribution according to the
number of virulence genes (virulence complexity) during the year 2005-06 and 2006-
07 is represented in Table 13 and 14. During these years, the pathotypes have 6 to 15
virulences genes (Fig. 8 and 9). In both the years, highest complexity value of 15 was
contained by Pathotype 70E16. The most frequent (41 isolates) were those with 10
virulences; 12 isolates had 11 virulences while 8 isolates encompassed 7 virulences
during 2005-06 (Table 13). It is evident from Fig. 8 that none of the isolates
contained 8, 13 or 14 virulences during the year 2005-06. The minimal complexity
value of six was articulated by four isolates.
During the year 2006-07, the most recurrent (39 isolates) were those that encompass
10 virulences followed by 13 isolates with 11 virulences while 9 isolates had 7
virulences (Table 14). No isolates possessed 8 and 13 virulences during the year
2006-07 (Fig. 9). The least complexity value of six was once again expressed by four
isolates.
59
Table 13 Grouping of Pst Pathotypes Collected from the Northern Punjab and
NWFP during 2005-06 based on the Frequency of Occurrence and
Virulence Gene
No of virulence genes
6 7 9 10 11 12 15 67E0(11)
6E16(2)
66E0(6) 68E0(1) 6E17(2)
2E0(4) 6E0(8) 6E1(2) 70E0(22) 71E0(11) 7E16(2) 70E16(2)
* Isolates number are indicated in parenthesis
0
10
20
30
40
50
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Complexity value
Fre
quen
cy (
%)
Fig. 8 Complexity (virulence loci numbers in each isolate) Distribution for the
Samples Pertaining to the Northern Punjab and NWFP during 2005-06
60
Table 14 Grouping of Pst Pathotypes Collected from the Northern Punjab and
NWFP during 2006-07 based on the Frequency of Occurrence and
Virulence Gene
No of virulence genes
6 7 9 10 11 12 14 15 67E0(9)
6E16(2)
66E0(3) 68E0(1) 6E17(2)
2E0(4) 6E0(9) 6E1(2) 70E0(25) 71E0(12) 7E16(2) 66E16(2) 70E16(4)
* Isolates number are indicated in parenthesis
0
10
20
30
40
50
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Complexity value
Fre
que
ncy
(%)
Fig. 9 Complexity (virulence loci numbers in each isolate) Distribution for the
Samples Pertaining to the Northern Punjab and NWFP during 2006-07
61
2.4.3.4 Pathogen abundance
A total of 73 isolates with a division of 6, 15, 6, 6, 13, 8, 12 and 7 isolates pertaining
to Sialkot, Rawalpindi, Chakwal, Attock, Nowshera, Charsadda, Peshawar and
Mardan were probed and grouped into 2, 3, 2, 3, 6, 3, 3 and 4 distinct pathotypes,
respectively during 2005 - 06 (Table 15). The pathotype abundance in each district
was calculated by dividing number of isolates of a particular pathotype with the total
number of isolates found in that particular district. The most abundant pathotype in
the Sialkot, Rawalpindi, Attock, Nowshera and Peshawar districts was 70E0
representing 66.7 (4/6), 40 (6/15), 66.7 (4/6), 23.1 (3/13) and 33.3 (4/12) percent of
the total isolates in that districts. In Chakwal district pathotype 2E0 represented 66.7
(4/6) percent of the isolates. It is evident from Table 15 that the most abundant
pathotype 67E0 in Charsadda and Mardan districts accounted for 50 (4/8) and 42.9
(3/7) percent of the total isolates in their respective districts. On the whole, pathotype
70E0, virulent to Yr6, Yr6+2, Yr7, Yr9, Yr17, Yr18, Yr24, YrSu, YrSk and YrA (Table
10) was the most dominant pathotype identified in six districts and accounted for
30.14 (22/73) percent of the total pathotype population (Table 15). The five most
abundant pathotypes identified during 2005-06 were 6E0, 70E0, 71E0, 66E0 and
67E0 comprised 79.45 percent of the total samples.
A total of 77 isolates with a division of 9, 13, 7, 10, 14, 6, 12 and 6 isolates
pertaining to Sialkot, Rawalpindi, Chakwal, Attock, Nowshera, Charsadda, Peshawar
and Mardan were investigated and classified into 2, 3, 2, 4, 6, 3, 3 and 4 distinct
62
Table 15 Numbers of Prevailing Pathotypes found in the Northern Punjab and
NWFP during 2005-06
Pathotypes Northern Punjab North West Frontier Province
Total Percent Sialkot Rawalpindi Chakwal Attock Nowshera Charsadda Peshawar Mardan
6E0 2 4 2 8 10.95 70E0 4 6 4 3 4 1 22 30.14
71E0 5 1 4 1 11 15.07
2E0 4 4 5.48
70E16 2 2 2.74
66E0 1 3 2 6 8.23
66E16 0 0
7E16 2 2 2.74
6E17 2 2 2.74
6E16 2 2 2.74
6E1 2 2 2.74
68E0 1 1 1.37
67E0 4 4 3 11 15.06
2/6 α 3/15 2/6 3/6 6/13 3/8 3/12 4/7 12/73 100 α indicate numbers of pathotypes / isolates
Table 16 Numbers of Prevailing Pathotypes found in the Northern Punjab and
NWFP during 2006-07
Pathotypes Northern Punjab North West Frontier Province
Total Percent Sialkot Rawalpindi Chakwal Attock Nowshera Charsadda Peshawar Mardan
6E0 3 3 3 9 11.69
70E0 6 5 3 4 6 1 25 32.47
71E0 5 3 3 1 12 15.58
2E0 4 4 5.19
70E16 2 2 4 5.19
66E0 2 1 3 3.90
66E16 2 2 2.60
7E16 2 2 2.60
6E17 2 2 2.60
6E16 2 2 2.60
6E1 2 2 2.60
68E0 1 1 1.30
67E0 3 3 3 9 11.68
2/9 α 3/13 2/7 4/10 6/14 3/6 3/12 4/6 13/77 100
α indicate numbers of pathotypes / isolates
63
pathotypes, respectively during 2006-07 (Table 16). During the same year, the most
abundant pathotype in the Sialkot, Rawalpindi, Attock, Nowshera and Peshawar
districts was 70E0 representing 66.7 (6/9), 38.5 (5/13), 30 (3/10), 28.6 (4/14) and 50
(6/12) percent of the total isolates in these districts. In Chakwal district, pathotype
2E0 represented 57.1 (4/7) percent of the total isolates.
The most abundant pathotype 67E0 accounted for 50 percent of the total isolates in
Charsadda and Mardan districts (Table 16). On the whole, pathotype 71E0, virulent
to Yr1, Yr6, Yr6+2, Yr7, Yr9, Yr17, Yr18, Yr24, YrSu, YrSk and YrA with virulence
complexity of 11 (Table 10) was the second most dominant pathotype after
pathotype 70E0 that was identified in four districts and accounted for 15.58 percent
of the total pathotype population (Table 16). The four most abundant pathotypes
identified during 2006-07 were 6E0, 70E0, 71E0 and 67E0 comprising 71.42 percent
of the total samples.
2.4.3.5 Cluster analysis
A dendrogram was constructed on similarity matrices basis by inserting pathotypes
differentiation data with respect to their effect on Yr resistance genes (Fig. 10). The
group similarity was then observed among isolates on the basis of unweighted pair-
group average percent disagreement: 2E0, 6E0, 6E1 and 6E16 (Fig. 10). Likewise,
isolates 71E0, 67E0 and 66E0 were similar with 68E0 and 70E0 at a linkage distance
of 0.06 (Table 17). Pathotypes 66E16, 71E0 and 70E0 from the Northern Punjab had
pair similarity with 70E16, 67E0 and 68E0, respectively which were identified from
64
Fig. 10 Tree Diagram for 13 Pathotypes Collected from the Northern Punjab
and NWFP during 2005-06 and 2006-07
Table 17 Percent Disagreement among the Pst Pathotypes Identified from the
Samples Collected during 2005-06 and 2006-07
Pathotypes 70E0 71E0 2E0 6E0 66E0 70E16 66E16 7E16 6E17 6E16 6E1 67E0 68E0
70E0 0.00
71E0 0.06 0.00
2E0 0.14 0.19 0.00
6E0 0.11 0.17 0.03 0.00
66E0 0.06 0.06 0.14 0.17 0.00
70E16 0.14 0.14 0.28 0.25 0.14 0.00
66E16 0.17 0.17 0.25 0.28 0.11 0.03 0.00
7E16 0.19 0.14 0.22 0.19 0.19 0.11 0.14 0.00
6E17 0.17 0.22 0.19 0.17 0.22 0.14 0.17 0.08 0.00
6E16 0.19 0.25 0.11 0.08 0.25 0.17 0.19 0.11 0.08 0.00
6E1 0.17 0.22 0.08 0.06 0.22 0.25 0.28 0.19 0.11 0.08 0.00
67E0 0.08 0.03 0.17 0.19 0.03 0.17 0.14 0.17 0.25 0.28 0.25 0.00
68E0 0.06 0.06 0.19 0.17 0.06 0.14 0.17 0.19 0.22 0.25 0.22 0.08 0.00
65
the isolates of NWFP. Four pathotypes from the Northern Punjab and six pathotypes
pertaining to NWFP had pair similarity. For instance isolates, 6E0 and 2E0, 66E16
and 70E16, as well as 6E17 and 7E16, 67E0 and 71E0, 68E0 and 70E0 were very
similar. Although, the pathotypes 66E16 and 70E16 formed a similar group with
pathotype 6E17 and 7E16 but a high linkage distance of 0.14 was observed (Table
17).
2.4.4 Wheat Stripe Rust Trap Nurseries
The present study evaluated the expression of rust resistance at seedling stage in the
glass house as well as appraised the occurrence and virulence of stripe rust in field
through establishing trap nurseries (WSRTN) at hot spots (Table 18). The near
isogenic lines with avocet background as Yr6, Yr7, Yr8, Yr17, Jupateco S, Jupateco
R, YrA (except at Sialkot) and Yr9 expressed high susceptibility at the WSRTN
locations. Similarly, the resistance gene, Yr1 showed moderate to high susceptible
response. Virulence factors for Yr1, Yr2, Yr6, Yr7, Yr9, Yr17, YrA and Yr27 have also
been reported at all trap nurseries (Rattu et al., 2007). The stripe rust resistance gene,
Yr24 was the only entry in the NILs that responded with MRMS type reaction
ranging from no reaction to 10MRMS during 2005-06 and 2006-07 (Appendix I).
Rattu et al. (2007) earlier reported that Yr24 reflected moderately resistant to
moderately susceptible reaction at Islamabad and Pirsabak and their results coincide
with the findings of this study as Yr24 expressed MRMS type reaction at PMAS-
AAU, Rawalpindi in addition to NARC and CCRI; while showed immune reaction at
66
Table 18 Virulence Analysis through Stripe Rust Reactions in NILs with
Avocet Background at WSRTN Sites of the Northern Punjab &
NWFP during 2005-06 & 2006-07*
Lines/Varieties Yr
Resistance
genes
Stripe Rust Reaction to Yr at Trap nurseries established at
PRS (Sialkot) PMAS-AAUR
(Rawalpindi)
NARC
(Islamabad)
CCRI
(Pirsabak)
AUP
(Peshawar)
NIFA
(Peshawar)
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07
Yr1/6* Avocet(s) 1 -* -* + + + + + + + + + +
Yr5/6* Avocet(s) 5 - - - - - - - - - - - -
Yr6/6* Avocet(s) 6 + + + + + + + + + + + +
Yr7/6* Avocet(s) 7 + + + + + + + + + + + +
Yr8/6* Avocet(s) 8 - - + + + + + + + + + +
Yr9/6* Avocet(s) 9 + + + + + + + + + + + +
Yr10/6* Avocet(s) 10 - - - - - - - - - - - -
Yr15/6* Avocet(s) 15 - - - - - - - - - - - -
Yr17/6* Avocet(s) 17 -* + + + + + + + + + + +
Yr18/3* Avocet(s) 18 -* -* + + + + + + + + + +
Yr24/3* Avocet(s) 24 - - -* -* -* -* -* -* - - - -
Yr26/3* Avocet(s) 26 - - - - - - - - - - - -
YrSP/3* Avocet(s) Sp - - - - - - - - - - - -
Yr27/3*Avocet
(CX 94.19.1.1) Sk - - + + + + + + + + + +
Jupateco R (Yr18) 18 - - + + + + + + + + + +
Jupateco S - - - + + + + + + + + + +
AVOCET + YrA A + + + + + + + + + + + +
*Field response of NILs based on disease severity against Pst is presented in Appendix I.
+ = Presence of virulence, - = Absence of virulence, -* = Virulence present but not
effective to overcome the gene at specified location. The presence or absence of the
virulences was calculated on the basis of coefficient of infection by multiplying the
response value given in Table 3 with the percent of rust infection as per modified
Cobb’s scale (Fig. 2).
67
PRS, Sialkot; AUP; and NIFA at Peshawar during both the years. The Yr26
expressed intermediate to moderate reactions as 5RMR at NARC, Islamabad (2006-
07), 10RMR at CCRI, Pirsabak and NIFA, Peshawar; Yr9, Yr2+ presented 10-
50MRMS type reactions at PMAS-AAUR, NARC, CCRI, AUP, and NIFA during
both the years. The NILs containing resistance gene Yr26 expressed resistance
during 2005 to 2007 wheat seasons in the WSRTN sites at PRS, PMAS-AAUR, and
AUP. Stripe rust was not observed at any WRSTN sites in the NILs with avocet
background having resistance genes Yr5, Yr10, Yr15 and YrSp.
Virulence response as tabulated in Table 18 was witnessed for stripe rust resistance
genes Yr6, Yr7, Yr9 and YrA in all WSRTN sites at AAUR, NARC, PRS, AUP,
NIFA and CCRI during the two years of study. Partial virulence was observed for Yr
resistance genes Yr1 and Yr18 during 2005-06 and 2006-07, while for Yr17 during
2005-06 at Sialkot. Likewise, virulence was present during these years, for Yr24 at
Rawalpindi, Islamabad and Pirsabak but was not effective to trounce the resistance
gene. Avirulence to Pst was observed in the study area for Yr resistance genes Yr5,
Yr10, Yr15, Yr26 and YrSp and also for Yr24 at PRS, Sialkot; AUP and NIFA,
Peshawar during both the years. It is evident from Table 18 that virulence for stripe
rust resistance genes Yr1, Yr8, Yr17, Yr18 and YrSk was prevalent at the WSRTN
sites Rawalpindi, Islamabad, Pirsabak and Peshawar during 2005-06 and 2006-07.
The virulence factor for Yr8 is present in Islamabad, Fatehjang and Pirsabak while
that of Yr18 is prevalent in Islamabad, FatehJang, Pirsabak, and Peshawar (Rattu et
68
al., 2007). The previous study on stripe rust resistance in Pakistan also demonstrates
that stripe rust development in natural field condition was satisfactory during 1999-
2000 whereby, 5-90S yellow rust response was observed for Yr1, Yr6, Yr6+7, Yr7,
Yr9 and YrA at CCRI Pirsabak as well as for Yr1, Yr6, Yr7 and Yr9 at PRS, Sialkot
(Anonymous, 2000). The results reported by Afshari et al. (2002) after testing 123
lines for identification of stripe rust resistance genes also confirms virulence for Yr6,
Yr7, and Yr6+7. During the present study, it was also noted that the trap nursery at
Pulses Research Station, Sialkot expressed immune reaction for all Yr genes except
for Yr1, Yr6, Yr7, Yr9, Yr17, Yr18 and YrA.
The disease severity ranged from 50 - 100S on Morocco at all the WSRTNs, which
is an indication that the disease level in the field was high enough to measure the
resistance in differential sets and the commercial wheat varieties. The adult plant
resistance genes identified in the present study were Yr5, Yr10, Yr15, Yr24, Yr26 and
YrSp as well as the Yr resistance genes present in all the European differentials.
Since virulence factor for these genes have not been observed, therefore, use of these
effective resistance genes for their incorporation in the national wheat breeding
program has also been recommended by Rattu et al. (2007) in addition to Tatara that
possesses Yr3 and YrCv resistance gene.
2.4.5 Field Response of Commercial Wheat Varieties to Pst
Stripe rust response of 58 wheat cultivars was explored by inducting them in the
established WSRTN (Table 19) at the six stripe rust hot spots during 2005-06 and
69
2006-07. The commercial varieties, Soorab-96 (barley), Tatara (Yr3) and GA-2002
were the only three that exhibited resistant response to stripe rust at all the WSTRN
sites during these two experimentation years (Appendix V). The resistant expression
of Soorab-96 and GA-2002 is in line with the findings of Rattu et al. (2007).
There were also five cultivars viz-a-viz Pavon-76, Kohsar-93, Fakhr-e-Sarhad, Iqbal-
2000 and Durum-97, which showed a combination of resistance and partial
resistance responses at the six locations (Table 19) (Appendix V). An unnamed slow
rusting gene in combination with Lr46 is responsible for the Pavon-76 slow rusting
resistance (Singh et al., 1998), which is currently believed to possess durable
resistance (Singh and Rajaram, 1994).
None of the commercial varieties showed reaction to stripe rust in the WSRTN
established at Sialkot during 2005-06 and 2006-07 except Kohistan-97 that offered
resistant reaction (Appendix V). Morocco was the only cultivar at WSRTN Sialkot
that proved susceptible reaction with 50 and 60 percent disease severity as per cobb's
scale during 2005-06 and 2006-07, respectively. Among the commercial wheat
cultivars incorporated in the trap nurseries, MH-97, Inquilab-91, Sindh-81, Zargoon,
Faisalabad-83, Faisalabad-85, Kaghan-93, Kirin-95, Kohinoor-83, LU-26,
Nowshera-96, Punjab-96, Sariab-92, Sarsabz, Tandojam-83, SH-2002, Pak-81,
Bahawalpur-97, Rothas-90, Suleman-96, WL-711, Zardana, Abadgar-93, Watan-94,
Moomal-2002 and Margalla-99 exhibited susceptible reactions at all locations except
at Sialkot (Table 19). Contrarily, Kohistan-97, Punjab-85, Bakhtawar-93, Blue Silver
70
Table 19 Stripe Rust Reaction of Commercial Varieties to Pst at Hot Spots of the
Northern Punjab and NWFP during 2005-06 & 2006-07
Sr. No
Differential Sets Yr Genes
Disease Reaction to Yr at Trap nurseries established at
PRS (Sialkot) PMAS-AAU Rwp NARC (I.abad) CCRI (Pirsabak) AUP (Peshawar) NIFA (Peshawar)
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 1 Morocco - - - - - - - - - - - - 2 Pavon 76 Yr29 APR + + + + ** ** + + + + + + 3 MH 97 + + - - - - - - - - - - 4 Inquilab-91 Yr27 + + - - - - - - - - - - 5 Kohistan 97 + + - - - - - - + + - - 6 Punjab 85 + + + ** ** - - - - - - - 7 Bakhtawar 93 Yr9+ + + - - - - - - + + - - 8 Blue Silver Yr6+A + + - - - ? - - + ** - - 9 Chakwal 86 + + - - - - - - + + - - 10 Sindh-81 + + - - - - - - - - - - 11 Zarghoon Yr6+ + + - - - - - - - - - - 12 Faisalabad 83 Yr7+2 APR + + - - - - - - - - - - 13 Faisalabad 85 Yr9+Yr4 + + - - - - - - - - - - 14 Kaghan 93 + + - - - - - - - - - - 15 Kirin 95 + + - - - - - - - - - - 16 Kohinoor 83 + + - - - - - - - - - - 17 LU-26 Yr6 + + - - - - - - - - - - 18 Nowshera 96 + + - - - - - - - - - - 19 Parwaz 94 Yr6+Yr7 + + - - - - - - + + - - 20 Pasban 90 + + - - - - - - + + - - 21 Pirsabak 85 + + - - - - - - + + - - 22 Punjab 96 + + - - - - - - - - - - 23 Sariab-92 Yr6+ + + - - - - - - - - - - 24 Sarsabz Yr7 + + - - - - - - - - - - 25 Shaheen 94 + + - - - - - - + - - - 26 Shahkar 95 Yr6+ + + ** - - - - - + ** - - 27 Soughat 90 Yr6+Yr7 + + - - - - - - ** ** - - 28 Tandojam 83 Yr6+ + + - - - - - - - - - - 29 SH-2002 + + - - - - - - - - - - 30 Pak 81 Yr9+Yr7 + + - - - - - - - - - - 31 Bahawalpur-97 + + - - - - - - - - - - 32 Kohsar 93 + + + + ** + ** ** + + ** ** 33 Rohtas 90 + + - - - - - - - - - - 34 Suleman 96 + + - - - - - - - - - - 35 WL 711 Yr2 + + - - - - - - - - - - 36 Zardana Yr7 + + - - - - - - - - - - 37 Abadgar 93 + + - - - - - - - - - - 38 Anmol-91 Yr9 + + - - - - - - + - + + 39 Bahawalpur-2000 + + - - - - - - + + - - 40 Bahkhar-2002 + + - - - - + + + - + + 41 Fakhr-e-Sarhad + + + + + + ** ** ** + ** ** 42 Marvi-2000 + + + + + ** - - - - - - 43 Mehran-89 Yr9 + + - - - - + - + ** + + 44 Soorab-96 (Barley) + + + + + + + + + + + + 45 Tatara Yr3 + + + + + + + + + + + + 46 Takbeer + + - - + - + ** + - + - 47 AS-2002 + + - - + - + - + ** + + 48 Iqbal 2000 Yr9 + + + + + + ** ** + + + + 49 Auqab-2000 Yr9 + + ** ** ** ** + + ** - - - 50 Chakwal-97 + + - - - - - - + ** ** ** 51 Durum-97 + + + ** ** + + + + + + + 52 Watan 94 + + - - - - - - - - - - 53 Moomal 2002 + + - - - - - - - - - - 54 Zarlashta Yr9 + + - - - - - - + - + + 55 GA-2002 + + + + + + + + + + + + 56 Wafaq-01 Yr9 + + - - - - - - + - - - 57 Margalla-99 + + - - - - - - - - - - 58 Manthar-3 Yr9 + + - - - - - - + + - -
+ = Resistant, - = Susceptible, ** = Partial Resistance, ? = Missed entry
71
and Chakwal-86 bestowed mix response from resistant to partial resistant and even
susceptible reactions at the trap nurseries during 2005-06 and 2006-07. Chakwal 86
was the single variety that presented MRMS type reaction at all the WRSTN sites
during these years.
Table 19 expose eight wheat commercial varieties viz. Kohistan-97, Bakhtawar-93,
Chakwal-86, Parwaz-94, Pasban-90, Pirsabak-85, Bahawalpur-2000 and Manthar-3
as completely resistant to stripe rust only at the WSRTN established at Peshawar
(AUP) and Sialkot (PRS) during 2005-06 and 2006-07.
Field WSRTN data illustrated that majority of the Yr resistance genes were
comparatively virulent at all WRSTN locations and expressed 5–100S yellow rust
reaction in Morocco, Inquilab-91 (Yr27), Bakhtawar-93 (Yr9+), Wafaq-01 (Yr9) and
MH-97 during both the years. Inquilab-91, Bakhtawar-93, Wafaq-01 and MH-97
illustrated immune reactions at Sialkot, however, Bakhtawar-93 (Yr9+) exhibited 20-
30MS at AAUR, 30MRMS-40S at NARC, 20S at CCRI and 50MSS-50S at NIFA
during 2005-06 and 2006-07, respectively. No susceptibility for this variety was
observed during these years at AUP. Wafaq-01 showed 20S at UAAR and NARC,
while 40MS at CCRI and 30S at NIFA while MH-97 revealed 30-80S response at the
WRSTN sites during 2005-06 and 2006-07 (Appendix V).
The variety Blue Silver (Yr6+YrA), which demonstrated susceptibility at four
WSTRN sites, expressed resistance at Peshawar (AUP) during 2005-06 while
showed partial resistance in 2006-07. Likewise, Soghat-90 (Yr6+Yr7) offered
72
susceptibility at all the trap nurseries excluding Sialkot but remained partially
resistant at Peshawar (AUP) during 2005-06 and 2006-07. Mehran-89 (Yr9) and AS-
2002 showed all types of reactions including resistance, partial as well as susceptible
responses at various locations during the experimentation years.
Table 20 pertaining to glasshouse virulence analysis of the isolates based on
infection types (ITs) showed that the varieties, Morocco and Inquilab-91 were the
most susceptible cultivars (IT 7 to 9), followed by Wafaq-01, MH-97 and
Bakhtawar-93 (IT 6 to 9).
Table 20: Stripe Rust Reactions at Seedling Stage on Commercial Varieties at CDRP, Sunny
Bank, Murree during 2005-06 & 2006-07
Sr.
No.
Varieties/
Cultivars
Yr
genes
Stripe Rust Reaction collected samples from WSRTN sites
Sialkot Rawalpindi Islamabad Pirsabak Nowshera Peshawar
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07
1 Morocco - 8 9 8 9 8 9 8 9 8 7 7 8 8 9 7 7 7 8 8 9
2 Inquilab-91 Yr27 7 8 7 8 8 9 8 9 8 7 7 8 8 9 8 6 7 8 9 8 9
3 Bakhtawar-93 Yr9+ 6 7 7 6 6 7 7 8 67 7 8 7 8 8 7 8 6 7
4 Wafaq-01 Yr9 7 8 7 8 9 7 8 6 7 6 7 8 8 9 7 8 6 7 8 7 8
5 MH-97 6 7 6 7 6 7 8 6 7 7 8 7 8 6 7 7 8 7 8 6 7
2.4.6 Field Response of Breeders Advance Lines to Pst
The results revealed that the intensity of disease infection during the year 2006-07
was higher as compared to the year 2005-06. This severity may be attributed to the
relative dry weather which prevailed during the year 2005-06. More rainfalls during
2006-07 favoured the intensity of disease in almost all varieties/lines. The results are
in line with the work done by TeBest et al. (2008) who reported that intensity of
73
C_7
3C
_65
C_2
0C
_15
C_1
88C
_187
C_1
86C
_185
C_1
84C
_183
C_1
82C
_179
C_1
78C
_177
C_1
76C
_175
C_1
74C
_173
C_1
72C
_170
C_1
68C
_167
C_1
66C
_165
C_1
64C
_163
C_1
62C
_160
C_1
59C
_158
C_1
57C
_156
C_1
55C
_154
C_1
53C
_152
C_1
50C
_149
C_1
51C
_148
C_1
47C
_146
C_1
45C
_144
C_1
41C
_180
C_1
40C
_139
C_1
38C
_137
C_1
36C
_135
C_1
34C
_132
C_1
31C
_130
C_1
29C
_128
C_1
27C
_126
C_1
25C
_124
C_1
23C
_122
C_1
21C
_120
C_1
19C
_118
C_1
17C
_116
C_1
15C
_114
C_1
13C
_112
C_1
11C
_110
C_1
09C
_108
C_1
07C
_106
C_1
05C
_104
C_1
03C
_102
C_1
01C
_100
C_9
9C
_98
C_9
5C
_94
C_9
3C
_92
C_8
9C
_88
C_8
5C
_84
C_9
0C
_83
C_8
2C
_171
C_9
1C
_81
C_7
9C
_78
C_7
7C
_75
C_1
69C
_74
C_7
2C
_142
C_7
1C
_143
C_8
6C
_70
C_6
9C
_68
C_6
7C
_66
C_6
4C
_63
C_6
2C
_60
C_5
9C
_58
C_5
7C
_56
C_5
5C
_54
C_5
3C
_52
C_5
1C
_50
C_4
9C
_47
C_4
6C
_43
C_4
2C
_133
C_8
7C
_61
C_4
1C
_39
C_3
8C
_37
C_3
5C
_34
C_3
3C
_32
C_3
1C
_181
C_7
6C
_26
C_2
4C
_23
C_2
2C
_28
C_2
7C
_30
C_2
9C
_11
C_9
6C
_48
C_2
1C
_19
C_2
5C
_18
C_1
7C
_16
C_4
5C
_44
C_1
4C
_97
C_3
6C
_13
C_1
2C
_10
C_9
C_8
C_1
61C
_80
C_4
0C
_7C
_6C
_5C
_4C
_3C
_2C
_1
0
20
40
60
80
100
120
(Dlin
k/D
max
)*10
0
Fig. 11 Cluster Analysis of 188 Commercial and Breeders' Lines based on
Disease Resistance Traits
74
stripe rust is favored by a model with temperature, humidity, and rainfall. It is
evident from (Appendix VI) that out of 188 varieties and lines, 77 showed zero
average coefficient of infection with highest Relative Resistance Index (RRI) valuing
9, 70 exhibited RRI values ranging 7.2 to 8.94, 26 cultivars showed RRI in the range
of 5.85 to 6.97 whereas remaining represented values of RRI below 5.
It was, therefore, concluded that most of the lines had a great potential to be used as
a source of resistant germplasm against the stripe rust disease. However, we should
not rely upon Inquilab-91 because it has shown greater susceptibility than the
previous years. The results are in line with those of Singh et al. (2004b) who
reported that Yr27 gene present in Inquilab-91 in Pakistan and PBW-343 in India has
broken down resulting in severe economic losses (Kisana et al., 2003). The data was
later on analyzed for cluster analysis. It is evident from the dendrogram (Fig. 11) that
the entire varieties could be divided into two clusters at 70 percent linkage distance,
wherein one cluster comprised only four cultivars whereas the remaining 184
varieties pertains to an other cluster. However, there was a split in this big cluster at
60 percent linkage distance where in we observed 4, 95 and 89 lines in cluster I, II
and III, respectively. A major break through was observed at 20 percent linkage
distance wherein the group of 4 was segregated and we observed 6 clusters vis-à-vis
Cluster I, II, III, IV, V and VI each comprising 3, 1, 57, 38, 61 and 28 lines
respectively (Fig. 11). It was observed that the cultivars generally grouped in clusters
according to their potential of resistance against the disease. The most susceptible
were grouped under cluster III (60 percent linkage distance). But under this cluster
75
these showed further split. It was amazing to note that the lines 124/2 and 101/25
falling under acceptable range grouped with lines 106/20 and 116/10 higher RRI
value. Likewise majority of the lines with highest RRI value of 9 fell under cluster-II
except the lines PASTOR/3/ALTAR 84/AEGILOPS SQUARROSA, TOB/ERA/
TOB/CNO67/3/PLO/4/VEE#5/5/KUAZ/6/, DHARWAR DRY/ NESSER, CHEN/
AEGILOPS/SQUARROSA(TAUS)//BCN/3/BAV92, CNDO/R143/ENTE/MEXI_2/
3/, ATTILA *2//CHIL/BUC were represented under cluster III at 60 percent linkage
distance. Such uneven distribution of cultivars containing lower RRI values and their
grouping with those having highest RRI values may be attributed to their
performance under moisture stress conditions of the arid zone. Our results are in
conformity with those of Ali et al. (2008) who observed that most of the lines exhibit
more resistance under high disease pressure as compared to the susceptible check.
Such lines may be exploited to explore and utilized for their resistance gene to
minimize damages caused by strip rust. We may, therefore, conclude that out of 188
cultivars 150 had RRI value >7≤9 and were in desirable range. Whereas, 28 cultivars
were placed among the acceptable range having RRI value ≥5 < 7 and only 10
cultivars showed RRI value < 5 and fell under undesirable range. Thus still there is a
great potential to exploit the existing sources of wheat germplasm to develop stripe
rust resistant varieties. For this purpose, process may be continued with more lines
and at more locations for determination of resistance against yellow rust as well as
for stability of the germplasm over years along with other desirable characteristics
before final approval of the variety.
76
Chapter II
2.5 DISCUSSION
2.5.1 Population Structure of Pst
The Pst pathotypes encompassing virulence to six or more of the following known
stripe rust resistance genes were identified through this study: Yr1, Yr4+, Y6, Yr7,
Yr8, Yr8+, Yr9, Yr12, Yr17, Yr18, Yr24, Yr26, YrSu, YrSk and YrA. The results
pertaining to presence of virulences in the study area are in agreement with the
yellow rust trap nurseries data reported by Mirza et al. (2004) about the presence of
virulence for the genes Yr1, Yr2, Yr6, Yr7 and Yr9 in Pakistan during 1998-99. In
addition to these, prevalence of virulence for genes Yr4, Yr8 and Yr18 has also been
documented by Rizwan et al. (2007) after collecting Yr isolates from the diseased
samples pertaining to environmentally diverse regions of the northern Pakistan
during 2003-04 and 2004-05. Presence of virulence for resistance genes Yr3, Yr5,
YrSp, YrSd and YrCv in the northern areas as reported by Rizwan et al. (2007) was,
however, not identified among the isolates collected from the study area of the
Northern Punjab and NWFP during 2005-06 and 2006-07. The virulence to Yr
resistance genes Yr6,2+, Yr7+, Yr9,2+, Yr10, Yr11, and Yr15 was also not detected
among the isolates collected from the Northern Punjab and NWFP during 2005-06
and 2006-07.
76
77
The isolates possessing virulence to Yr1, Yr4+, Y6, Yr7, Yr8, Yr9, Yr17, Yr18, Yr24,
and YrA identified in this study were not unique to Northern areas of Pakistan as
virulence to Yr6, Yr7 and Yr9 is there in most wheat producing regions of the world
(Chen et al., 2002). Virulence has also been accounted to stripe rust genes Yr2, Yr3,
Yr4, Yr6, Yr7, Yr8, Yr9, Yr17, YrSd, YrSu and YrCv in Germany (Flath and Bartels,
2002). Virulence near 100 percent was observed to Yr1 in a similar study conducted
in Europe (Flath and Bartels, 2002; Wozniak-Strzembicka, 2003).
The stripe rust resistance genes Yr2+, Yr3V, Yr3N, Yr5, Yr6,2+, Yr7+, Yr9,2+, Yr10,
Yr11, Yr15, YrSd, YrCv and YrSp were found effectual against the isolates collected
from the wheat cultivated areas of the Northern Punjab and NWFP. The findings
were in accordance with the reports of Chen (2005), Afshari (2008) and Chunmei et
al. (2008) that virulences to genes Yr5 and Yr15 occur as a rare phenomenon in
wheat growing regions of the globe. The most extensively stripe rust resistance gene
to wheat illustrated so far is Yr5 that occur in T. spelta album, located on
chromosome 2BL (Macer, 1966) and has expressed high degree of resistance against
all or most of the Yr isolates in North America, Turkey and China (Macer, 1966;
Wang et al., 1996; Zeybeck and Yigit, 2004; Chen, 2005).
Combinations or use of these resistance genes in alone may facilitate to trim down
yield losses associated with stripe rust in Pakistan. The gene pyramiding tactic,
22however, has an edge to develop commercial wheat varieties possessing durable
resistance (Jacobs and Parlevliet, 1993). Yr genes Yr8,19,APR, Yr18 and Yr24
78
(Appendix I, II and III) offer moderate resistance which can also be deemed for gene
pyramiding in future breeding program. Their minimum protective efficiency to the
prevailing stripe rust population, however, necessitates exploring novel resistance
sources.
Alteration in the virulences of the stripe rust population can occur either due to
mutations, migrations, re-combinations or direct selections. High similarity between
pathotypes was observed when classification and grouping of pathotypes, based on
the presence of number of virulence genes in each isolate was made (Tables 10, 13
and 14). The pathotype with 15 virulences (70E16) thus differs from the pathotype
with 14 virulences (66E16) by the presence of virulence to gene Yr6+2 (Table 10).
Likewise, pathotype with 14 virulences (66E16) differs from the pathotype with 12
virulences (6E17) by the presence of the Yr1, Yr24, Yr26 and YrSu virulences, while
absence of Yr4+ and Yr6+2 virulences. The number of virulences (12) in 7E16 is
increased after subtraction of virulence to Yr24 and YrSu but addition of virulences
to Yr8, Yr8+ and Yr12 in to pathotype 68E0 and 71E0 with 11 virulences. Similarly
the most frequent pathotype 70E0 in the major-group with 10 virulence genes differs
from pathotype 6E1 with 9 virulences by absence of virulence to Yr4+ and Yr12 but
having additional virulence to Yr18, Yr24 and YrSu. This pathotype differ from
pathotype 71E0 with 11 virulences by lacking virulence to Yr1. This single Yr gene
difference among the pathotypes might defend a single step mutation pattern.
79
A significant difference in the pathotypes composition pertaining to various regions
was observed (Table 15 and 16). Only four and five out of the 13 pathotypes were
detected in three or more locations during the year 2005-06 and 2006-07,
respectively. The most frequent pathotype 2E0 pertaining to Chakwal was not
identified at any other district. Similarly the four most common pathotypes in the
Nowshera district (7E16, 6E17, 6E16 and 6E1) were not detected in any of the other
seven districts during both the surveyed years (Table 15 and 16). Likewise 68E0 was
identified only at Charsadda during 2005-06 and 2006-07. The pathotype 66E16 was
not identified in the Northern Punjab and NWFP during 2005-06 but was found only
in Attock during 2006-07. On the contrary, pathotype 66E0 that was found in
districts Attock, Charsadda and Mardan during 2005-06 (Table 15) was not
identified in Attock district in the later year (Table 16). Moreover, pathotype 70E16
identified during 2006-07 in Attock, was not found in the previous year. The
occurrence of pathotypes 66E16 and 70E16 during 2006-07 at Attock, which was not
identified during 2005-06 may be because of the cultivation of different varieties
than sown during 2005-06. A single step mutation in 66E0 also suggests for the
identification of pathopype 66E16 in Attock during 2006-07. The prevalence of
pathotype 70E16 was noticed during both the study years in Nowshera, the district
that share it boundries with sistrict Attock (Fig. 5). Presence of pathotype 70E16 in
Attock during 2006-07, therefore, suggests for its air borne movement from
Nowshera to Attock. During the research period, the pathotype 67E0 remained
confined to districts Charsadda, Peshawar and Mardan. In the same way pathotype
80
6E0 was found in three districts of the Northern Punjab viz. Sailkot, Rawalpindi and
Chakwal during both the study years.
The difference in pathotype pattern was not observed over seasons in the NWFP with
the same 10 pathotypes identified over the two seasons per locality. Same was the
situation in district Sialkot, Rawalpindi and Chakwal where no difference in
pathotype pattern was identified. The reason may be attributed to the adoption of
same wheat varieties in the respective districts during both the stripe rust sampling
years.
The most frequent pathotype in three districts of the Northern Punjab (70E0) was
also identified in the three districts of NWFP with a pathotype abundance of 30.14
and 32.47 percent during 2005-06 and 2006-07, respectively. In the same years, the
second most abundant pathotype (71E0) was identified in the four districts out of
eight with 15.07 and 15.58 percent in the same order. The provincial differences in
the pathotypes composition specify that the breeding programs for wheat
improvement may capitalize on the approach that focus these provincial pathotype
populations’ variations to developed commercial varieties for specific areas. It is,
however, worth mentioning that considering a comparatively small sample size used
in this research, extensive studies encompassing more areas are imperative. The
composition of pathogen populations can transform with time as well, which must be
taken up under consideration in the wheat breeding programs. Accordingly, the
81
upcoming stripe rust studies in the country must focus on special and temporal
dynamics of rust population within and between the years.
In plant pathogen populations that are exposed to continuous directional selection
from increasing virulence, it is inevitable that the most complex pathotypes will
eventually dominate provided absence of other selective factors. In the present study
pathotype with a moderate complexity value of 10 is more frequent. The lowest
complexity value of 0 to 5 is not contained by any pathotypes (Fig. 8 and 9). These
results are in accordance with the findings of Chen (2005) who in the recent years
uncovered the predominance of Pst with a broad virulence spectrum as compared to
those with a narrow virulence band in North America. This phenomenon, however,
does not back up the general perception that isolates with limited virulence genes are
more belligerent and encompass better vigor than isolates with more virulence genes
(Vanderplank, 1963; Line and Qayoum, 1992). Lately, Chen (2004) demonstrated
this notion with P. striiformis f. sp. hordei in North America. The prevalence of a
broader or confined virulence spectrum is most likely influenced by selection force
from the genotypes of the host population at a particular location. Chen (2005)
proposed that, if the host population possesses comparatively limited resistance
genes then races containing just the virulence genes that match these resistance genes
ought to be encouraged by selection and therefore, should be inclined to be
predominant. Conversely, when the host population contains a number of resistance
genes then the races having potential to overcoming most of these genes must
become predominant.
82
In general, the pathotypes identified from NWFP collections and from Nowshera
district in particular, had higher genetic diversity when compared with the
pathotypes from the northern Punjab. The same may be attributed to the production
of both indigenous landrace cultivars and improved commercial wheat varieties in
NWFP. The wheat genotypes in this area are known to have high genetic diversity as
most of the existing Pakistani commercial wheat varieties contain resistance genes
Yr3, Yr6, Yr7, Yr9 and Yr27, individually or in combination. The isolates with high
complimentary genetic diversity, therefore, coexist with a broad range of genotypes.
The correlation is advocated by Manisterski et al. (2000) by noting strong influence
of genes in cultivated commercial wheat varieties, landraces as well as wild and
relative species on diversity of virulence in a pathogen population of P. recondita
f.sp. tritici in Israel.
Genetically few diverse stripe rust pathotypes were identified from the northern
Punjab in contrast to NWFP areas, which could be attributed to the cultivation of
wheat varieties with a little degree of genetic diversity. In districts of the Punjab
province, bread wheat mono-cropping system is the dominating production mode
year after year. The time has thus played a noteworthy role in sinking crop diversity
in the commercial fields (Ensermu et al., 1998). Inquilab-91 is the single largest
wheat variety that is being cultivated throughout the north-western Pakistan and due
to monoculture on a large scale, has now turned highly vulnerable to the stripe rust
(Singh et al., 2004a). Most of the commercially grown wheat varieties are deprived
83
of genetic variation against stripe rust because of similar genetic background (Gebre-
Mariam, 1991; Badebo, 2002).
Some pair similarities were observed for the populations from the Northern Punjab
and NWFP at a linkage distance of only 0.03 and 0.06 (Table 17). This has also been
illustrated by the cluster analysis that exhibited group resemblance within and
between pathotypes pertaining to these two provinces. It can, therefore, be suggested
that presence of stripe rust population in these two regions exhibit high adaptation to
commercial wheat varieties that are cultivated under both small-scale and
commercial state farmers.
2.5.2 Wheat Varietals Response to Stripe Rust
These were few varieties in trap nurseries, Punjab-85 at Islamabad, Shahkar-95 at
Rawalpindi while Shaheen-94 and Auqab-2000 at Peshawar that showed resistance
or partial resistance during 2005-06 but expressed susceptibility in the following
year. This suggests that virulence was not present or if present, was not effective
enough to overcome resistance. In the following year resistance seems to collapse
and this short term resistance failure led to a boom-and-bust syndrome (Kilpatrick,
1975).
Among the wheat rust resistance breeding programs, some achievements have,
however, been seen for some years as 1BL.1RS wheat-rye translocation (Zeller,
1973), linked with Yr9, Lr26, Sr31 and PM8 remained effective throughout the
wheat world till 1999. Presently, this translation is available in a number of high
84
yielding wheat cultivars, including Faisalabad-85 and Pak-81. Although virulence is
prevailing to Yr9 at the NWFP, however, some cultivars like Anmol-91, Zarlashta
and Wafaq-01, known to possess Yr9, showed resistant response during 2005-06 at
Peshawar (AUP) but exhibited susceptibility at the same site during 2006-07. Marvi-
2000, which demonstrated resistant to partial resistant response in the Northern
Punjab, fell susceptible in NWFP during 2005-06 and 2006-07 (Table 19, Appendix
V).
The commercial varieties in Pakistan that encompass the range between MRMS to
MSS type reaction might eventually express susceptibility after appearance of
virulence. Most of the commercial varieties contain either single or combinations of
Yr6, Yr7 and Yr9 while Inquilab-91 possesses Yr27 resistance gene. Data pertaining
to Isogenic lines given in Table 18 illustrate that virulence do occur in nature for all
these genes. The glasshouse experiments for virulence determination (Appendix III
and IV) visualizes that these genes also possess virulence at the seedling stage and
meanwhile confirms that virulence for these genes occur in nature.
The seedling stage performance of Inquilab-91 possessing Yr27, explained that it
remained susceptible to all race groups that were identified in the collection made
during 2003-04 and 2004-05 (Ahmad and Kazi, 2005). Such behavior of Inquilab-91
still persists when it fell susceptible to all the 13 race groups identified during 2005-
06 and 2006-07 (Table 9). On the other hand, the commercial cultivars SH-2002 and
Bakhtawar that were found resistant to the race group collection of 2003-04 (Ahmad
85
and Kazi, 2005) proved a contrary situation when tested under this study during
2005-06 and 2006-07. SH-2002 and Bakhtawar-93 was found resistant at Sialkot and
Peshawar (AUP), respectively during 2005 to 2007, while expressed susceptibility at
other WSRTNs. The commercial cultivar Marvi-2000 is worth to be mentioned here
as it showed resistant response (Rattu, 2007) and also expressed no disease reaction
at WSRTN Sialkot (2005-06 and 2006-07), Rawalpindi and Islamabad during 2005-
06. The variety Marvi exhibited TR to 10MS type disease reaction at Rawalpindi and
Islamabad, respectively during the following year and a susceptible type reaction
ranging from 10-40S at the WRSTN sites in the NWFP during both the
experimentation years (Appendix V). This is indicative of the fact that stripe rust
pathotypes have the potential to overcome resistance with time and accordingly have
rendered the varieties susceptible, which were considered to be resistant earlier on.
The key to control the cereal rusts is to use resistant cultivars (Johnson, 1981). The
cultivars remain resistant to rust for five or a bit more years depending upon the
agronomic lifespan, when a lively breeding program subsists. Some varieties fell to
rust as soon as they are cultivated. Most of such cases are attributed to the failure that
happen due to inadequate knowledge about the prevailing virulences in the pathogen
population. In some instances, mutations or possibly recombination in existing
virulences render the wheat crop susceptible.
Perusal of the compiled results of pathotype variation under glass house and TRAP
data from field exhibited that virulence was present for most of the known resistant
86
genes used in the present study. Although the virulence frequency for some of the
genes remained low, yet the presence of virulence against them is alarming under the
circumstances when genetic base of resistance is stumpy in the presently cultivated
varieties. It would, therefore, be imperative to incorporate the resistance gene in the
Pakistani cultivars, as identified in the present investigations.
2.5.3 Epidemiological Zone of the Indian Sub-continent
This epidemiological zone includes India, Pakistan, Bangladesh, Nepal and
Afghanistan (Saari and Prescott, 1985) where because of cooler temperatures
requirement, the hot summer's survival of rust is at northern higher elevations and
Southern Nilgiri hills in India. The zone that was once considered to have many
resistant cultivars with infrequent rust epidemics (Knott, 1989a) has now gone other
way.
2.5.3.1 Migration and introduction of stripe rust pathogen
Like in numerous other airborne fungal pathogens, the distant air dispersal and
infrequently by human activities enables spread of stripe rust pathogens to new
geographic vicinity. The urediniospore dispersal route has been described as the
“Puccinia path”. Seasonal dispersal of cereals rust pathogens over long-distance has
been narrated by Nagarajan and Singh (1990), Eversmeyer and Kramer (2000) as
well as Brown and Hovmøller (2002). The wheat stripe rust pathogen has been in
Europe and Asia for thousands of years and has been reported on the American
87
continents for over a century (Stubbs, 1985; Line, 2002). The pathogen was not in
Australia and New Zealand until 1979 when it was first reported in Australia
(O’Brien et al., 1980). Pst pathogen was possibly introduced from Europe through
urediniospore-contaminated clothing (Wellings and McIntosh, 1987). It probably
spread by wind dispersal to New Zealand the following year from eastern Australia
(Wellings and McIntosh, 1990). In central Africa, stripe rust was first reported in
northern Zambia in 1958 (Angus, 1965), but the disease was not found in South
Africa until 1996 when it was detected in the Western Cape (Pretorius et al., 1997).
From there it spread to most of South Africa in 1997 (Boshoff et al., 2002).
The resistance gene Yr9 was transferred from rye to wheat through the 1B/1R
chromosomal substitution and the 1RS/1BL translocation (Zeller, 1973). The Yr9
(Macer, 1975) has extensively been used in the wheat breeding programs through out
the world (Bartos, et al., 1973; Jing, et al., 1992; Johnson and Bimb, 1996;
McIntosh, et al., 1998; Saari, 1996; and Zeller, 1973). The Yr9 is tightly linked to
Lr26 (leaf rust), Sr31 (stem rust) and Pm8 (powdery mildew) resistance genes
(McIntosh, et al., 1998). The stripe rust races with Yr9 virulence were recorded first
of all in the former Soviet Union in 1973 and later on in the Netherlands, China, East
Africa, Middle East, South America, Mexico and in Australia in 1974 (Stubbs et al.,
1977), 1977 (Wang et al., 1986), 1986 (Saari, 1996), 1989 (Hakim and Mamluk,
1996), 1988 (Stubbs and Yang, 1988) and 1991 (Wellings and Burdon, 1992),
respectively. The Yr9 virulence was accounted in Iran, Pakistan and India during
1992, 1994 and 1996, respectively (Nazari and Torabi, 2000; Saari, 1996).
88
As the stripe rust fungus requires host crops with comparatively cool temperatures
than leaf and stem rusts, some epidemic exposed districts such as Nowshera, Attock,
Rawalpindi and Islamabad may serve as inoculum sources for recipient areas.
Seasonal migration of urediniospores in both directions between mountainous
regions and plains has been reported in China (Brown and Hovmøller, 2002; Wan et
al., 2004). The research study demonstrated that a single Pst colonel population of
70E0 with similar virulence pattern exists throughout the Northern Punjab (Sialkot,
Rawalpindi and Attock) and NWFP (Nowshera, Peshawar and Mardan) and that
Nowshera might be the recipient of migrant urediniospores from adjoining countries.
The assumption is in line with the study conducted by Hovmøller et al., (2002)
whereby a single race exists in Denmark, France, Germany and the United Kingdom
while Denmark was found the migrant spores’ recipient country. Prevailing air
movements, environmental conditions as well as crop seasons influence the seasonal
distant migration of rust spores in the Northern Punjab and NWFP. The northern
areas of Pakistan, might have acted as the recipient zones of urediniospores and the
distant dispersal of urediniospores would spread new races to the districts of Attock,
Rawalpindi and to Sialkot.
The Pst survival on grasses or wheat crop in the Himalayan mountain range might
also be the source of rust spore survival in summers and its dispersal occurs during
early winter with the prevailing northern cool winds towards plains of the Northern
Punjab. The presence of alternate host for the Pst has not been confirmed in this
region so far but study on these lines is required. The ability of the stripe rust
89
pathogen to survive near wheat fields through the non-cropping season plays an
important role in disease onset and the increase of inoculum during early growth
stages. The summer and autumn survival of the stripe rust fungus is dependent on
susceptible volunteer or self-sown wheat plants and, to a lesser extent, on grass
species (Sharp and Hehn, 1963; Shaner and Powelson, 1973; Wellings and
McIntosh, 1981; Stubbs, 1985; Dennis and Brown, 1986; Nazari et al., 1996).
Virulence on grasses provides greater opportunity for the survival and increases
stripe rust (McIntosh and Brown, 1997) which may play a role in the occurrence of
epidemics (Mardoukhi and Torabi, 1998). Factors that may lead to the rapid
evolution of aggressive pathogen races are increased fecundity and a microclimate
more conducive for disease development (Coakley et al., 1999).
The dispersal of urediniospores against the prevailing wind cannot be over sighted
and is quite possible, as demonstrated by the westward spread of P. striiformis f. sp.
hordei from southern Texas to California and the Pacific Northwest of the United
States from 1991 to 1995 (Chen et al., 1995). The southern winds during summer
may transport the Pst urediniospores from the plains of Northern Punjab towards
high mountainous areas of the country. The wind movement in both the directions
prevails in this part of the globe, which might facilitate the survival, occurrence,
distribution and development of the stripe rust pathogens.
The study thus conducted on the identification of races by collecting the samples
from various districts of the Northern Punjab and NWFP support these findings.
90
Advance studies by using molecular markers are, however, imperative to
authenticate findings of the envisaged research.
2.5.3.2 Influence of environmental factors on stripe rust
Like other diseases where three factors are essential in the disease triangle, the
development of stripe rust also relies strongly on a very precise weather conditions in
the presence of inoculum and susceptible host. The three imperative weather factors
that affect stripe rust epidemics include wind, moisture and temperature.
Wind is a key to spread or disperse stripe rust spores from pustules that develop on
infected leaves. Spores can live for a few days once released from a leaf and can
germinate and infect the leaf if land on another living wheat, provided the conditions
are suitable. Infection requires high humidity for 4 to 6 hours at 10 to 15°C, with
increasing time required at lower and higher temperatures. Infection seldom occurs
below about 2°C and ceases above 23°C (Murray et al., 2005). After infection, the
pathogen grows within the leaf and derives its nutrients from living wheat cells.
Growth is most rapid at 12 to 15°C that reduce to almost nil at 3°C and above 25°C.
If temperatures are outside the range for growth at any part of the day, rust stops
growing for that time but resumes growth when the temperatures become favourable
again at other times of the day. At 12 to 20°C, the fungal pathogen grows for about
14 days (shorter in some highly susceptible varieties) before the pustules erupt
through the leaf, with longer times of up to 80 days at 3°C and cessation of growth
much above 25°C (Murray et al., 2005). The time between infection and appearance
91
of symptoms is termed the latent period. Spore production is favoured by high
humidity and similar temperatures to the other stages of growth. Thus fresh spores
are usually seen in the morning because cooler temperature and still air are more
conducive for sporulation.
Moisture has an impact on spore survival, germination and infection. During high
humidity in winter, most spores remain in small clumps, which are relatively heavy
and fall out of air quickly, so their spread is mostly over very short distances. In
lower humidity, spores disperse singly in the air and can travel for much longer
distances (Murray et al., 2005). On the plant surface, at least three hours of
continuous moisture is required by the urediniospores to encourage stripe rust
(Rapilly, 1979). The regions where frequent moist conditions persist during the
growing season are likely to encourage stripe rust development. Pirsabak
(Nowshera), Attock, the twin cities of Rawalpindi and Islamabad are the areas where
a cool-moist weather pattern prevails during the month of February (Table 21) which
encourage stripe rust infection every year.
In rain-fed areas of the northern Punjab and NWFP, light showers exhibit perfect
conditions for infection as high humidity in the air and soil creates dew for several
nights during February 2006 and 2007 (Table 21). The rain drops act as a tool to
disperse spores and to spread the disease-urediniospores either by direct collision or
by splashing (Rapilly, 1979). Although high moisture encourages disease by
supporting spore germination but the extended or violent rain events wash off spores
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(Roelfs et al., 1992) already deposited on leaves and be detrimental to the spread of
disease (Sache, 2000).
Temperature impinges on spore germination, infection, dormant period, sporulation,
survival and the host resistance. P. striiformis has lower temperature optima so
prefers cool weather, which restricted the pathogen to temperate and tropical areas of
high elevation. The findings of the executed survey envisage that the prevalence of
stripe rust was more frequent in districts Nowshera, Attock and Rawalpindi where
the required set of environmental conditions persisted (Table 21). The studies are in
lines with the reviews narrated by Rapilly (1979) and Line (2002). The variation
found among the isolates may be attributed to the temperature response of P.
striiformis.
2.5.4 Pst Distribution in the Northern Punjab and NWFP
Stripe rust of wheat has become increasingly important in the Northern Punjab and
NWFP during the recent years. This situation has put the disease under constant
watch of the rust pathologists at CDRP, NARC, Islamabad through collaborators by
field and disease nurseries survey throughout Pakistan.
In Nowshera district, prevailing environmental conditions such as high humidity and
cool temperature (Table 21) are complimentary for stripe rust occurrence every year.
Presence of susceptible cultivars is imperative for stripe rust epidemics with
prevalence of favorable environmental conditions. Out of the 13 identified races, six
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were detected in the diseased samples pertaining to a single Nowshera district
including two races that are also virulent to Hybrid 46 (Yr4+).
Table 21 Environmental Conditions Persisted during 2005-06 and 2006-07 in
the Northern Punjab and NWFP
Years 2006 2007
Month
Temperature (°C) Rainfall
(mm) Humidity
(%)
Temperature
(°C) Rainfall (mm)
Humidity (%)
Min Max Min Max
Sialkot
February 11.9 26.0 81.3 88 9.3 19.7 89.3 92
March 13.4 26.7 55.6 80 12.7 25.1 176.5 86
April 17.9 34.8 0.5 54 20.1 36.0 3.0 61
Rawalpindi
February 9.7 25.0 25.6 61 6.6 19.3 93.6 76
March 11.4 26.2 45.5 65 9.0 23.1 143.2 65
April 15.3 32.7 20.3 37 15.9 33.9 19.6 44
Attock
February 7.7 25.9 10.0 66 6.6 18.6 288.0 74
March 11.4 26.2 39.0 65 9.2 23.0 189.0 68
April 14.7 33.0 17.0 41 15.7 34.4 52.0 49
Peshawar
February 11.9 25.9 17.5 65 8.9 18.5 159.1 75
March 12.6 25.2 27.4 65 11.9 23.5 80.9 66
April 16.5 32.7 15.3 45 18.8 34.4 14.6 54
Cherat (Nowshera)
February 9.0 16.4 21.0 61 5.7 9.9 440.5 74
March 7.9 16.3 82.0 67 5.4 14.6 116.4 65
April 10.2 24.8 13.0 45 18.6 27.0 40.5 46 Source: National Agro-Meteorology Centre, Islamabad
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The distribution of stripe rust was extensive in the Northern Punjab and NWFP in
2006-07 as compared to the pervious year. The winters in the northern regions of
Pakistan during 2005-06 and 2006-07 remained favorable for establishment of stripe
rust. The cool spring weather with more than 60 percent humidity in March (Table
21) during both the experimentation years boosted rust development (Chen et al.,
2002) and its spread in all the 8 districts of the study area (Fig. 5).
Occurrence of wheat stripe rust was observed throughout the four districts of the
Northern Punjab during 2005-06 and 2006-07 during March attributed to the perfect
environmental conditions (Table 21). The districts of Sialkot and Chakwal exhibited
trace to five percent disease severity while the districts of Rawalpindi and Attock
responded with stripe rust severity from trace to 20% especially where Inquilab-91
was under cultivation (data not given).
Wheat stripe rust was observed to be severe in district Nowshera with 10 to 40
percent disease severity and up to 30 percent disease incidence in late March during
both the years. Cool temperatures and more rains than normal in 2006-07 (Table 21)
provided humid conditions that allowed stripe rust to establish on wheat varieties
cultivated in the area. Favourable weather conditions also allowed wheat stripe rust
to spread in commercial fields of several cultivars around the trap nursery in
Pirsabak and Nowshera. In Peshawar, Charsadda and Mardan, stripe rust was light
ranging from trace to 10 percent in farmers' wheat fields. The reason for less disease
severity in farmers’ fields may be attributed to less inoculum pressure as compared
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to that in the trap nurseries where bulk stripe rust inoculum is tested year after year
under various experiments. Use of fungicides as well as adoption of resistant
varieties would also be a reason for low disease severity in the farmers’ fields. High
severity levels of wheat stripe rust were, however, observed in trap nurseries
established at Pirsabak and Peshawar (Appendix V). Although, stripe rust is
prevalent in the late March and early April of both the years but development was
slower than normal. This was due to dry conditions with increased temperature
especially in the month of April (Table 21), which were not conducive for rust
development (Chen et. al., 2002).
2.5.4.1 Race studies in the Northern Punjab and NWFP
Race frequency is determined by two opposing forces. Virulence is the first force
essential for obligate parasites, such as P. striiformis, to infect host plant, grow and
reproduce. The more virulence genes a race has, the more cultivars it is capable of
infecting, increasing its frequency in the pathogen population. The second force is
the cost of unneeded virulence. A wide virulence spectrum may result in reduced
fitness and aggressiveness (Vanderplank, 1963; Line and Qayoum, 1992).
Many of the wheat genotypes that have been used to differentiate races of Pst have
more than one gene for resistance. The presence of two or more genes in a single
differential genotype makes it difficult to quickly detect new races that have
virulence to a single resistance gene. Wellings et al. (2004) at the Plant Breeding
Institute, University of Sydney, Cobbity, New South Wales, Australia, developed
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near-isogenic lines in the ‘Avocet Susceptible’ background for 13 Yr genes including
Yr1, Yr5, Yr6, Yr7, Yr8, Yr9, Yr10, Yr15, Yr17, Yr18, Yr24, Yr26 and Yr32, which
have been used throughout the world for monitoring virulence genes of Pst. More
effort is needed to develop near-isogenic lines for the other Yr genes which would
eventually replace the current differential genotypes used for identifying races.
The existence of stripe rust races 66E0, 67E0 and 70E0 identified by Kirmani (1980)
from samples collected during 1969 indicate that all three races still exist and are
stable in nature. The large region of the Northern Punjab and NWFP may be
considered as a single epidemic region under 70E0 pathotype, which was detected in
this region four decades back. In 2006 and 2007, Pst race 70E16, similar to 70E0 and
71E0 with additional virulence to Compair, was detected in the region. The
unwavering race composition could be due to few wheat varieties that are
extensively cultivated over the years with similar genetic background as the previous
wheat breeding programs have not generally been concerned with the resistance to
stripe rust. Since, less frequent occurrence was witnessed for stripe rust in this
region; therefore, required priority was not extended in the previous wheat breeding
programs. After the dawn of new millennium, the race composition has became a
complex due to introduction of a number of wheat cultivars such as SH-2002,
Bahawalpur-2000, Bahkar-2002, Marvi-2000, AS-2002, Iqbal-2000, Auqab-2000,
Moomal-2002, Zarlashata, GA-2002, Wafaq-01 and Manthar.
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The situation of stripe rust of wheat in the northern region of Pakistan also illustrates
virulence transformation in the stripe rust. The Pst races 66E0, 67E0 and 70E0
reported by Kirmani (1980) were again identified among the nine physiological races
from the samples collected during 2003-04 and 2004-05 cropping season from
different wheat growing areas of northern Pakistan (Ahmad and Kazi, 2005). We
could speculate that initially a single race might be introduced into the country
through the northern winds following the Puccinia pathway of the epidemiological
zones in the Indian Subcontinent (Saari and Prescott, 1985). These three pathotypes
identified during 1969, belong to the same race group as evident from Fig. 10, spread
to NWFP and the Northern Punjab during the following years. These races might
have evolved into the identified 13 races within last four decades. The assumption
can be associated with the findings of Wellings and McIntosh (1990) as Pst that
introduced in Astralia in 1979 spread to New Zealand in 1980 as a single race, which
evolved in 15 races with in just 10 years.
Of the 13 races identified from isolates of the disease samples collected during 2005-
06 and 2006-07 in the eight districts, one was new. Generally, the stripe rust isolates
collections pertaining to the Northern Punjab were similar in virulence to the
collections from NWFP. Of the 13 races, one (6E1) and four (70E16, 66E16, 7E16
and 6E16) pathotypes were identified through their virulent response on Hybrid 46
and Compair, respectively. These races have recently been identified in Nowshera
district while 70E16 and 66E16 have been reported for the first time in the district
Attock. Among the identified pathotypes, 6E17 was the one that broke resistance for
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genes Yr4+ and Yr8,19 APR. The identified Pst races during 2005-06 and 2006-07
had various virulence combinations to Yr1, Yr6, Yr7, Yr8, Yr9, Yr12, Yr17, Yr18,
Yr24, Yr26, YrSk and YrA but all the races possess virulence on Yr6, Yr7, Yr9, Yr17,
YrSk, YrA and Jupateco S. The races 70E0 and 71E0, virulent on Yr6, Yr7, Yr9, Yr17,
Yr18, Yr24, YrSk and YrA remained widely distributed in the Northern Punjab and
NWFP during the wheat season of 2005-06 and 2006-07.
Most of the races virulent on Yr18 identified in the present investigations have the
virulence on Yr1, Yr8, Yr12 and Yr24. The resistance gene Yr6, Yr7 and Yr9 are
dominant genes incorporated in the genetic makeup of the Pakistani wheat varieties
(Table 19). During 2005-06 and 2006-07, all the commercial Pakistani wheat
varieties possessing Yr9 resistance genes remained free of stripe rust at the
established WSRTN only in Sialkot (Table 19). In Pakistan, Yr27 was not widely
used in commercial wheat cultivars and due to this reason Inquilab-91 survived for
the longest period in the major stripe rust zone of the Northern Punjab and NWFP till
2003-04.
The resistance gene Yr8 originally pertains to Triticum comosum (Sibth. & Sm.)
Richter and translocated on chromosome 2D of wheat in ‘Compair’, ‘Hobbit Sib’,
and ‘Maris Widgeon’ (McIntosh, et al., 1998). Another resistance gene, Yr17 was
originally from Triticum ventricosum Ces., which was transferred to a number of
wheat cultivars including Mandsen and Hyak and the gene Yr26 originated from
Triticum turgidum L. and was mapped on chromosome 1BS (Ma et al., 2001).
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Likewise, the most effective Yr resistance gene, Yr15 was derived from T.
dicoccoides and is located in chromosome 1BS and expressed a linkage of 34 cm
with Yr10 (McIntosh, et al., 1996).
2.5.4.2 Stripe rust occurrence during 2005-06 and 2006-07
Stripe rust was first observed in first week of March 2006 on the susceptible cultivar
Morocco in the WRSTN established at Nowshera followed by Rawalpindi during the
second week of March (Fig. 12 a, b and c). In the following year, heavy and above-
average rain showers together with lower than average minimum and maximum
temperatures recorded during February and March 2007 (Table 21) contributed
significantly to the occurrence of stripe rust.
Due to prevalence of conducive environmental conditions, the expression of stripe
rust was much earlier in the WRSTN established at Nowshera as the disease was
detected in the third week of February 2007, while at Rawalpindi during the 1st week
of March 2007.
The universally susceptible cultivar Morocco was 1st at all the WRSTN sites that
expressed susceptibility to stripe rust. During both the years, subsequent surveys at
the time of detection showed that the disease was well established in wheat fields of
NWFP and the northern parts of the Punjab. The initial location and time of
outbreak, therefore, could not be determined.
100
Fig. 12 a. Susceptible cultivar – Morocco exhibiting stripe rust in the WRSTN. b. Urediniospores of Pst. c. A glance at WRSTN established in experimental area at PMAS-AAU, Rawalpindi.
a
c
b
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Table 19 reflect that almost all the commercially grown cultivars at the five WRSTN
sites remained susceptible to stripe rust except at the WRSTN established at Sialkot
during 2005-06 and 2006-07. The susceptible cultivar Morocco and Kohistan 97
were the only two cultivars that responded with 80S-90S and 10R-5R reaction,
respectively, during 2005-06 and 2006-07 at the WRSTN established at Sialkot. No
disease was expressed by any other commercial variety at this site during both the
years. The disease was also prevalent in the commercial fields of the stripe rust prune
hot spots in Rawalpindi, Islamabad, Nowshera and Peshawar. Due to hot and dry
weather conditions in these regions, the incidence of stripe rust was low after the first
week of April 2005-06 and 2006-07.
After the first time detection of Pst on wheat in the district Nowshera during March
2006 and February 2007, the pathogen intruded to the important wheat-production
areas in NWFP and the Northern Punjab. The disease subsequently established itself
as an endemic disease with an enormous impact on wheat production in the cooler
regions of the country.
Out of 8.303 million hectares, more than 5.8 million hectares (70 percent) of wheat
production area is under the threat of stripe rust (Singh, et al., 2004b). The wheat
production dry land areas central and southern Punjab largely escapes stripe rust
epidemics. Factors that contributed to the escape are less dense stands due to low
seeding rates (80-100 kg ha-1), drought conditions with low humidity during early
growth stages, followed by high day temperatures after the first spring rains. High
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day temperatures experienced during February and March in the irrigated areas
combined with presence of cultivar resistance may have contributed to the absence
of disease. Park (1990) has defined those days favorable for stripe rust when the
mean temperature is within the range of 12.4 to 18.4 °C and the minimum is between
7.3 to 14.6 °C. The condition for both minimum and mean temperatures in the
Northern Punjab and NWFP fulfilled during February and March of 2006 and 2007
(Table 21), when the stripe rust was expressed. The highest rainfall in the two years
of study was recorded in Cherat (Nowshera) and Attock during February 2007 as
440.5 mm and 288 mm (Table 21), respectively that contributed to the early
establishment and buildup of stripe rust pathogen. The temperature becomes lowered
and leaf wetness was prolonged due to high rainfall, which contributed towards
disease spread because of rain splash (Geagea, et al., 1999). Surveys identified 13
stripe rust pathotypes in Northern Punjab and NWFP. The susceptible reaction of
Inquilab-91 against Pst shifted some cultivation to other resistant cvs like Bakhtawar
and SH-2002, which remained resistant during 2003-04 but fell susceptible to
pathotypes 71E0 and 34E0, respectively in the following year (Ahmad and Kazi,
2005). Similarly, AS-2002 which showed susceptible reaction to Pst races 66E0 and
70E0 in 2003-04 also expressed susceptibility to 68E0 and 34E0 during 2004-05.
Another commercial variety, Marvi-2000 when released was ranked among the
resistant cultivar also proved susceptibility to race 6E0 in year 2003-2004 collection
(Ahmad and Kazi, 2005).
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The high selection pressure consequently resulted in the development of pathotype
66E16, which seems to be a single-step mutation process in 66E0 thereby resulting
in additional virulence for Yr8, Yr12 and Yr26. The identified pathotype 6E16 has
previously been detected in East and North Africa, the Middle East, and Western
Asia (Badebo, et al., 1990; Louwers, et al., 1992). It is quite possible that few
variations in this pathotype may occur some where as many resistance genes have
not been reflected in the classification of pathotypes worldwide. With the available
data, it is not possible to speculate as to the likely origin of pathotype 6E16. The
pathotype was, however, expressed virulence to the resistance genes Yr6, Yr7, Yr8,
Yr9, Yr12, Yr17, YrSk and YrA.
From the field data (Appendix I), it is evident that three differential cvs. in the world
set Moro (Yr10), Strubes Dickkopf (YrSd) and T. spelta album (Yr5), all the eight
European differentials and the NILs with Avocet-S carrying Yr5/6*, Yr10/6*,
Yr15/6* and YrSp/3* resistance genes that are still effective against the prevailing
pathotypes at the stripe rust hot spots. The resistance gene Yr27 present in Inquilab-
91 produced a compatible seedling reaction when scored 15 to 17 days after
inoculation. Based on the glasshouse and field data, it is obvious that the seedling
gene Yr27 is not effective. Supplemental lines with Avocet-S background that
exhibited heterogeneous field reactions were Yr24/3* and Yr26/3*. The absence of
most of the effective seedling genes from commercial cultivars (Pretorius, 1998) and
the resulting low selection pressure preclude any predictions of their durability to
local stripe rust pathotypes. Seedling genes with a hypersensitive response to
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avirulent isolates of Pst are well known for their vulnerability to the development of
more virulent pathotypes (Wellings and McIntosh, 1990; Danial, et al., 1995; Ma
and Singh, 1996; McIntosh and Brown, 1997; Shan, et al., 1999). The seedling genes
Yr8 and YrSk have become ineffective due to changes in the pathogen population
without the presence of these genes in Pakistani wheat (Wellings and McIntosh,
1990; McIntosh and Brown, 1997), however, the resistance genes Yr5 and YrSp are
still effective. Adult plant resistance against Pst has been known to last longer than
seedling genes (Park and Rees, 1989; Johnson, 1992; Broers, et al., 1996; Ma and
Singh, 1996; McIntosh and Brown, 1997). The adult plant gene Yr18 present in
Jupateco R and Yr18/3*Avocet-S now exhibits low level of resistance under
Northern Punjab and NWFP environment. Although the adult plant genes Yr29
present in cultivar Pavon-76 showed an MS type reaction only at Rawalpindi and
Islamabad, it might still have relative level of resistance compared with other
susceptible varieties cultivated in Pakistan. It can, therefore, still be used as a
potential cultivar and the responses are expected to provide adequate crop protection.
The key determinants of wheat stripe rust epidemic development are the biotic and
abiotic factors. The former pertains to frequencies of resistance and virulence genes
while the later deals with the climatic conditions. The history of speciation and
global spread of wheat stripe rust is still in the dark. Recent laboratory work
conducted in France and Denmark, however, described stripe rust populations with
high pathotype diversity in China and Pakistan as compared to Europe (Pope, 2008).
105
A big leap towards understanding of the rust pathogen spread and its adaptation
would not be possible without identification of its origin center. Although the
continental diffusion of Pst is on record (Brown and Hovmøller, 2002) but its center
of origin is still unknown. The studies conducted for barley pathogens envisage that
the origin centre of pathogens may vary from the origin center of the corresponding
host (Zaffarano et al., 2006; Brunner et al., 2007). Middle East is the center of origin
for cultivated triticale, whereas the center of diversity for wheat stripe rust is
supposed to be in Transcausia (Stubbs, 1985). A very low genetic diversity among
Pst populations has been reported so far in most parts of the globe, (Hovmøller et al.,
2002; Enjalbert et al., 2005) but in contrast, huge diversity has been reported in
Chinese populations because of parasexuality and over-summering in highland areas
(Shan et al., 1998). In Pakistan, the stripe rust population has shown a diversity of an
intermediate level. The situation thus raises a query about an Asiatic location for the
origin centre of Pst, which can be tackled by conducting global study on the diversity
of Pst populations by using common molecular markers and developing samplings in
non-adequately described areas.
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Chapter II
2.6 CONCLUSIONS
Stripe rust of wheat caused by an obligate pathogen Pst stands among the most
important diseases of wheat worldwide. Despite frequent occurrence of the stripe
rust epidemics in the northern region of the country, the exploration corresponding to
the structures of pathogen population is still lacking. In the present study, pathotype
variation of Pst population that occur naturally in the major wheat growing areas of
Pakistan were analyzed. The outcomes were highlighted in the context of prevailing
virulences and identification of the Yr resistance genes that are still effective. These
resistant sources can play a groundbreaking role in stripe rust management through
their exploitation in the wheat breeding program for development of resistant
cultivars. The summary of the major findings are outlined as follows:
The virulence and diversity of the stripe rust population prevalent in the Northern
Punjab and NWFP was probed using differential genotypes with recognized Yr
genes. The physiological races were classified into 13 different pathotypes identified
from 150 isolates. There were marked differences in the composition of individual
sub-populations in the study area of northern Pakistan. Out of the 13 diverse
pathotypes, only four and five of them were detected in more than three districts
during 2005-06 and 2006-07, respectively. In 2005-06 and 2006-07, 12 previously
identified races were confirmed while one new race was identified. Five and two, out
of the 12 identified races were found virulent on Compair (8,19,APR) and Hybrid 46
106
107
(4+), respectively. Pathotypes 70E16, 66E16, 7E16, 6E17 and 6E16 were virulent on
Compair (8,19,APR) have recently been identified in Nowshera district while 70E16
and 66E16 have been reported for the first time in the district Attock. The races 70E0
and 71E0, virulent on Yr6, Yr7, Yr9, Yr17, Yr18, Yr24, YrSk and YrA remained
widely distributed in the Northern Punjab and NWFP during the wheat season of
2005-06 and 2006-07.
Substantial genetic variation was detected with in and between populations on the
basis of areas of collection. The Pst 70E0 was the most frequent pathotype that was
identified in six out of eight districts during 2005-06 and 2006-07. None of the eight
districts, however, shared a single common stripe rust pathotype. Such variation in
the pathotypes might specify that wheat varieties could have been developed and
released for cultivation on a climatic or provincial basis.
The measure of resistance genes effectiveness termed as virulence gene frequency,
ranged from 0 to 100%, where as the number of the virulent genes out of 29 referred
as virulence complexity, fluctuated from 6 to 15.
The stripe rust resistance genes Yr2+, Yr3V, Yr3N, Yr5, Yr6,2+, Yr7+, Yr9,2+, Yr10,
Yr11, Yr15, YrSd, YrCv and YrSp were uncovered as effective against all isolates. To
deploy these Yr resistance genes either singly or in combination in the upcoming
wheat breeding program could play an effective role to lessen yield losses inflicted
by stripe rust. The gene pyramiding approach has, however, an edge to develop
varieties possessing long-lasting resistance.
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Although resistance genes Yr4+, Yr8+, Yr26 and Yr29 (Pavon 76) expressed partial
virulence they still have prospective for exploitation. Varieties encompassing these
stripe rust resistance genes can be opted for their deployment in combinations as
sources of resistance in wheat rust breeding programs to assimilate their genetic
traits in the national germplasm. This integration will assist in exploitation of future
high yielding durable rust tolerant wheat genotypes with a broad genetic base
thereby ensuring robust wheat production besides availability of resistant germplasm
resource, which could be exploited as and when required.
The genes Yr6, Yr7, Yr9 and Yr27, present in the genetic background of most of the
commercial Pakistani wheat cultivars, previously characterized as effective source of
resistance against stripe rust, have now articulated high virulence frequency (100
percent). This may advocate drastic change of Pst pathotype among the population
that occurred in the recent past.
Virulent pathotypes have already been observed to most of the Yr genes used in
Pakistani wheat varieties. A quest for hunting new and efficient sources of stripe rust
resistance genes is imperative to tangle the alteration in the host-pathogen
interaction. The genes, Yr6, Yr7 and Yr9 were the most frequently encountered stripe
rust resistance gene either alone or in a blend with other Yr genes, investigated in the
commercial wheat varieties. These previously resistance genes now lack sufficient
protection against an extensive series of Pst isolates collected across the major wheat
cultivated regions of the northern Pakistan. Non-replacement of these cultivars by
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the ones with more effective genes would endanger the golden grains harvest in an
ample quantity.
The epidemics that witnessed during 1977-78 and 1992-93 cautions that stripe rust
could be a serious threat for wheat in Pakistan whenever the pre-requisites of disease
triangle is accomplished. Production losses associated with the disease and non-
affording of high input costs due to the repeated fungicidal application necessitate
the development of sustainable control measures against yellow rust. Under the
present growing circumstances of predominantly dry-land wheat production coupled
with uncertain rainfall, breeding for resistant cultivars is emphasized and it will offer
the most reliable and cost-effective channel for controlling stripe rust. Effective
breeding approach, therefore, strongly relies upon the understanding of genetic
variation in both host and pathogen. Effective disease control approach further
necessitates an epidemiological understanding of the pathogen. This includes the
potential of stripe rust pathogen to persist during non-crop season and disease
occurrence probability in different wheat-cultivated areas of the Northern Punjab and
NWFP. The same will have a restraining influence on the release of cultivars that are
susceptible in hot spot areas, which will be an important aspect in selection of a
cultivar by the farmers.
The higher elevation areas in North of Pakistan that comprise of a different growing
season than the main wheat areas in Northern Punjab and NWFP, provide a zone for
the over-summering of stripe rust. These sources of inoculum can be minimized by
110
planting resistant cultivars, especially in the northern areas where summer rain and
mild summer temperatures can significantly contribute to the summer survival of
stripe rust. Stripe rust control strategies in the Northern Punjab and NWFP should be
directed at reducing the probability of epidemics and magnitude of losses. The latter
may be obtained by avoiding the release of cultivars containing only genes for
seedling resistance and the planting of stripe rust resistant cultivars in the more rust-
prone areas. It is imperative to continue monitoring of the stripe rust population for
pathotype changes so that new pathotypes with the potential to overcome resistance
genes currently deployed can be detected early.
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Chapter III
ESTIMATION OF WHEAT YIELD LOSSES DUE TO STRIPE RUST
3.1 INTRODUCTION
Plants contribute around 93 percent of the food to feed the people of the world, two-
third of which is contributed by the cereals (wheat, maize, rice, barley, sorghum, and
millet). About 80 percent of the global cereal production is shared by wheat, maize,
and rice. Among the cereals, wheat is the largest one being cultivated in 27 countries
of the developing world. Of the two principle types of wheat i.e. Bread and Durum,
90 percent of the world’s wheat production is attributed to bread wheat (Stubbs,
1985). Wheat is used mainly for human consumption and is sole source of energy for
nearly 35 percent of the world population (Dreisigacker, 2004). It is estimated that
more than 75 percent of the world’s population consumes wheat as part of their diet
daily. In parts of northern Africa and in the newly independent republics of the
Caucuses region, annual consumption per person is the highest at around 200 kg
(Maarten and Ogbonnaya, 2006) whereas in Pakistan, per capita wheat consumption
is around 124 kg/annum (MINFAL, 2007).
To achieve self-sufficiency, sustainable productivity of wheat is of paramount
importance in the context of food security. Accordingly, there is a dire need to bridge
this gap to ensure food security for the country’s inhabitants. The foremost trouble
with our agriculture is lack of innovation in agricultural research. During the last few
111
112
decades, the researchers have not been able to come with any high-yielding and
disease resistant variety. Hence, the wheat growers have no option except to cultivate
the varieties introduced in the early nineties. A wheat research program is, therefore,
vital whereby the scientists would make collaborative efforts for the genetic
improvement and development of new disease and pest resistant varieties. At the
same time, it is imperative to acquire maximum yield from the existing varieties
through better agronomic and plant protection measures. Huge losses in wheat yield
are attributed to the invasion of various diseases, which have caused enormous yield
losses in the past few years. Planting resistant varieties is a simple solution if such
varieties have been introduced because breeding for disease resistance is a
continuous phenomenon as new races of rusts evolve and overcome resistance with
the passage of time. A wheat variety that has been resistant in the past may not
remain resistant to new races of rusts (Brian, 2006).
Limiting factors in wheat production are associated with several biotic and abiotic
stresses. Of these in the former category, rust diseases are the most significant wheat
diseases, which have continued to ravage this crop since ancient times. Stripe rust,
also known as yellow rust, caused by the parasitic fungus P. striiformis, is one of the
most damaging diseases of wheat in many areas around the world (Zadoks, 1961;
Chen, 2005; Fu et al., 2008). The obligate parasitic fungus has devastated cereal
production worldwide due to rapid systemic infection of affected plants resulting in
defoliation and shrivelled kernels. Stripe rust probably occurred long before wheat
was grown for food, but Gadd from Europe first described it in 1777 (Eriksson and
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Henning, 1896). Hassebrauk (1965), Stubbs (1985), Line (2002), and Li and Zeng
(2003) were among those who reported occurrence, damage caused by the stripe rust
and its distribution around the world. By now, stripe rust of wheat has been reported
in more than 60 countries of the globe.
The stripe rust attacks all the above-ground parts of the wheat plant and is
characterized by powdery masses (pustules) of yellow spores, which form stripes on
the leaf surface. The pustules occur in blotches covering large areas of affected
leaves on young plants while pustules are arranged in parallel lines, giving affected
leaves a characteristic striped appearance on older plants. Infection of leaf sheaths
and stems can also occur, but spore production on these parts is less than on leaves.
Stripe rust can also attack the heads. The rust fungus infects the glumes and awns at
flowering, which results in an accumulation of spores in the florets and on the
surface of the developing grain (Grant et al., 2005).
Through the envisaged studies, an attempt has also been made to assess the effect of
stripe rust towards yield loss in the commercially grown wheat varieties in rain-fed
areas of Pakistan. The study was imperative to evaluate the economic impact of this
disease thereby predicting the losses, if the disease occurs in epidemics. The research
was executed keeping in view the following objective:
1. To ascertain wheat yield losses due to stripe rust in field under disease stress
conditions.
114
Chapter III
3.2 REVIEW OF LITERATURE
Literature has depicted that stripe rust of wheat reduces the weight and quality of the
kernel and forage. Seed produced from crop damaged by stripe rust are shriveled,
have low vigor and thus express poor emergence after germination. Stripe rust can
cause 100 percent yield loss if infection occurs very early and the disease continues
to develop during the growing season provided the cultivars are susceptible. In most
wheat producing areas, yield losses caused by stripe rust have ranged from 10-70
percent depending upon the susceptibility of cultivar, earliness of the initial
infection, rate of disease development, and duration of the disease (Chen, 2005).
Severe epidemics may result in substantial losses as annual losses due to yellow rust
varied averaging 2 to 15 percent in Danish wheat trials during 1988 to 1990 (Ullerup,
1991), and for the most susceptible cultivar, losses of almost 50 percent (equal to 4
t/ha) were recorded in 1990 (average of 236 trials). Losses of the same magnitude
were reported for susceptible cultivars in Asia (Sharma et al., 1985). An early
investigation conducted by Conner and Kuzyk (1988) showed that wheat yield
reduction in the Lethbridge area was up to 75 percent and reached 45 percent at
Lacombe based on their recent observations. Yield reductions in some wheat plots
during 2004 and 2005 were undoubtly high as disease severity reached 100 percent
at Lacombe, Olds and Trochu in Alberta (Kequan Xi et al., 2007).
114
115
Estimates of yield losses of cereals caused by rusts since 1918 are maintained by the
United States Department of Agriculture, Agricultural Research Service, Cereal
Disease Laboratory in St. Paul, Minnesota (Chen et al., 2004). They have further
reported that wheat yield losses caused by stripe rust were, however, not recorded
until 1958 when a loss of 2.9 million bushels (about 78,996 tons and 4 percent of
production) was recorded in Washington State. In this State, the most severe yield
losses recorded were 25 percent (591,108 tons) in 1960 and 17 percent (787,236
tons) in 1976. In the past 46 years in the Washington State (from 1958 to 2003),
stripe rust occurred almost every year. Severe damage (a statewide yield loss of 5
percent or more) occurred in 12 of the 46 years, all of which occurred before 1990
and most of which before 1984.
Chen (2004a) reported that yield losses were generally low after the early 1980s
because of the use of resistant cultivars and the application of effective fungicides.
Afterwards, in 2002, due to extremely favorable weather conditions for stripe rust in
Washington State, the disease affected 440,000 acres (70 percent of the acreage) of
the spring-wheat crop and 45,000 acres (2.5 percent of the acreage) of the winter-
wheat crop, 20 percent of the total wheat acreage. A cost of more than US$2.5
million was incurred to spray fungicides on an area of about 170,000 acres without
which stripe rust could have caused a yield loss of 20 – 25 percent (155,268 –
193,404 tons), valued at about US$26 to US$33 million. He has further intimated
that frequent epidemics have occurred in Oregon and Idaho because of their
116
geographic proximity to Washington State having similar weather conditions and
cropping systems.
Chen (2004a) further signified the damage caused by stripe rust by adding that the
disease was not recorded in California (USA) until 1974, when a yield loss of 8
percent was estimated. Later on from 1975 to 1997, yield losses in California were
below one percent but in 1998, stripe rust became an increasingly severe problem
which in 2003 caused 25 percent yield loss. He added that by this time the epidemic
was so severe and widespread that fungicides were used for the first time in
California. In addition to the favorable weather conditions in 2003, the major reason
for the severity of the epidemic was that the major wheat cultivars with race-specific
resistance were overcome by new races of P. striiformis f. sp. tritici Eriks.
Chen (2004a) reported that in the United States, stripe rust was primarily a problem
in the Washington, Oregon and Idaho but since 2000, the disease has become
increasingly important in the south central states (Texas, Louisiana, Oklahoma,
Arkansas, and Mississippi) and the central great plains (Kansas, Colorado, Missouri,
and Nebraska).
In Arkansas, the only significant epidemic that was recorded before 2000 occurred in
1987 with a statewide yield loss of 2.5 percent. The disease caused yield losses of 7,
5, and 3 percent in 2000, 2002, and 2003, respectively, plus the millions of dollars
that were spent on fungicide applications each year. Stripe rust seldom caused
significant damage in Kansas before 2000, but the State suffered yield losses of 7.3
117
percent (706,606 tons) in 2001 and 10.6 percent (1,573,110 tons) in 2003. Similarly,
Colorado had an 8 percent (156,358 tons) yield loss in 2001, and Nebraska had a 10
percent (256,328 metric tons) yield loss in 2003. Monetary losses due to the
reduction of wheat yield caused by stripe rust in the central great plains of the US
especially Kansas, were estimated to be $27M, $119M, $24M, and $267M for 2000,
2001, 2002 and 2003, respectively (Chen et al., 2004). Including the cost of
fungicide applications, losses due to stripe rust amounted to more than US$300
million during 2003 (Chen, 2005).
In USA, Small Grain Workgroup carried out a thorough probe into epidemic and
losses caused by the stripe rust. It was observed that stripe rust was devastating to the
California wheat crop in 2003, while many other states across the country also
experienced severe stripe rust attack. The disease developed early and was more
severe than normal throughout the U.S. The workgroup demonstrated the impact of
the disease in California as highly susceptible varieties suffered >75 percent loss due
to the stripe rust in fields receiving no fungicide applications; whereas susceptible
varieties sustained 50-70 percent loss; moderately susceptible varieties 20-40 percent
loss; and moderately resistant varieties 5-15 percent loss (Anonymous, 2004).
The wheat growers faced a loss of about 8 and 40 million AU$ during 1983 and 2003,
respectively for fungicide application to control stripe rust during severe epidemics in
Australia (Wellings and Luig, 1984; Wellings and Kandel, 2004). In Ethiopia, yellow
rust epiphytotics were recorded in 1977, 1980-1983, 1986, 1988 and 1990 with
118
severe losses during 1988 in bread wheat (Badebo and Bayu, 1992). In recent years,
stripe rust caused severe damage to wheat in several other countries. Although the
disease was observed for the first time in South Africa in 1996, it caused a loss of
$22.5 millions due to widespread epidemic in spring wheat that year because of
cultivar susceptibility and favorable weather conditions (Pretorius, 2004). Epidemics
occurred again in the central and western Free State in 1997, the eastern Free State in
1998, and all wheat-growing areas characterized by a weather pattern of summer
rainfall in 2002. The 1998 epidemic on about 42 000 ha of winter wheat in the
eastern Free State resulted in losses of ZAR 12 million (~US$2.25 million)
(Pretorius, 2004).
Major epiphytotics were recorded in Egypt once in each decade since 1960s with the
most recently reported in the Delta region during 1995. The average grain yield loss
ranged from 14 to 26 percent in this region, while the national loss was about 10
percent (El-Daoudi et al., 1996). Similarly, the southern region of West Asia also
witnessed severe epidemics of yellow rust in the past as the yield losses remained
10-50 percent in Yemen during 1991-96 (Bahamish et al., 1997).
Stripe rust was a dominant disease in Central Asian Muslim states of Kazakhstan,
Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan in the late 1990s and early
2000s, accounting for yield losses of 20 – 40 percent in 1999 and 2000 (Morgounov
et al., 2004). During the last decade, several yellow rust epidemics in most of the
wheat-growing areas of Iran caused over 30 percent crop loss which was estimated to
119
be 1.5 and 1.0 million tons of grain losses in 1993 and 1995, respectively (Torabi et
al., 1995).
In the Cukurova area of Turkey, a loss of over 0.5 million tons was recorded due to
epidemics of yellow rust on the cultivar “Seri 82” (Dusunceli et al., 1996). An
epidemic of stripe rust occurred in China in 2002 and affected about 6.6 million
hectares of wheat in 11 provinces and caused a yield loss of 1.3 million tons (Wan et
al., 2004).
Grant et al. (2005) reported from Australia that in Victoria, stripe rust can reduce
yield by up to 50 percent and leaf rust by greater than 20 percent in susceptible
varieties. Colson and Wilkinson (2006) also witnessed this disease throughout the
wheat growing regions of Queensland in 2005.
Rust scenario in Pakistan revealed that the pathogen exhibits its severity mainly in
the country’s foothills, northern areas and upland of Baluchistan, which can
jeopardize wheat production when it develops in an epidemic form, as occurred in
1992-93. Out of the total 8.35 million hectares of wheat production area in Pakistan,
70 percent is prone to stripe rust, which encompasses an area of about 5.8 million
hectares (Singh et al., 2004b). In Pakistan, the rust epidemics have occurred in 1947-
48, 1953-54, 1958-59 and 1977-78. Mild epidemic of stripe and leaf rusts also
occurred in 1972-73 and 1975-76.
Pathogenic activity of rusts can either be curtailed by resistance varieties or by use of
chemicals. Use of fungicide is more effective when rust disease is identified on
120
susceptible varieties early in the growing season but is a least employed method of
stripe rust management in the northern Punjab and NWFP. Although, the
recommended wheat yield potential of a variety would be 1,680-1,866 and 2613-
2799 Kg ha-1 for rain-fed and irrigated areas, respectively in order to justify fungicide
treatment (Anonymous, 2009) but the use of fungicide is merely a dream as majority
of the resource poor farmers’ in the barani area could only attain an average yield of
1,755 Kg ha-1 (Anonymous, 2008) due to rain-fed agriculture. Stripe rust of wheat is
presently being managed by cultivation of resistant varieties, which at present is the
only viable option for the farmers.
Hassan et al. (1979) accounted 10.1 percent yield loss to the tune of 0.83 million
tons valuing US$86 million during 1977-78 due to leaf rust. Likewise, Ahmad et al.
(1991) reported a revenue loss attributing to stripe rust valuing US$8 million in just
three districts of Baluchistan.
Khan and Mumtaz (2004) also witnessed rust epidemic appearance during 1995 on
wheat varieties Pak-81, Pirsabak-85 and also on Inquilab-91 during 2003. A loss of
Rs. 2 billion in Pakistan was intimated by Hussain et al. (2004) during the year 1997
and 1998 due to progressive increase in virulence of pathotypes attached to cultivars
possessing YrA (Bahawalpur-79, Chenab-79, Nuri-70), YrA and Yr6 (LU-26,
Lyallpur-73, Pari-73, Sandal-73, Yecora etc.) and Yr22 (Blue Silver, Sonalika, WL-
711). Eventually, heavy losses inflicted by wheat rust in the past have upset the
economy of the country thereby, giving a serious jolt to the planners.
121
Chapter III
3.3 MATERIALS AND METHODS
The present investigations were executed at the experimental area of the Pir Mehr
Ali Shah Arid Agriculture University (PMAS-AAU) located in Rawalpindi at
geographical location (33o 38’50.68”N, 73o 04’57.82”E) and average annual rainfall
1,000 – 1,200 mm/annum Fig.13.
Fig. 13 Global location of experimental site in the PMAS-Arid Agriculture
University, Rawalpindi
122
The studies were accomplished during the years 2005-06 and 2006-07 to ascertain
wheat yield losses in terms of 1,000 grain weight and yield of wheat in Kg ha-1
inflicted by the stripe rust.
3.3.1 Soil Analysis
The composite soil samples from the experimental area of the PMAS-AAUR were
collected before sowing during each year and were analyzed for their physical and
chemical characteristics as indicated in Table 22.
Table 22 Physical and Chemical Properties of the Experimental Site
Location Clay (%)
Silt (%)
Sand (%)
Bulk density (mg m-3) pH
EC (dSm-1)
Organic matter
(%)
PMAS-AAUR (2005)
15 14 71 1.45 7.7 0.25 0.71
PMAS-AAUR (2006)
15 14 71 1.40 7.7 0.26 0.73
Field was kept summer fallow before wheat plantation during both the experimental
years. Soil was prepared by plowing the experimental area with tractor mounted
cultivator for four times followed by a planking operation. Same site was used
during both the experimentation years.
3.3.2 Experimental Layout
Wheat-fallow-wheat was the cropping pattern adopted at the experimental area since
2000-01. Wheat seeds of four varieties viz. Bakhtawar, Inquilab-91, Wafaq-2001 and
123
Morocco (universal susceptible cultivar) collected from Crop Disease Research
Program (CDRP), National Agricultural Research Centre (NARC), Islamabad were
sown at the experimental area by adopting Randomized Complete Block Design
(RCBD). Bakhtawar and Wafaq-2001 are the commercial varieties of barani areas
while Inquilab-91 was selected as it is most popular variety adopted by the farming
community and is cultivated in all the wheat growing areas of Pakistan. These three
varieties which now rank among Moderately Susceptible to Susceptible varieties
were resistant to stripe rust when released for commercial cultivation. Each of the
three commercial wheat varieties and the Morocco as a susceptible cultivar, was
sown in lines with drill in 6 rows where each row length was kept 5 meter by
maintaining a row to row distance of 30 cm. Eight replications were used for each
variety by using seed rate of 100 kg ha-1 in each planting. The trials were managed
with optimum nutrient application @ 100 Kg Nitrogen and 50 Kg Phosphorous per
hactare. Since there was no significant incidence of insect attack during the last five
years of wheat cultivation in and around the experimental area, therefore, insecticide
was not used while weed and hoeing were practiced manually during both the
experimentation years.
3.3.3 Trials Conducted under the Research Study
The entire trial was subdivided into two experiments. In experiment-1, the stripe rust
epidemic was initiated by inoculating plants of all varieties with equal doses from the
bulk inoculum of urediniospores of Pst collected at CDRP, NARC, Murree
124
possessing virulence for stripe rust resistance genes Yr1, Y6, Y6+2, Yr7, Yr9, Yr17,
Yr18, Yr24, YrSu, YrSk and YrA. Artificial inoculation was executed with this bulk
inoculum containing 250 mg uderiniospores suspended in one litre of distilled water
having 2 drops of Tween-20 as dispersant by spraying with pressure hand sprayer to
non protected plots after sun set as described by Roelfs et al. (1992). In experiment-
2, wheat plots were sprayed with excellent stripe rust control providing fungicide –
Triadimefon (Murray et al. 2005) registered under trade name Bayleton 125 EC,
Bayer Crop Science Pvt. Ltd. @ 500 ml ha-1 in a volume of 200 litres water per
hactare (Bayer, 2007) to maintain disease free wheat plants enabling us to compare
wheat yield in diseased and disease free experimental units. The fungicide spray was
applied with knap sac sprayer with an aim to protect green area of flag and flag-1
leaves (Bedggood and Hollaway, 2007) to minimize losses from stripe rust (Murray
et al., 2005). Spray droplets were of fine to medium range thereby ensuring
fungicide penetration in the wheat crop canopy. The triadimefon after application is
absorbed rapidly into the plant, become systemic but move only in the direction of
water flow. Within the wheat plant, the fungicide move to the leaf tips but do not
move from lower to upper leaves or the head, therefore, portions of leaves, whole
leaves and heads that develop after spraying are not protected (Murray et al., 2005).
Accordingly, triadimefon was applied four times between emergence of flag leaf to
fully emergence of heads (Hunger and Edwards, 2009) during 2006 and 2007 on
various dates but the first application was always made on the 1st February of both
the experimentations years keeping in view the past experience of weather conditions
125
that prevailed in the previous years. Second, third and forth fungicidal applications
were made on 16th February, 3rd March and 21st March during 2006 while on 15th
February, 5th March and 22nd March during 2007.
3.3.4 Observation of Rust Severity
The observations on stripe rust severity were recorded as percentage of leaf aea
covered with rust according to the modified Cobb’s scale (Peterson et al., 1948) from
February till end of March in non-protected plots at approximately seven days
interval after appearance of the first symptoms. The final disease severity in
percentage at non-protected plots for 2005-06 and 2006-07 after taking mean of three
reading in each replication is given at Appendix XI and XII, respectively. As stripe
rust severity is determined by visual observation, therefore, reading cannot be
absolutely perfect (CDRP, 2004). When the severity was below 5 percent, the
intervals used were trace (T) to 2 while 5 percent intervals were used for higher
readings. The host response to infection was recorded by the letters O, R, RMR, MR,
MR-MS, MS, MS-S and S as mentioned at Table 3 (Fig. 3).
3.3.5 Yield Parameters
To draw a precise conclusion for yield assessment, all the threshing operation after
harvesting was executed manually. The wheat grain yield pertaining to following
two parameters was observed:
126
3.3.5.1 Thousand grain weight (gms)
Wheat spikes for each replicate were thrashed separately and after counting 1,000
grains, were weighed by using a digital analytical balance. Yield loss assessment was
made by comparing 1,000 kernel weight and yield difference for each variety in
diseased and disease free plots.
3.3.5.2 Grain yield (kg ha-1)
Grain yield was worked out initially by randomly throwing standard (1m2) quadrate
in each replicate of the varietal plot and harvesting all the plants within the quadrate.
Wheat bundles of each replicate were sun-dried and then thrashed separately. The
grain weight of each replicate was recorded in kg and subsequently the data was
transformed into kg ha-1.
3.3.6 Statistical Analyses
The data was statistically computed by MSTAT-C program (MSTATC, 1991) and
MS-Excel program was availed for regression and correlation analysis. The
correlation and regression values have been worked out on the average values for
disease severity and yield in respect of each variety for both years and the regression
coefficient (r2) was obtained by squaring the correlation values. The comparison of
individual means was accomplished by Least Significant Difference (LSD) at 0.01
percent level of probability / significance according to Steel et al. (1997).
127
Chapter III
3.4 RESULTS & DISCUSSION
3.4.1 Effect of Environmental Conditions in Expressing Stripe Rust
The effect of three environmental conditions viz-a-viz rainfall, temperature and
humidity (Appendix VII-X) on the development of stripe rust was observed in field
conditions. Weekly record of stripe rust severity (percentage) at the non-protected
plots was maintained from February to March of both the experimentation years.
Data pertaining to maximum disease severity, after taking mean of three readings in
each replication for all the four varieties under trial, given at Appendix XI and XII is
plotted in Figure 14a and 14b for the year 2006 and 2007, respectively.
3.4.1.1 Development of wheat stripe rust in experimental area during 2005-06
To inhibit the incidence of stripe rust, control field plots in the experimental area
were sprayed with Bayleton 125EC in the first week of February 2006. Stripe rust
was not detected in the experimental plots till the third week of February 2006 as
evident from Fig 14a. Stripe rust on wheat was first noticed on 20th February in the
Morocco, followed by Inquilab-91 on 22nd February 2006. During the third week of
February to the mid of second week of March, the stripe rust severity was at low
levels in fields (Fig. 14a) even after inoculation of fresh urediniospores. Stripe rust
was observed to be scattered throughout the inoculated diseased plots in low
incidence of around 5 percent. The disease took a boost from the second week of
127
128
Fig. 14 Maximum Disease Expression by Commercial Varieties during 2006 (a)
and 2007 (b)
0.0
10.0
20.0
30.0
40.0
50.0
60.0
70.0
80.0
90.0
Feb.
1st
2nd 3rd 4th
Mar.
1st
2nd 3rd 4th
Apr. 1
st 2nd
Months / Weeks during 2005-06 a
Dis
ease
Sev
erit
y (%
)
Wafaq 2001 Inquilab 91 Bakhtawar Morocco
0.0
10.0
20.0
30.0
40.0
50.0
60.0
70.0
80.0
90.0
Feb.
1st
2nd 3rd 4th
Mar
. 1st 2n
d 3rd 4th
Apr. 1s
t2n
d
Months / Weeks during 2006-07 b
Dis
ease
Se
verit
y (%
)
129
March 2006 (Fig. 14a) as the pustules on infected plants was seen sporulating
profusely during prevalence of high humid condition (Fig. 15 and 16) with low
temperature (Fig. 17) favouring disease development. After the mid of March, wheat
stripe rust infected leaves which were exhibiting not more than 30 percent disease
severity, expressed up to 60 percent on Morocco (Fig. 14a).
During the last week of March 2006, wheat stripe rust was turned to low levels on
the flag leaves in disease plots (Fig. 14a). The dry (Fig. 15) and hot (Fig. 17)
conditions after last week of March 2006 obstructed further development of stripe
rust in experimental fields and accordingly, in the first week of April, no stripe rust
was observed in diseased plots (Fig. 14a). This was the time when the infectious
stage (Uderiniospores) turned into resting stage (Teliospores). The flag leaves which
were exhibiting yellowish color in stripes turned in to black.
To inhibit stripe rust development in the disease free plots, Bayleton 125EC was
applied for second, third and fourth time in the 3rd week of February, 1st and 3rd week
of March 2006, respectively.
Although rainfall during the last days of second week in March 2006 (Fig. 15)
provided high moisture conditions but the warmer day and night temperatures (Fig.
17) restricted further stripe rust development (Long et al., 2004) at the end of March
2006. Accordingly, in early April, stripe rust exhibited very light reactions in the
diseased plots (Fig. 14a). During the ideal period for stripe rust expression in
inoculated plots, up to 60 percent severities on susceptible cultivar – Morocco was
recorded in one out of the eight replications.
130
0
20
40
60
80
100
120
140
160
Jan Feb March April May
Months
Rai
nfal
l (m
m)
2006 2007
Fig. 15 Distribution of Rainfall in Rawalpindi dur ing 2006 and 2007
0
10
20
30
40
50
60
70
80
Jan Feb Mar Apr May
Months
Hu
mid
ity
(%)
2006 2007
Fig. 16 Prevalence of Humid Conditions during 2006 and 2007
131
0.00
5.00
10.00
15.00
20.00
25.00
30.00
35.00
Jan Feb March April May
Months
Mea
n T
empe
ratu
re (C
elci
us)
Min
Max
Fig. 17 Minimum and Maximum Temperature Regime at Rawalpindi during 2006
0.00
5.00
10.00
15.00
20.00
25.00
30.00
35.00
Jan Feb March April May
Months
Mea
n Tem
pera
ture
(C
elci
us)
Min
Max
Fig. 18 Minimum and Maximum Temperature Regime at Rawalpindi during 2007
132
Maximum stripe rust severity was observed in the second fortnight of March 2006
(Fig. 14a) causing the real yield damaging effect of the stripe rust, which copiously
reduced the photosynthetic area of the flag leaves. The effect on grain yield loss
would have been enormous provided the conducive environmental conditions
prevailed for prolonged period (Murray et al., 2007).
3.4.1.2 Expression of wheat stripe rust in experimental area during 2006-07
Since the start of February 2007, all the replications in control plots were sprayed
with fungicide to maintain rust free crop. Neither establishment nor development of
wheat stripe rust was noticed till the second week of February. Since, the conditions
seemed appropriate for disease establishment in February due to high rain fall (Fig.
15), increased humidity (Fig. 16) and low temperature (Fig. 18), Bayleton 125EC
was applied for the second time at the start of 2nd fortnight of February. On the other
hand, in un-sprayed plots, stripe rust appeared in traces during second week of
February 2007 (Fig. 14b). The disease appeared in substantial amounts in non-
protected plots from forth week of February 2007 and by that time, 50, 20, 25 and 10
percent stripe rust severities were expressed by Morocco, Inquilab-91, Wafaq-2001
and Bakhtawar, respectively (Fig. 14b).
Disease control plots were sprayed for the third time in the first week of March 2007
to maintain disease-free. In the mid - March, wheat stripe rust was much intense,
severe and extensive than last year in all the plots inoculated with the rust pathogen.
The increased disease severity was attributed to high rainfall (Fig. 15), increased
133
humidity (Fig. 16) with lower temperature (Fig. 18). Murray et al. (2005) suggested
that spraying is effective before the stripe rust approaches five percent in the flag leaf
area which becomes difficult to manage if exceeds this level.
The disease lasted till the first week of April 2007 (Fig. 14b) as the dry and
inauspicious environmental conditions impeded further development of stripe rust.
After first week of April, hot temperatures (Fig. 18) coupled with reduced rainfall
(Fig. 15) and low humid conditions (Fig. 16), arrested stripe rust development on all
the four wheat varieties under yield loss trial. Before the mid of April, stripe rust
development had ceased in the disease plots (Fig. 14b) due to non-conducive high
temperatures at night for stripe rust amplification (Long et al., 2004).
The appearance of stripe rust was a bit earlier in 2007 (Fig. 14b) as compared to
2006 (Fig. 14a), due to humid weather in the early February 2007 (Fig. 15 and 16)
that enhanced infection in the presence of low temperature (Fig. 18). The stripe rust
infections were not only on the top leaves but the lower leaves also expressed
enormous disease load. In early March, wheat stripe rust was observed to be
developing very rapidly in replicated lines of susceptible varieties. Some replications
in diseased plots had incidence levels of 80 percent stripe rust and severity levels of
30 percent or more.
The cultivars Morocco and Inquilab-91 exhibited more susceptibility to stripe rust
infections as over 50 and 30 percent, respectively (Fig. 14b) in all the eight
replications for each variety. The other two commercially grown wheat varieties
134
expressed moderate susceptibility to stripe rust from 20 to 35 percent. The cool
temperatures in the presence of high humidity levels were found conducive for stripe
rust development in both the wheat seasons and the study is in line with the
observations made by Long et al. (2004) that low temperatures with high moisture
levels were favorable for development of stripe rust in spring wheat fields in north
central region of Minnesota.
In 2005-06, yield losses from stripe rust were considerably less than in 2006-07
because of the late development of heavy rust infections. One concern in 2007 was
that new rust races might have developed which were more virulent and aggressive
as compared to those of 2006. Another possibility is that stripe rust over summering
might have increased with greater survival of stripe rust population, which appeared
in higher frequency during next season. Beard et al. (2005) suggested that the
amount of rust prevalence in a given year is important in determining the risk of
stripe rust as there is more chance of carryover to the next season. Some cultivars
may have the potential to withstand rust losses due to the presence of high
temperature adult plant resistance (Long et al., 2004).
The disease might be favored by moist conditions that prevailed during 2007 with
increased rainfall (Fig. 15), humidity (Fig. 16) and lower temperature (Fig. 18) as
compared to the environmental conditions persisted during 2006 (Fig. 15, 16 and
17). Watkins (2005) also concluded greater risk for stripe rust when there are above
average rainfalls and reduced chances of disease under dry conditions.
135
3.4.2 Disease Severity Recorded in Wheat Varieties during 2005-06 & 2006-07
The data pertaining to expression of disease severity during 2005-06 and 2006-07
was subjected to Analysis of Variance (Table 23 and 24). The results of analysis of
variance depicted highly significant differences among different varieties under
trials.
Table 23 ANOVA Expressing Response of Wheat Varieties to Incidence of
Stripe Rust during 2005-06
Source of variance df SS MS F-value P
Wheat varieties 3 9733.59 3244.53 555.50 0.000
Replicate 7 117.97 16.85 2.89 0.028
Error 21 122.66 5.84
Total 31 9974.22
Table 24 ANOVA Expressing Response of Wheat Varieties to Incidence of
Stripe Rust during 2006-07
Source of variance df SS MS F-value P
Wheat varieties 3 10706.3 3568.8 123.62 0.000
Replicate 7 225.0 32.1 1.11 0.391
Error 21 606.3 28.9
Total 31 11537.5
The mean values regarding disease severity in wheat varieties corresponding to the
crop years 2005-06 and 2006-07 compared by Least Significant Difference (LSD)
Test at α = 0.01 are presented in Table 25.
136
Table 25 Comparison of mean values for disease severity and percent increase
in disease severity
Varieties Disease Severity (%)
2005-06 2006-07 Pooled Average % Increase*
Bakhtawar 13.12 bc 21.88 c 17.50 c 66.77
Inquilab-91 8.12 c 36.25 b 26.25 b 346.43
Wafaq-2001 25.00 b 31.88 b 28.44 b 27.52
Morocco 53.12 a 57.50 a 55.31 a 8.25
* Disease severity 2nd year – Disease severity base year (1st year)
x 100 Disease severity base year
It is evident from Table 25 that universally susceptible variety Morocco exhibited
maximum vulnerability to stripe rust by showing the disease severity as 53.12 and
57.50 percent during 2005-06 and 2006-07, respectively. The other three commercial
varieties vis-à-vis Wafaq-2001, Bakhtawar and Inquilab-91 followed Morocco with
25, 13.12 and 8.12 percent as well as 31.88, 21.88 and 36.25 percent disease
severity, for the years 2005-06 and 2006-07 (Table 25). All the varieties responded
with significant differences in expressing towards disease severity during both the
experimental years. Table 25 further elucidates increase in stripe rust severity in all
the varieties whereby Inquilab-91 exhibited maximum disease severity as 346.43
percent during 2006-07 over the last year (2005-06). Similarly, there was an increase
of 66.77 and 27.52 percent disease severity expressed by Bakhtawar and Wafaq-
2001, respectively during 2006-07. Favourable environmental conditions for stripe
rust during 2006-07 also enhanced the disease severity in Morocco to 8.25 percent.
137
The figures pertaining to comparison of mean values for disease level tabulated in
Table 25 describe that disease severity response of all the varieties tends towards
higher side during the second year of experimentation. The abrupt increase in the
disease severity of Inquilab-91 from 8.12 percent in 2005-06 to 36.25 percent in
2006-07 cautions the growers to avoid monoculture of a single variety on a large
scale in the wheat growing areas of the country. The reason may be attributed
towards increased virulence of the P. striiformis tritici that coupled with the presence
of favorable environmental conditions like increased rainfall (Fig. 15) and high
humid conditions (Fig. 16) with low temperature (Fig. 18) during the second year.
The invariable change in the race structure of the stripe rust pathogen besides
recurrent appearance of new races can overcome genetic resistance thereby rendering
ineffectiveness of the resistant cultivars (Kequan Xi et. al., 2007).
The results mentioned in Table 25 are the indication of the build up of disease
pressure (pathogen population size) under favourable environmental conditions
because of the year after year cultivation of same varieties with narrow genetic base,
which has made the stripe rust pathogen aggressive against presently cultivated
commercial varieties. It is also evident that during 2006-07, Wafaq-2001 and
Bakhtawar depicted higher severity to stripe rust (Fig. 14b) as compared to the
previous year (Fig. 14a). No doubt, the resistance influences individual crop risk but
the response of all the varieties towards serious stripe rust outbreak during 2006-07
may be attributed to favorable weather factors (Beard et al., 2005) that prevailed for
138
a longer period of time. The phenomenon may be credited to more rainfall (Fig. 15)
during 2006-07 in comparison with 2005-06. The results are in line with the previous
work, which indicate that disease severity is strongly affected by meteorological
factors such as temperature, rainfall and humidity. High rainfall coupled with
increased humid conditions under the prevalence of optimum temperature regime
favored early disease expression as the stripe rust spores require moisture to infect
leaves (Beard et al., 2005). The epidemic continued pathogen development till
decreased humidity (Fig. 16) and rise in night temperature (Fig. 17 and 18) during
April.
3.4.3 Performance of Wheat Varieties against Disease Severities on 1,000
Grain Weight during 2005-06 & 2006-07
The data pertaining to exposure of wheat varieties for 1,000 grain weight against
disease severities during 2005-06 and 2006-07 were subjected to Analysis of
Variance (Table 26 and 27). Results of analysis of variance depicted highly
significant differences among 1,000 grain weight of different varieties in response to
the stripe rust attack, however, no significant effect among blocks/replicates was
observed during 2005-06. Almost identical results were observed during 2006-07
except the effect of blocks/replicates was found significant, which may be attributed
to the effect of more rainfall, low temperature, and high humidity thereby resulting in
more disease pressure.
139
Table 26 ANOVA Exhibiting Effect of Disease Severities on 1,000 Grain
Weight of Different Wheat Varieties (2005-06)
Source of variance df SS MS F-value P
Replications 7 15.51 2.215 1.52 0.100
Wheat varieties 3 1038.83 346.28 229.43 0.000
Disease level 1 4251.20 4251.20 2816.74 0.000
Variety * Disease level 3 796.56 265.52 175.93 0.000
Error 49 69.01 1.40
Total 63 6171.11
Table 27 ANOVA Exhibiting Effect of Disease Severities on 1,000 Grain
Weight of Different Wheat Varieties (2006-07)
Source of variance df SS MS F-value P
Replications 7 90.71 12.95 3.23 0.0025
Wheat varieties 3 582.2 194.1 37.86 0.000
Disease level 1 5651.3 5651.3 1102.42 0.000
Variety * Disease level 3 830.6 276.9 54.01 0.000
Error 49 196.38 4.0
Total 63 7351.1
The mean values regarding expression of wheat varieties in terms of 1,000 grain
weight against disease severities corresponding to the year 2005-06 and 2006-07
compared by Least Significant Difference (LSD) Test at α = 0.01 are presented in
Table 28.
140
Table 28 Comparison of Mean Values for 1,000 Grain Weight in Diseased Plots
Varieties Mean values of 1,000 grain weight
2005-06 2006-07 Combined Average
Bakhtawar 32.54 a 23.55 b 28.05 b
Inquilab 91 34.57 a 33.61 a 34.09 a
Wafaq-2001 27.76 a 24.61 b 26.19 b
Morocco 15.35 b 15.20 c 15.28 c
Comparison of mean values in Table 28 revealed that a non-significant difference
was observed in 1,000 grain weight of the three commercial varieties except
Morocco during 2005-06. Some significant differences per 1,000 grain weight were,
however, found among the wheat varieties during the 2nd year of yield loss trials. The
expression of wheat stripe rust severity varied with the variety, which resultantly
yielded 1,000 grain weight in all the varieties used in experimentation during 2005-
06 and 2006-07. Perusal of means indicated that universally susceptible variety
yielded minimum 1,000 grain weight with 15.35 gms and 15.20 gms during 2005-06
and 2006-07, respectively. The other three commercial varieties vis-à-vis Wafaq-
2001, Bakhtawar and Inquilab-91 followed Morocco with 27.76 gms, 32.54 gms and
34.57 gms as well as 24.61 gms, 23.55 gms and 33.61 gms per 1,000 grain weight,
for the years 2005-06 and 2006-07 in that order.
The Table 28 envisages that reduction of 8.99 gms per 1,000 grain weight was
recorded in Bakhtawar during 2006-07 due to expression of almost double disease
severity than the preceding year. The reduced 3.15 gms per 1,000 grain weight of
141
Wafaq-2001 was calculated during the year 2006-07 in comparison with that of the
1,000 grain weight during 2005-06. The 1,000 grain weight for Inquilab-91 and
Morocco remained almost same with a very minor difference of 0.91 gms and 0.15
gms, respectively from the previous year. The results of the study are in line with the
work done by Smith et al. (1986) that correlated increased disease severity with
lesser grain weight.
3.4.4 Expression of Wheat Varieties for 1,000 Grain Weight in Control during
2005-06 & 2006-07
The data pertaining to exposure of wheat varieties for 1,000 grain weight in the
control plots with zero percent disease during 2005-06 and 2006-07 was subjected to
Analysis of Variance (Table 29 and 30). Results of analysis of variance depicted
highly significant differences among 1,000 grain weight of different varieties.
Table 29 ANOVA Showing Varying Genetic Potential of Wheat Varieties
in 1,000 Grain Weight in Control during 2005-06
Source of variance df SS MS F-value P
Wheat varieties 3 51.173 17.058 22.22 0.000
Replicate 7 1.906 0.272 0.35 0.918
Error 21 16.119 0.768
Total 31 69.198
142
Table 30 ANOVA showing varying genetic potential of wheat varieties in
1,000 grain weight in control during 2006-07
Source of variance df SS MS F-value P
Wheat varieties 3 51.32 17.11 4.81 0.011
Replication 7 61.58 8.80 2.47 0.051
Error 21 74.71 3.56
Total 31 187.62
The mean values regarding expression of wheat varieties in terms of 1,000 grain
weight in control against zero percent stripe rust pertaining to the year 2005-06 and
2006-07 compared by Least Significant Difference (LSD) Test at α = 0.01 are
presented in Table 31.
Table 31 Comparison of Mean Values for 1,000 Grain Weight in Control
Varieties Mean values of 1,000 grain weight
2005-06 2006-07 Combined Average
Bakhtawar 42.76 a 42.53 ab 42.65 a
Inquilab-91 44.49 a 41.45 b 42.97 a
Wafaq-2001 45.61 a 44.98 a 45.30 a
Morocco 42.56 a 43.23 ab 42.90 a
143
Comparison of mean values in Table 31 revealed that all the varieties expressed non-
significant differences in 1,000 grain weight during the year 2005-06, however,
offered some significant difference during 2006-07. Wafaq-2001 articulated
maximum 1,000 grain weight as 45.61 gms and 44.98 gms followed by Inquilab-91
with 44.49 gms and 41.45 gms during both the experimental years. The cultivars
Morocco and Bakhtawar exhibited 42.56 gms, 43.23 gms and 42.76 gms, 42.53 gms
per 1,000 grain weight for the year 2005-06 and 2006-07, respectively.
The Table 31 reflected that all the varieties except Inquilab 91 responded with almost
similar 1,000 grain weight. All the three commercial varieties exhibited maximum
1,000 kernel weight during 2005-06 but the universally susceptible variety Morocco
performed contrarily by expressing 0.67 gms more weight during 2006-07. Inquilab-
91 reacted with 3.04 gms lower weight for 1,000 wheat grains during 2006-07.
Assessment of the pooled average for all the four varieties for the year 2005-06 and
2006-07 envisages that Wafaq-2001 responded with maximum 1,000 grain weight as
45.30 gms followed by Inquilab-91, Morocco and Bakhtawar with 42.97 gms, 42.90
gms and 42.65 gms, respectively. The results of the study are in line with the
calculation based on 232 samples made by Khan (2003) for assessment of 1,000
kernel weight of different wheat varieties in the Punjab, the values of which ranged
between 26.4 – 56.2 gms during 1999 to 2001. It was further suggested that 1,000
kernel weights of wheat varieties in the Northern Punjab varied from 39.2 – 49.8
gms under favorable environmental conditions for the plant growth.
144
Table 32 ANOVA showing impact of stripe rust on 1,000 grain weight of
different wheat varieties during 2005-06 and 2006-07
Source of variance df SS MS F-value P
Replications 7 79.2 11.3 0.36 0.923
Wheat varieties 3 10197.5 3399.2 355.18 0.000
Years 1 395.5 395.5 41.33 0.000
Disease level 1 25736.1 25736.1 2689.16 0.000
Varieties * Years 3 22.5 7.5 0.78 0.506
Varieties * Disease level 3 10197.5 3399.2 355.18 0.000
Years * Disease level 3 395.5 395.5 41.33 0.000
Varieties * Years * Disease level 3 22.5 7.5 0.78 0.506
Error 105 992.7 9.45
Total 127 48038.9
ANOVA results in Table 32 showed that there were significant differences among
1,000 grain weight of different wheat varieties. The cropping years and disease level
had also significant impact on 1,000 grain weight of different wheat varieties sown
in the irrigated and rain-fed areas of Pakistan. Similarly, interaction of disease level
with the varieties as well as with years was also found statistically significant. The
interaction between varieties with years as well as varieties with years and disease
level were, however, found statistically non-significant.
The average 1,000 kernel weight of wheat obtained from eight replications
pertaining to research study carried out for two consecutive years under disease free
145
situation envisages that the same are in line with the yield potential of the
commercial varieties. The proposal for approval of wheat variety Inquilab-91 made
by WRI, Faisalabad (1991) predicted the yield potential of this variety as 44.4 gms
while the average 1,000 kernel weight of the variety ranged 41.45-44.49 gms.
Likewise, genetic potential for 1,000 kernel weight of Wafaq 2001 and Bakhtawar
was reported as 40 gms (NARC, 2001) and 46 gms (NIFA, 1992), respectively while
the observed 1,000 kernel weight ranged between 44.98 – 45.61 gms and 42.53 –
42.76 for these varieties in that order.
3.4.5 Response of Wheat Varieties against Disease Severities in Yield (Kg ha-1)
during 2005-06 & 2006-07
Data regarding impact on yield in Kg ha-1 when wheat varieties were exposed against
disease severities during 2005-06 and 2006-07 were subjected to Analysis of
Variance (Table 33 and 34). The ANOVA results depicted highly significant
differences among yield in Kg ha-1 of different varieties to the stripe rust under trials.
Table 33 ANOVA Showing Response of Wheat Varieties against Disease
Severities in Yield (Kg ha-1) during 2005-06
Source of variance df SS MS F-value P
Wheat varieties 3 7110848 2370283 1238.50 0.000
Replication 7 21510 3073 1.61 0.189
Error 21 40191 1914
Total 31 7172549
146
Table 34 ANOVA Showing Response of Wheat Varieties against Disease
Severities in Yield (Kg ha-1) during 2006-07
Source of variance df SS MS F-value P
Wheat varieties 3 7647915 2549305 1835.12 0.000
Replication 7 15221 2174 1.57 0.200
Error 21 29173 1389
Total 31 7692309
The mean values attributing to expression of wheat varieties in terms of yield in Kg
ha-1 against disease severities corresponding to the year 2005-06 and 2006-07
compared by Least Significant Difference (LSD) Test at α = 0.01 are presented in
Table 35.
Table 35 Comparison of Mean Values for Yield (kg ha-1) in Diseased Plots
Varieties Mean values of yield Kg ha-1
2005-06 2006-07 Combined Average
Bakhtawar 1,640.7 b 1,187.41 c 1,414.06 b
Inquilab 91 2,336.7 a 2,271.81 a 2,304.26 a
Wafaq-2001 1,418.0 b 1,257.09 b 1,337.55 b
Morocco 1,041.7 c 1,031.52 d 1,036.61 c
Comparison of mean values in Table 35 revealed that there are significant
differences among yield of all the wheat varieties used in the trials during 2005-06
and 2006-07. Assessment of means indicated that universally susceptible Morocco
responded with the lowest yield with 1,041.7 kg ha-1 and 1,031.52 kg ha-1 during
2005-06 and 2006-07, respectively. The results are in line with the findings of Beard
147
et al. (2005) as the yield losses in susceptible varieties will be the highest especially
when early infection is there. The other three commercial varieties vis-à-vis Wafaq-
2001, Bakhtawar and Inquilab-91 showed higher yield as 1,418 kg ha-1, 1,640.7 kg
ha-1 and 2,336.7 kg ha-1 as well as 1,257.09 kg ha-1, 1,187.41 kg ha-1 and 2,271.81 kg
ha-1 for the years 2005-06 and 2006-07, respectively.
It is evident from Table 35 that during 2006-07, Bakhtawar and Wafaq-2001
responded with significant reduction in kg ha-1 yield as compared to Morocco and
Inquilab 91. The yield for Morocco remained almost the same with a very minor
difference of 10.18 kg ha-1 from the previous year. The Table 35 also visualizes that
considerable reduction in kg ha-1 yield was accredited to Bakhtawar during 2006-07
with 453.29 kg ha-1 as compared to the previous year. Least yield loss in terms of kg
ha-1 was witnessed in case of Inquilab-91 as 64.89 kg ha-1, which indicates that
although the stripe rust resistance has been broken yet the variety has potential to
express substantial yield even with greater degree of disease severity.
The combined averages envisaged that maximum kg ha-1 yield was expressed by
Inquilab 91 as 2,304.26 kg ha-1 followed by Bakhtawar, Wafaq 2001 and Morocco as
1,414.06 kg ha-1, 1,337.55 kg ha-1 and 1,036.61 kg ha-1. The study results are in line
with the work done by Salman et al. (2006) who correlated increased disease
severity with lesser yield. Working on similar lines, Ash and Brown (1990)
correlated total yield loss of the wheat cultivar with timing of epidemic and yield
potential of the crop.
148
The combination of near perfect weather conditions during the wheat growing season
of 2006-07, ideal for the stripe rust development and its spread besides the genetic
adaptability of the pathogen to evolve new races having capability to infect a wide
range of host plants, resulted in a significant yield loss in the susceptible varieties
(Wright, 2003).
3.4.6 Yield Response of Wheat Varieties in Control Plots during 2005-06 &
2006-07
The data pertaining to yield in kg ha-1 among different wheat varieties against zero
percent stripe rust during 2005-06 and 2006-07 was subjected to Analysis of
Variance (Table 36 and 37). The results of analysis of variance depicted highly
significant differences among kg ha-1 yield in different varieties in control against
zero percent stripe rust during the two years of experimentation.
Table 36 ANOVA Showing Yield Response of Wheat Varieties in Control
Plots during 2005-06
Source of variance Df SS MS F-value P
Wheat varieties 3 4084708 1361569 855.21 0.000
Replication 7 31780 4540 2.85 0.030
Error 21 33434 1592
Total 31 4149922
149
Table 37 ANOVA Showing Yield Response of Wheat Varieties in Control
Plots during 2006-07
Source of variance Df SS MS F-value P
Wheat varieties 3 2780600 926867 1840.65 0.000
Replication 7 3957 565 1.12 0.386
Error 21 10575 504
Total 31 2795132
The mean values concerning to expression of wheat varieties in terms of kg ha-1 yield
in control against zero percent stripe rust pertaining to the years 2005-06 and 2006-
07 compared by Least Significant Difference (LSD) Test at α = 0.01 are presented in
Table 38.
Table 38 Comparison of Mean Values for kg ha-1 Yield in Control Plots
Varieties Mean values of yield kg ha-1
2005-06 2006-07 Combined Average
Bakhtawar 1,928.00 b 1,917.62 b 1,922.81 b
Inquilab-91 2,480.00 a 2,310.54 a 2,395.27 a
Wafaq-2001 1,518.70 c 1,497.72 d 1,508.21 d
Morocco 1,730.30 bc 1,757.53 c 1,743.92 c
Perusal of mean values in Table 38 revealed that all the varieties showed
significantly different responses in terms of kg ha-1 in disease free circumstances
during 2005-06 and 2006-07. The data indicated that all the varieties expressed
significant differences in yield during both the experimentation years but minor
within a variety. The widely adopted Inquilab-91 responded with maximum yield
150
during both the years as 2,480 kg ha-1 and 2,310.54 kg ha-1. The combined average
for both the years also credited maximum yield to Inquilab-91 as 2,395.27 kg ha-1
followed by Bakhtawar, Morocco and Wafaq-2001 with 1,922.81 kg ha-1, 1,743.92
kg ha-1 and 1,508.21 kg ha-1, respectively.
The average kg ha-1 yield of wheat obtained after calculating yield from eight
replications pertaining to research study carried out for two years under disease free
situation indicated that the same are in line with the yield potential of the commercial
varieties. The proposal for approval of wheat variety Inquilab-91 made by WRI,
Faisalabad (1991) predicted the yield potential of Inquilab-91 as 6,088 kg ha-1 while
the variety under this study responded with 2,310.54 – 2,480 kg ha-1. Likewise, the
genetic potential of wheat varieties, Wafaq-2001 and Bakhtawar, that was reported
as 5,167 kg ha-1 (NARC, 2001) and 6,700 kg ha-1 (NIFA, 1992), responded with
yield that ranged between 1,497.72 – 1,518.70 kg ha-1 and 1,917.62 – 1,928 kg ha-1,
respectively in the executed yield loss study.
It was suggested that wheat plant continues robust growth for a longer time at
optimum temperature i.e., 25oC, which enhances grain fill period and resultantly
express maximum yield. The growth continues at lower temperatures (3-4oC) or at
higher temperatures (30-32oC) but lower temperatures retards the growth while
higher temperatures accelerate maturity (Khan, 2003).
During 2006-07, Bakhtawar, Inquilab-91 and Wafaq-2001 responded with slightly
lower yields as compared to the previous year trials as evident from Table 38.
151
Morocco, however, exhibited 27.23 kg ha-1 more yield during 2006-07 as compared
to the yield given by this variety during 2005-06. The unexpected increase in case of
Morocco may be attributed to the climatic factor that not only favoured disease
development but also resulted in yield increase.
The average minimum and maximum crop season temperatures prevailed in
Rawalpindi during 2006 and 2007 are shown in Fig. 19 and 20, respectively. The
straight line in yellow depicts optimum growth temperature. The minimum and
maximum temperature lines are outside the temperature limits in both the figures,
which indicate that the temperature during experimentation seasons was below the
optimum limits. At the terminal end, maximum temperature is heading above the
optimum requirements that accelerated the crop maturity without providing sufficient
time for proper grain filling. A yield difference of 10.38, 20.98 and 169.46 kg ha-1
observed during 2006-07 among Bakhtawar, Wafaq-2001 and Inquilab-91,
respectively with zero percent disease may be attributed to this environmental factor.
The gap between optimum and existed minimum-maximum temperature was found
widened during 2007 (Fig. 20) as compared to 2006 (Fig. 19). This gap might have
resulted in expression of lower yields in case of the three commercial varieties
during 2007. The studies are in accordance with the findings of Khan (2003) who
suggested that shorter growing season with lack of optimum temperature
requirement in the country is a cause for expression of lower yields as compared to
some of the best wheat yielding countries.
152
0.0
5.0
10.0
15.0
20.0
25.0
30.0
Jan. 1s
t 2n
d3rd 4th
Feb. 1
st 2n
d3r
d 4th
Mar
. 1st
2nd
3rd 4t
h
Apr. 1
st2n
d3r
d 4th
Months / Weeks
Tem
pera
ture
C
Min. Temp. 2006 Max. Temp. 2006 Optimum Temp.
Fig. 19 Weekly Average Minimum and Maximum Temperatures at Rawalpindi during 2006
0.0
5.0
10.0
15.0
20.0
25.0
30.0
Jan.
1st
2nd
3rd
4th
Feb.
1st
2nd
3rd
4th
Mar
. 1st
2nd
3rd 4t
h
Apr. 1
st2n
d3r
d 4th
Months / Weeks
Tem
pera
ture
C
Min. Temp. 2007 Max. Temp. 2007 Optimum Temp.
Fig. 20 Weekly Average Minimum and Maximum Temperatures at Rawalpindi during 2007
153
Table 39 ANOVA showing impact of stripe rust on wheat yield (kg ha-1) of
different wheat varieties during 2005-06 & 2006-07
Source of variance df SS MS F-value P
Replications 7 65163 9309 0.05 1.000
Wheat varieties 3 18810213 6270071 870.62 0.000
Years 1 325337 325337 45.17 0.000
Disease level 1 4535074 4535074 629.71 0.000
Varieties * Years 3 246700 82233 11.42 0.000
Varieties * Disease level 3 1920040 640013 88.87 0.000
Error 109 785004 7202
Total 127 26687531
ANOVA results in Table 39 showed that there were significant differences among
wheat yield (kg ha-1) of different varieties tested under in the research during both
the trial years. The cropping years and disease level also had significant impact on
wheat yield of the three commercial varieties and Morocco cultivar. Similarly,
interactions between varieties with experimentation years and with the disease level
were also found statistically significant. It was further evident from the ANOVA
table that blocks/replicates showed significant variation. The phenomenon may be
attributed to the rainfall and other climatic patterns that prevailed during both the
experimentation years.
154
3.4.7 Correlation between Disease Severity and Yield
The correlation pertaining to disease levels and yield was studied statistically during
2005-06 and 2006-07.
3.4.7.1 Correlation between disease level and yield during 2005-06
The graph indicated a strong negative correlation (-0.87003) between the disease
levels and wheat yield (Fig. 21). Similar trend was observed when correlation
statistics was performed between disease severity and wheat yield per hectare.
y = -23.516x + 2193.5
R2 = 0.7569
0
500
1000
1500
2000
2500
0 10 20 30 40 50 60
Disease Severity (%)
Yie
ld K
g/ha
Fig. 21 Graph Showing Relationship between Averages of Disease Severity &
Wheat Yield 2006
155
y = -6.2813x + 1668.6
R2 = 0.0279
0
500
1000
1500
2000
2500
0 10 20 30 40 50 60
Disease Severity (%)
Yie
ld K
g/h
a
Fig. 22 Graph Showing Relationship between Averages of Disease Severity &
Wheat Yield 2007
3.4.7.2 Correlation between disease level and yield during 2006-07
Analysis of the data pertaining to the year 2006-07 indicated a strong negative
correlation (-0.16698) between the disease level as well as 1,000 kernel weight (Fig.
22). Similar trend was obvious when correlation statistics was performed between
disease severity and wheat yield per hectare.
The reduced value of adjusted R2 is due to the unexpected response of Inquilab-91,
which showed more than 4 times disease severity as compared to the 2005-06. The
data was correlated with the environmental factors which showed that incidence of
rain was greater during 2006-07 which not only boosted the disease severity but also
helped the crop in expression of its enhanced yield potential.
156
3.4.8 Regression Analysis
The dependence of yield, expressed by different wheat varieties in the trials, under
the disease severities was studied during both the years 2005-06 and 2006-07.
3.4.8.1 Regression analysis studies during 2005-06
It is evident from the Table 40 that wheat yield was strongly dependent upon the
disease severity levels showing values of F= 907.16 at 0.000 percent level of
probability during the year 2005-06. The statistical calculations are in line with the
yield data, which has also indicated that increased level of diseases caused
proportional reduction in wheat yield and thousand grain weight.
Table 40 Regression analysis showing dependence of wheat yield on disease
level (2005-06)
A N O V A
S.O.V. df SS MS F Significance F
Regression 1 18579.77 18579.77 907.1601 0.0000
Residual 62 1269.837 20.48125
Total 63 19849.61
157
0
10
20
30
40
50
60
70
Control Disease Control Disease Control Disease Control Disease
Bakhtawar Inquilab-91 Wafaq Morocco
Disease Severity 1000 Grain Weight (grams)
Fig. 23 Comparison of 1,000 Grain Weight of Wheat Varieties in Control and
Diseased Plots during 2005-06
0
10
20
30
40
50
60
70
Control Disease Control Disease Control Disease Control Disease
Bakhtawar Inquilab-91 Wafaq Morocco
Disease Severity 1000 Grain Weight (grams)
Fig. 24 Comparison of 1,000 Grain Weight of Wheat Varieties in Control and
Diseased Plots during 2006-07
158
3.4.8.2 Regression analysis studies during 2006-07
The table 40 presents that wheat yield was strongly dependent upon the disease
severity levels showing values of F=334.52 at 0.000 % level of probability during the
year 2006-07. The statistical calculations are in line with the yield data as
proportional reduction in wheat yield and thousand grain weight are attributed to the
increased level of disease.
Table 41 Regression analysis showing dependence of wheat yield on disease
level (2006-07)
A N O V A
S.O.V. df SS MS F Significance F
Regression 1 23448.01 23448.01 334.5291 0.0000
Residual 62 4345.74 70.09259
Total 63 27793.75
Comparison of mean values in Table 42 revealed that stripe rust significantly
affected 1,000 kernel weights of all the varieties used in the experiment. Appraisal of
means indicated that maximum reduction in 1,000 grain weight was observed in case
of Morocco with 27.62 gms. The mean value of 1,000 grain weight loss pertaining to
Bakhtawar and Wafaq 2001 showed 14.60 and 19.11 gms, respectively whereas,
lowest 1,000 grain weight loss was observed in Inquilab-91 despite of exhibiting
26.25 percent of stripe rust severity.
159
Table 42: Comparison of Mean Values for Stripe Rust Severity and 1,000 Grain
Weight Loss during 2005-06 and 2006-07
* Severity was recorded as percent of rust infection as per modified Cobb’s scale
MS-S+: Moderately Susceptible-Susceptible, S++: Susceptible
The varietals response of Wafaq-2001 and Morocco against stripe rust infection
remained “Susceptible (S)” for both the experimental years while Bakhtawar
responded with “Moderately Susceptible – Susceptible (MS-S)”. Inquilab-91 showed
MS-S to S response during the year 2005-06 and 2006-07, respectively.
Table 43 indicated a comparison of wheat yield, total and percent yield loss due to
stripe rust in different varieties. It has been found that maximum yield loss of 707.31
kg ha-1 (40.56 percent) was observed in Morocco followed by Bakhtawar showing
508.75 kg ha-1 loss (26.46 percent).
Wheat
Varieties
Disease
Severity *
(%)
Host
response to
infection
1,000 Grain wt (gms)
Control
plot
Disease
plot
Kernel wt.
loss
Bakhtawar 17.50 MS-S+ 42.65 28.05 14.60
Inquilab-91 26.25 MS-S+ to S++ 42.97 34.09 8.88
Wafaq-2001 28.44 S++ 45.30 26.19 19.11
Morocco 55.31 S++ 42.90 15.28 27.62
160
Table 43 Yield reduction in different wheat varieties due to stripe rust
during 2005-06 and 2006-07
Variety Yield in control
Kg ha-1
Yield in
diseased field
Kg ha-1
Yield loss
Kg ha-1
Yield loss
%
Morocco 1,743.92 1,036.61 707.31 40.56
Bakhtawar 1,922.81 1,414.06 508.75 26.46
Wafaq-2001 1,508.21 1,337.55 170.66 11.32
Inquilab-91 2,395.27 2,304.26 91.01 3.78
The other two varieties vis-à-vis Wafaq-2001 and Inquilab-91 showed wheat loss to
the tune of 11.32 and 3.78 percent, respectively that has separately been shown year
wise in Fig. 25 and 26.
Though all the wheat varieties belong to the same Triticum aestivum L., species, yet
highly significant differences were found among wheat yield as well as disease
levels. The overall performance of wheat varieties presented in Table 42 indicated
that Inquilab-91 exhibited minimum reduction of 8.88 grams per 1,000 kernel weight
in spite of the fact that maximum disease severity was observed in this variety after
Morocco. Results have proved high yielding potential of Inquilab-91 even under
disease stress.
161
y = -11.033x + 44.36R2 = 0.8035
0 10 20 30 40 50 60
Morocco
Bakhtawar
Wafaq-2001
Inquilab-91
Disease level % Yield Loss kg/ha
Linear (Yield Loss kg/ha)
Fig. 25 Estimated yield losses in response to different levels of stripe rust 2006
y = -14.09x + 59.51R2 = 0.9377
0 10 20 30 40 50 60
Morocco
Bakhtawar
Wafaq-2001
Inquilab-91
Disease level % Yield Loss kg/ha
Linear (Yield Loss kg/ha)
Fig. 26 Estimated yield losses in response to different levels of stripe rust 2007
162
It was further evident from figure 26 that disease severity in all the varieties
increased during 2006-07 as compared to 2005-06 (Fig. 25) wherein, 53 percent
disease severity was observed in case of Morocco followed by Wafaq-2001,
Bakhtawar and Inquilab-91 with 25, 13 and 8 percent, respectively (Afzal et al.,
2007) against 57, 31, 21 and 36.25 percent recorded during 2006-07 (Afzal et al.,
2008).
The present investigations are supported with the work done by Smith et al. (1986)
who found in their investigations that stripe rust cause a 51 percent loss in grain yield
on the well watered plots from (4.9-2.1 t ha-1) and a 46 percent reduction on rainfed
plots (2.8-1.5 t ha-1). Similarly, Lee in 2003 observed that stripe rust was most severe
in the California wheat fields receiving no fungicide application. Highly susceptible
cultivars suffered 75 percent or more yield loss; susceptible cultivars, 50-70 percent
yield loss; moderately susceptible cultivars, 20-40 percent yield loss, and moderately
resistant cultivars, 5-15 percent loss. Very few cultivars maintained resistance
throughout the season. Other researchers have also conducted trials on stripe rust and
have established correlation between the disease epidemic and yield loss of wheat
crop. After conducting one of such study, Ash and Brown (1990) observed that total
yield loss depends upon the cultivar, epidemic timing and the yield potential of the
crop. The yield loss experimentation carried out during 2005-06 and 2006-07,
envisaged that occurrence of early stripe rust exert enormous effect on yield as
compared to late epidemic. The results of the present study (Table 43) reflected
40.56 percent yield loss in case of Morocco, which coincide with the findings of Ash
163
and Brown (1990) as they have reported grain yield losses up to 50 percent in
susceptible cultivars.
It has been established that out of five disease components infection efficiency (IE),
sporulation capacity (SC), lesion expansion rate (LE), latent period (LP) and
infectious period (IP), the LE, IE and SC were found to be the most important
components, however, disease was more strongly influenced by weather and initial
disease (Luo and Zeng, 1995). Salman et al. (2006) reported that yield losses
increase proportionately with the increase in severity of the disease. According to
their investigations, varieties like Morocco, WL-711, SA 75, SA 42 and Chakwal
exhibited maximum losses (52-57 percent) against the leaf rust. Some other workers
also reached the same conclusion that slow ruster varieties / lines usually suffer less
yield losses as compared to the fast rusters like Morocco etc., in which losses were as
high as 52-57 percent.
Chen (2005) concluded that a susceptible cultivar could sustain a total loss when
environmental conditions are favorable for disease. The studies are also supported by
Bowden (2006) who reported that a variety would sustain 40 percent or more losses
provided the same is susceptible and there is early infection of stripe rust. A
susceptible barley cultivar suffered 72 percent grain yield loss in the US (Marshall
and Sutton, 1995) besides the stripe rust markedly reduced malting quality (Line,
2002).
164
3.4.9 Changes in Varietal Susceptibility
The tremendously diverse nature of the rust pathogens has been well documented.
New races able to cause disease on previously resistant cultivars are readily evolved.
The changes in susceptibility of Inquilab-91 that occurred during 2007 can be
attributed to favorable conditions for disease that occurred, eventually producing an
abundant inoculum load and the opportunity for mutations to higher virulence to
develop.
3.4.10 Disease Severity and Yield Loss of Wheat Cultivars during 2005-06 and
2006-07
Data offered in Table 44 and 45 confirm huge loss estimates where disease levels
were very high (Table 25). The most severely diseased cultivars had the lowest
yields (Lee, 2003), like the universally susceptible cultivar Morocco, which has a
reasonable yield potential in the absence of stripe rust, had extremely low yields
when stripe rust was severe. The statement also holds true as far as Bakhtawar and
Wafaq-2001 were concerned but in contrast, Inquilab-91, which is now susceptible
to stripe rust, had very high yields at control sites but expressed a loss of only 7.84
gms per 1,000 grain weight (Table 44) and 38.73 kg ha-1 (Table 45) when the disease
severity was 36.25 percent (Table 25) in 2007 as compared to the loss exhibited by
the variety during 2006 with 9.92 gms per 1,000 grain weight (Table 44) and 143.3
kg ha-1 (Table 45) when the disease severity was just 8.12 percent (Table 25). The
present study reveals that Inquilab-91 exhibited better performance under high
165
disease pressure when compared with susceptible check. The results are in line with
the findings of Sajid et al. (2007) whereby Inquilab-91 responded in a similar
fashion. Stripe rust was also severe against Bakhtawar and Wafaq-2001 during 2007
at the PMAS-AAUR test sites in Rawalpindi and had a dramatic affect on yield
(Table 45).
Table 44 Comparison of 1,000 Grain Weight in Wheat Varieties due to
Stripe Rust during 2005-06 and 2006-07
Varieties
Mean values of 1,000 grain weight
2005-06 2006-07
Control
plots
Disease
plots
Yield
difference
Control
plots
Disease
plots
Yield
difference
Bakhtawar 42.76 32.54 10.22 42.53 23.55 18.98
Inquilab-91 44.49 34.57 9.92 41.45 33.61 7.84
Wafaq-2001 45.61 27.76 17.85 44.98 24.61 20.37
Morocco 42.56 15.35 27.21 43.23 15.20 28.03
Table 45 Comparison of Yield difference in Wheat Varieties due to Stripe
Rust during 2005-06 and 2006-07
Varieties
Mean values of yield kg ha-1
2005-06 2006-07
Control
plots
Disease
plots
Yield
difference
Control
plots
Disease
plots
Yield
difference
Bakhtawar 1,928.00 1,640.70 287.30 1,917.62 1,187.41 730.21
Inquilab-91 2,480.00 2,336.70 143.30 2,310.54 2,271.81 38.73
Wafaq-2001 1,518.70 1,418.00 100.70 1,497.72 1,257.09 240.63
Morocco 1,730.30 1,041.70 688.60 1,757.53 1,031.52 726.01
166
Considerable yield loss was observed in 2007 as compared to the losses estimated
during 2006. The conducive environmental factor that persisted for more than four
weeks prolonged the infectious stage of the stripe rust pathogen during 2007. Greater
yield losses were observed in 2007 as the stripe rust was developed earlier (Hollaway
and Brown, 2005) and the varieties were susceptible (Bowden, 2006).
Two factors, therefore, significantly influenced the grain yield as the loss increased
with the increase in epidemic length and decreased as the temperature mounted
during grain development. The phenomenon is in line with the studies conducted by
Murray et al. (2007). The experimental sites where fungicide – Bayleton 125 EC was
applied, yields of susceptible cultivar such as Morocco were about 688.6 – 726.01 kg
ha-1 higher than at the diseased sites, with the difference attributed to the fungicide
application.
Yield losses also depended on the varietal disease resistance and were pronounced in
susceptible varieties (Beard et al., 2005). Wide spread of disease in field caused 30
to 40 percent yield losses by shriveling the grains (Fig. 27) due to reduction of
photosynthetic leaf surface (Pollock, 2005).
167
Inquilab-91 Bakhtawar
Wafaq-2001 Morocco
Fig. 27 Shriveling of Wheat Grains due to Stripe Rust Disease Intensities:
Wheat Grains in a. Control; b. up to 20%; c. More than 20 but than
40%; d. more than 40%
a
b c
d a
b c
d
a
b
c
d
a
b c
d
168
Chapter III
3.5 CONCLUSIONS
The present study exposes that although Inquilab-91 is placed among those varieties
that have been declared susceptible to stripe rust, still exhibit better yield even under
high disease pressure as compared to the other two commercial varieties of the
rainfed areas i.e., Bakhtawar and Wafaq-2001. Inquilab-91 exhibited a loss of only
38.73 kg ha-1 when the stripe rust severity was 36.25 percent in 2007, while
Bakhtawar and Wafaq-2001 expressed 730.21 and 240.63 kg ha-1 when the disease
severity was just 21.88 and 31.88 percent, respectively. There was a considerable
increase in disease severity during 2006-07 because of built up of high inoculum
pressure under favourable environmental conditions.
The use of appropriate fungicides although controls stripe rust, also adds cost to
wheat production. Their use is a huge burden for the resource poor farming
community especially in the developing countries like Pakistan. The use of fungicide
exerts health problems for users, affect the environment adversally but also result in
the selection of fungicide – resistant strains of the pathogen.
Adoption of resistant varieties is the ever economical and appropriate measure to
minimize stripe rust losses. The producers must apply varietal complementation
concept while selecting wheat varieties, which envisages selection of varieties that
differ in disease reaction as well as parentage and maturity. Since there is an absence
168
169
of a single perfect variety, complementation permits growers to counter balance the
potential weaknesses in every variety, thereby enhancing the opportunity to stabilize
yield of the whole production system.
Keeping in view the above results, it is evident that there is a dire need to avoid rapid
ruster and susceptible varieties. The study envisages contribution of host resistance
or susceptibility when the pathogen is virulent under favorable environment, to the
progress of an epidemic as such relationships are fundamental to understand and
manage yield and crop losses. Yield losses due to stripe rust were substantial;
therefore, all wheat breeding programs entrusted under plant breeding departments
should be encouraged, appreciated as well as accounted for to continuously monitor
rust situation through Plant Pathologists. Such programs must focus towards the goal
for evolving new resistant cultivars thereby ensuring sustainable food security of the
country. The impact of disease towards loss of crop yield is undeniably a loss of
human and animal resources which must be quantified in order to make decisions
towards stripe rust prevention or control.
170
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202
Appendix I Field response of differential set and NILs to Pst at hot spots in the Northern Punjab and NWFP during 2005-06 & 2006-07
Ent. No. Lines/Varieties
Resistance genes
Stripe Rust Reaction to Yr at Trap nurseries established at
PRS (Sialkot) PMAS-AAUR
(Rwp) NARC (Islamabad) CCRI (Pirsabak) AUP (Peshawar) NIFA (Peshawar)
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07
World set
1 Chinese 166 Yr1 TR TR 70S 70S 50S 50S 50S 50S 10S 5S 60S 60S
2 Lee Yr7 5S 5S 70S 70S 80S 80S 70S 80S 40S 60S 80S 70S
3 Heines Kolben Yr6,Yr2 5S 5S 50MSS 40MSS 20S 70S 70S 60S 30S 60S 80S 70S
4 Vilmorin Yr3V 0 0 0 5MRS 0 0 5MS 0 0 0 5MS 0
5 Moro Yr10 0 0 0 0 0 0 0 0 0 0 0 0
6 Strubes Dickkopf YrSd 0 0 0 0 0 0 0 0 0 0 0 0
7 Suwon92 x Omar YrSu,Yr4 10S 10S 80S 80S 80S 80S 30MSS 30MSS 20MSS 20MS 30MSS 30MSS
8 Clement Yr9,Yr2+ 0 0 10MRMS 10MRMS 10MRMS 20MRMS 40MRMS 50MRMS 10MRMS 10MRMS 10MRMS 10MRMS
9 Triticum spelta Yr5 0 0 0 0 0 0 0 0 0 0 0 0
European set
10 Hybrid 46 Yr4+ 0 0 0 0 0 0 0 0 0 0 0 0
11 Reichersberg 42 Yr7+ 0 0 0 0 0 0 0 0 0 0 0 0
12 Heines Peko Yr6,Yr 2+ 0 0 0 0 0 0 0 0 0 0 0 0
13 Nord Desprez Yr3N 0 0 0 0 0 0 0 0 0 0 0 0
14 Compare Yr8,19 APR 0 0 0 0 0 0 0 0 0 0 0 0
15 Carstens V YrCv 0 0 0 0 0 0 0 0 0 0 0 0
16 Spalding Prolific YrSp 0 0 0 0 0 0 0 0 0 0 0 0
17 Heines VII Yr2+ 0 0 0 0 0 0 0 0 0 0 0 0
Near Isogenic Lines with Avocet background
18 Yr1/6* Avocet(s) Yr1 5S 5S 20S 30S 60S 40S 50MSS 40MSS 20S 30S 50MSS 40S
19 Yr5/6* Avocet(s) Yr5 0 0 0 0 0 0 0 0 0 0 0 0
20 Yr6/6* Avocet(s) Yr6 40S 50S 90S 90S 90S 80S 70S 70S 30S 60S 80S 60S
21 Yr7/6* Avocet(s) Yr7 20S 30S 40S 40S 30S 80S 80S 70S 30S 50S 80S 80S
22 Yr8/6* Avocet(s) Yr8 0 0 80S 80S 80S 80S 40S 40S 10S 10S 40S 50S
23 Yr9/6* Avocet(s) Yr9 40S 40S 30S 40S 40S 70S 70S 80S 30S0 30S 70S 60S
24 Yr10/6* Avocet(s) Yr10 0 0 0 0 0 0 0 0 0 0 0 0
25 Yr15/6* Avocet(s) Yr15 0 0 0 0 0 0 0 0 0 0 0 0
26 Yr17/6* Avocet(s) Yr17 5S 20S 70S 70S 50S 60S 80S 70S 20S 50S 60S 50S
27 Yr18/3* Avocet(s) Yr18 5MS 10MS 20S 30S 70S 70S 50S 60S 30S 30S 60S 70S
28 Yr24/3* Avocet(s) Yr24 0 0 10MRMS 10MRMS 5MRMS 10MRMS 5MRMS 10MRMS 0 0 0 0
29 Yr26/3* Avocet(s) Yr26 0 0 0 0 0 5RMR 10RMR 10RMR 0 0 10RMR 10RMR
30 YrSP/6* Avocet(s) YrSp 0 0 0 0 0 0 0 0 0 0 0 0
31 Yr27/3*Avocet (CX 94.19.1.1)
YrSk 0 0 40S 50S 80S 80S 40MSS 40MSS 20MS 30MS 50MSS 50MS
32 Jupateco R (Yr18) Yr18 0 0 80S 80S 60S 50S 80S 50S 30S 40S 60S 60S
33 Jupateco S - 0 0 90S 90S 90S 90S 80S 90S 20S 40S 80S 90S
34 AVOCET + YrA YrA 10S 30S 90S 90S 50S 70S 70S 80S 30S 40S 80S 80S
203
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection
Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Sialkot 06-S-4,18 07-S-25,32,38 06-S-1,5,9,13 07-S-22,28,30,34,36,37 05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 L L L L
2 2 Lee 7 H H H H
3 4 Heines Kolben 2,6 H H H H
4 8 Vilmorin 3V L L L L
5 16 Moro 10 L L L L
6 32 Strubes Dickkopf Sd,25 L L L L
7 64 Suwon92 / Omar 4,Su L L H H
8 128 Clement 2,9,25,Cle L L L L
9 256 Triticum spelta album 5 L L L L
European set
10 1 Hybrid 46 4+ L L L L
11 2 Reichersberg 42 7,25 L L L L
12 4 Heines Peko 2,6,25 L L L L
13 8 Nord Desprez 3N L L L L
14 16 Compair 8, 19 APR L L L L
15 32 Carstens V Cv L L L L
16 64 Spaldings Prolific Sp L L L L
17 128 Heines VII 2,25,HVII L L L L
PATHOTYPE α 6E0 6E0 70E0 70E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotype identified as 6E0 (from 2 and 3) and 70E0 (from 4 and 6) isolates tested during 2005-06 and 2006-07, respectively.
204
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection
Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Rawalpindi
06-R-6,12,22,25
07-R-43,46,47
06-R- 2,3, 9,14,
26,30
07-R-38,40,48,
54,67
06-R-18,21,
28,33,35
07-R-39,42,49,
51,63
05-06 06-07 05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 L L L L H H
2 2 Lee 7 H H H H H H
3 4 Heines Kolben 2,6 H H H H H H
4 8 Vilmorin 3V L L L L L L
5 16 Moro 10 L L L L L L
6 32 Strubes Dickkopf Sd,25 L L L L L L
7 64 Suwon92 / Omar 4,Su L L H H H H
8 128 Clement 2,9,25,Cle L L L L L L
9 256 Triticum spelta album 5 L L L L L L
European set
10 1 Hybrid 46 4+ L L L L L L
11 2 Reichersberg 42 7,25 L L L L L L
12 4 Heines Peko 2,6,25 L L L L L L
13 8 Nord Desprez 3N L L L L L L
14 16 Compair 8, 19 APR L L L L L L
15 32 Carstens V Cv L L L L L L
16 64 Spaldings Prolific Sp L L L L L L
17 128 Heines VII 2,25,HVII L L L L L L
PATHOTYPE α 6E0 6E0 70E0 70E0 71E0 71E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotype identified as 6E0 (from 4 and 3), 70E0 ( from 6 and 5) and 71E0 (from 5 each year) isolates tested during 2005-06 and 2006-07, respectively;
205
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Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Chakwal 06-Ck-1,6,7,10
07-Ck-13,16,17,23
06-Ck- 9,11
07-Ck-14,20,27
05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 L L L L
2 2 Lee 7 H H H H
3 4 Heines Kolben 2,6 L L H H
4 8 Vilmorin 3V L L L L
5 16 Moro 10 L L L L
6 32 Strubes Dickkopf Sd,25 L L L L
7 64 Suwon92 / Omar 4,Su L L L L
8 128 Clement 2,9,25,Cle L L L L
9 256 Triticum spelta album 5 L L L L
European set
10 1 Hybrid 46 4+ L L L L
11 2 Reichersberg 42 7,25 L L L L
12 4 Heines Peko 2,6,25 L L L L
13 8 Nord Desprez 3N L L L L
14 16 Compair 8, 19 APR L L L L
15 32 Carstens V Cv L L L L
16 64 Spaldings Prolific Sp L L L L
17 128 Heines VII 2,25,HVII L L L L
PATHOTYPE α 2E0 2E0 6E0 6E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotype identified as 2E0 (from 4 each year) and 6E0 (from 2 and 3) isolates tested during 2005-06 and 2006-07.
206
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Attock
06-A-7,9,18
07-A-20,24,38
06-A-13
07-A-22,27,34
06-A-17
07-A-26,41
06-A- 3
07-A-31,37
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 L L H H L L H H
2 2 Lee 7 H H H H H H H H
3 4 Heines Kolben 2,6 H H H H H H L L
4 8 Vilmorin 3V L L L L L L L L
5 16 Moro 10 L L L L L L L L
6 32 Strubes Dickkopf Sd,25 L L L L L L L L
7 64 Suwon92 / Omar 4,Su H H H H H H H H
8 128 Clement 2,9,25,Cle L L L L L L L L
9 256 Triticum spelta album 5 L L L L L L L L
European set
10 1 Hybrid 46 4+ L L L L L L L L
11 2 Reichersberg 42 7,25 L L L L L L L L
12 4 Heines Peko 2,6,25 L L L L L L L L
13 8 Nord Desprez 3N L L L L L L L L
14 16 Compair 8, 19 APR L L L L L H L H
15 32 Carstens V Cv L L L L L L L L
16 64 Spaldings Prolific Sp L L L L L L L L
17 128 Heines VII 2,25,HVII L L L L L L L L
PATHOTYPE α 70E0 70E0 71E0 71E0 70E0 70E16 66E0 66E16
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotype identified as 70E0 (from 4 and 3), and 71E0 (from 1 and 3) isolates tested during 2005-06 and 2006-07, respectively whereas 70E16 and 66E16 ( from 2 isolates each) tested during 2006-07 and 66E0 ( from 1) isolate tested during 2005-06.
207
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection
Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Nowshera
06-N-2,3
07-N-32,38
06-N-1,12
07-N-51,59
06-N-4,18
07-N-44,52
06-N-6,13
07-N-35,48
06-N-7,10
07-N-36,45
06-N-15,21,30
07-N-47,49,56,
60 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07
World set
1 Chinese 166 1 H H L L L L L L L L L L
2 Lee 7 H H H H H H H H H H H H
3 Heines Kolben 2,6 H H H H H H H H H H H H
4 Vilmorin 3V L L L L L L L L L L L L
5 Moro 10 L L L L L L L L L L L L
6 Strubes Dickkopf Sd,25 L L L L L L L L L L L L
7 Suwon92 / Omar 4,Su L L L L L L L L H H H H
8 Clement 2,9,25,Cle L L L L L L L L L L L L
9 Triticum spelta
album 5 L L L L L L L L L L L L
European set
10 Hybrid 46 4+ L L H H L L H H L L L L
11 Reichersberg 42 7,25 L L L L L L L L L L L L
12 Heines Peko 2,6,25 L L L L L L L L L L L L
13 Nord Desprez 3N L L L L L L L L L L L L
14 Compair 8, 19 APR H H H H H H L L H H L L
15 Carstens V Cv L L L L L L L L L L L L
16 Spaldings Prolific Sp L L L L L L L L L L L L
17 Heines VII 2,25,HVII L L L L L L L L L L L L
PATHOTYPE α 7E16 7E16 6E17 6E17 6E16 6E16 6E1 6E1 70E16 70E16 70E0 70E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotypes identified as 7E16, 6E17, 6E16, 6E1 and 70E16 ( from 2 isolates in each year) and 70E0 (from 3 and 4 isolates) tested during 2005-06 and 2006-07, respectively.
208
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection
Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Charsadda
06-Cd-4
07-Cd-23
06-Cd-6,11,12,17
07-Cd-22,28,34
06-Cd-9,15,18
07-Cd-24,30
05-06 06-07 05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 L L H H L L
2 2 Lee 7 L L H H L L
3 4 Heines Kolben 2,6 H H L L H H
4 8 Vilmorin 3V L L L L L L
5 16 Moro 10 L L L L L L
6 32 Strubes Dickkopf Sd,25 L L L L L L
7 64 Suwon92 / Omar 4,Su H H H H H H
8 128 Clement 2,9,25,Cle L L L L L L
9 256 Triticum spelta album 5 L L L L L L
European set
10 1 Hybrid 46 4+ L L L L L L
11 2 Reichersberg 42 7,25 L L L L L L
12 4 Heines Peko 2,6,25 L L L L L L
13 8 Nord Desprez 3N L L L L L L
14 16 Compair 8, 19 APR L L L L L L
15 32 Carstens V Cv L L L L L L
16 64 Spaldings Prolific Sp L L L L L L
17 128 Heines VII 2,25,HVII L L L L L L
PATHOTYPE α 68E0 68E0 67E0 67E0 66E0 66E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotype identified as 68E0 (from 1 isolate in each year), 67E0 (from 4 and 3) and 66E0 (from 3 and 2) isolates tested during 2005-06 and 2006-07.
209
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Peshawar
06-P-9,20,21,23
07-P-27,29,32
06-P-2,5,7,10
07-P-33,41,43,44,
45,49
06-P-12, 15,16,18
07-P-24,25,28
05-06 06-07 05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 H H L L H H
2 2 Lee 7 H H H H H H
3 4 Heines Kolben 2,6 L L H H H H
4 8 Vilmorin 3V L L L L L L
5 16 Moro 10 L L L L L L
6 32 Strubes Dickkopf Sd,25 L L L L L L
7 64 Suwon92 / Omar 4,Su H H H H H H
8 128 Clement 2,9,25,Cle L L L L L L
9 256 Triticum spelta album 5 L L L L L L
European set
10 1 Hybrid 46 4+ L L L L L L
11 2 Reichersberg 42 7,25 L L L L L L
12 4 Heines Peko 2,6,25 L L L L L L
13 8 Nord Desprez 3N L L L L L L
14 16 Compair 8, 19 APR L L L L L L
15 32 Carstens V Cv L L L L L L
16 64 Spaldings Prolific Sp L L L L L L
17 128 Heines VII 2,25,HVII L L L L L L
PATHOTYPE α 67E0 67E0 70E0 70E0 71E0 71E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α identified as 67E0 and 71E0 (from 4 and 3 isolates) and 70E0 (from 4 and 6 isolates) tested during 2005-06 and 2006-07, respectively.
210
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Appendix II Pathotype Determination of P. striiformis f. sp. tritici through Seedling Infection Types Produced on world and European Differentials during 05-06 & 06-07
Sr. No.
Race Nomenclature Differentials
Yr resistance
gene(s)
Accession Nos. of Pst Isolates collected from District Mardan
06-M-5,6
07-M-16
06-M-1,12,13
07-M-21,24, 28
06-M- 9
07-M-19
06-M- 2
07-M- 27
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07
World set
1 1 Chinese 166 1 L L H H H H L L
2 2 Lee 7 H H H H H H H H
3 4 Heines Kolben 2,6 L L L L H H H H
4 8 Vilmorin 3V L L L L L L L L
5 16 Moro 10 L L L L L L L L
6 32 Strubes Dickkopf Sd,25 L L L L L L L L
7 64 Suwon92 / Omar 4,Su H H H H H H H H
8 128 Clement 2,9,25,Cle L L L L L L L L
9 256 Triticum spelta album 5 L L L L L L L L
European set
10 1 Hybrid 46 4+ L L L L L L L L
11 2 Reichersberg 42 7,25 L L L L L L L L
12 4 Heines Peko 2,6,25 L L L L L L L L
13 8 Nord Desprez 3N L L L L L L L L
14 16 Compair 8, 19 APR L L L L L L L L
15 32 Carstens V Cv L L L L L L L L
16 64 Spaldings Prolific Sp L L L L L L L L
17 128 Heines VII 2,25,HVII L L L L L L L L
PATHOTYPE α 66E0 66E0 67E0 67E0 71E0 71E0 70E0 70E0
L refers to Infection Type 0-5 (Avirulent Reaction); and H refers to Infection Type 6-9 (Virulent Response). α Pathotype identified as 66E0 ( from 2 and 1), 67E0 (from 3 isolates in each year) while 71E0 and 70E0 ( from 1 isolate in each year) tested during 2005-06 and 2006-07, respectively;
211
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Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Sialkot Rawalpindi
06-S-1 06-S-4 06-S-5 06-S-9 06-S-13 06-S-17 06-S-18 06-S-20 06-R-2 06-R-3
1 2 3 4 5 6 7 8 9 10
1 Yr1/6* Avocet(s) 1 4 4 5 4 5 4 5 4 4 4 5 5 4 5 4 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 7 6 6 7 6 7 7 7 6 7 7 8 8
4 Yr7/6* Avocet(s) 7 7 6 7 6 7 6 6 7 6 7 6 7 6 6 7 6 7
5 Yr8/6* Avocet(s) 8 1 2 2 2 1 2 1 2 2 1 2 2 2 3 2 3
6 Yr9/6* Avocet(s) 9 6 7 6 7 6 6 7 6 7 7 7 6 7 6 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 6 6 7 6 7 6 7 6 7 7 6 7 6 7
12 Yr18/3* Avocet(s) 18 6 4 6 6 7 6 4 5 3 4 4 7 7
13 Yr24/3* Avocet(s) 24 6 7 4 5 6 7 6 6 3 4 4 5 4 5 6 6
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 6 7 6 6 7 6 7 6 7 6 6 6
17 Jupateco R (Yr18) 18 6 7 3 4 6 6 7 6 3 4 3 4 3 4 6 7 7
18 Jupateco S - 6 7 7 6 6 6 7 6 7 6 6 7 8 7 8
19 Avocet + YrA A 5 6 6 7 6 6 6 5 6 6 5 6 7 7
Infection Type 0 - 5 = Avirulent Reaction, Infection Type 6 - 9 = Virulent Reaction
211
213
Sheet 2 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Rawalpindi
06-R-6 06-R-7 06-R-9 06-R-12 06-R-14 06-R-18 06-R-19 06-R-21 06-R-22 06-R-25
11 12 13 14 15 16 17 18 19 20
1 Yr1/6* Avocet(s) 1 4 4 5 5 5 5 6 4 6 7 4 4 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 8 7 8 7 8 6 7 6 7 7 7 8 7 8 8 7 8
4 Yr7/6* Avocet(s) 7 6 7 6 7 8 7 8 8 7 7 6 7 6 7 7
5 Yr8/6* Avocet(s) 8 6 7 6 7 2 3 6 7 2 3 3 4 6 7 4 6 6 7
6 Yr9/6* Avocet(s) 9 6 6 6 7 7 6 6 7 6 7 6 6
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 7 6 7 7 7 6 7 7 6 7 7 7 6 7
12 Yr18/3* Avocet(s) 18 4 5 3 4 6 3 4 7 7 8 6 7 7 3 4 4
13 Yr24/3* Avocet(s) 24 4 4 5 6 5 6 6 4 6 7 5 4 5
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 7 6 6 6 7 7 6 7 6 6 7 6
17 Jupateco R (Yr18) 18 4 6 7 6 3 4 6 7 7 6 7 6 7 3 4 3
18 Jupateco S - 6 7 6 7 7 7 6 7 6 7 7 6 7 7 6 7
19 Avocet + YrA A 8 7 8 8 8 7 8 8 8 8 8 7 8
Infection Type 0 - 5 = Avirulent Reaction, Infection Type 6 - 9 = Virulent Reaction
212
214
Sheet 3 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Rawalpindi Chakwal
06-R-26 06-R-28 06-R-30 06-R-31 06-R-33 06-R-35 06-R-36 06-Ck-1 06-Ck-3 06-Ck-4
21 22 23 24 25 26 27 28 29 30
1 Yr1/6* Avocet(s) 1 3 4 6 7 4 5 4 5 6 6 7 4 5 4 5 5 5 6
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 8 7 8 8 8 7 8 7 8 7 8 6 7 6 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 7 7 6 7 7 6 7 6 7 6 7 6 7 6 7
5 Yr8/6* Avocet(s) 8 2 3 4 3 4 6 3 4 4 5 6 7 5 6 6 6
6 Yr9/6* Avocet(s) 9 6 6 6 7 6 7 6 7 6 6 6 6 6
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 7 6 7 7 7 6 7 7 7 6 7 7 6 7
12 Yr18/3* Avocet(s) 18 6 6 7 7 7 7 8 7 7 8 3 4 6 7 3 4
13 Yr24/3* Avocet(s) 24 6 7 6 6 4 5 6 7 7 4 5 5 6 5 6 6
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 7 6 6 7 6 7 7 7 6 7 6 6 6 7
17 Jupateco R (Yr18) 18 6 7 6 6 7 6 6 6 7 6 7 3 4 6 4
18 Jupateco S - 7 8 7 7 8 7 7 8 7 8 7 6 7 7 7
19 Avocet + YrA A 8 7 8 8 8 7 8 8 8 7 8 7 8
213
215
Sheet 4 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Chakwal Attock
06-Ck-6 06-Ck-7 06-Ck-9 06-Ck-
10 06-Ck-
11 06-A-3 06-A-7 06-A-9 06-A-13 06-A-14
31 32 33 34 35 36 37 38 39 40
1 Yr1/6* Avocet(s) 1 5 5 4 5 4 5 4 5 6 7 5 6 5 6 6 6 7
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 6 7 6 7 6 7 7 6 7 6 7 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 6 7 6 7 6 6 7 6 7 7 7 6 7 7
5 Yr8/6* Avocet(s) 8 5 6 5 6 2 3 5 6 2 3 4 3 4 3 4 4 6 7
6 Yr9/6* Avocet(s) 9 6 6 7 6 6 6 7 6 7 7 6 7 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 6 7 6 7 7 6 7 7 6 7 6 7 7 6 7
12 Yr18/3* Avocet(s) 18 3 4 3 4 4 2 3 3 4 6 6 6 7 6 7 6 7
13 Yr24/3* Avocet(s) 24 5 6 5 6 5 5 6 4 5 6 7 6 7 7 7 6 7
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 6 6 7 6 6 6 6 7 6 7 6
17 Jupateco R (Yr18) 18 5 6 3 4 4 5 6 3 4 6 7 7 6 7 6 7 7
18 Jupateco S - 7 6 7 6 7 7 6 7 8 7 8 8 7 8 8
19 Avocet + YrA A 7 8 7 7 8
7 8 8 7 8 7 7 8 7 8 7
214
216
Sheet 5 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Attock Nowshera
06-A-17 06-A-18 06-N-1 06-N-2 06-N-3 06-N-4 06-N-6 06-N-7 06-N-10 06-N-12
41 42 43 44 45 46 47 48 49 50
1 Yr1/6* Avocet(s) 1 5 6 5 6 5 7 6 7 4 5 4 5 7 6 7 4 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 7 7 8 7 6 7 7 6 7 6 7 7
4 Yr7/6* Avocet(s) 7 7 6 7 7 6 7 6 7 6 7 6 7 6
5 Yr8/6* Avocet(s) 8 3 4 4 6 7 6 7 7 6 7 3 4 6 6 6 7
6 Yr9/6* Avocet(s) 9 6 7 7 7 6 7 6 7 7 6 7 6 7 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 6 7 7 6 7 6 7 7 6 7 7 7
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 8 7 8 7 6 7 7 6 7 7 6 7 7 7
12 Yr18/3* Avocet(s) 18 6 6 6 7 6 7 6 4 5 4 5 6 7 6 7 7
13 Yr24/3* Avocet(s) 24 6 7 7 5 6 4 5 4 5 6 4 5 6 6 5 6
14 Yr26/3* Avocet(s) 26 0 0 5 6 4 5 4 5 4 5 4 5 6 7 7 5 6
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 6 7 6 6 7 6 6 6 7 6 6 7
17 Jupateco R (Yr18) 18 6 7 6 7 5 6 6 7 6 4 5 4 5 6 6 7 5 6
18 Jupateco S - 7 8 7 7 8 7 7 8 8 7 8 8 7 8 7 8
19 Avocet + YrA A 7 7 8 8
7 8 8 7 8 8 7 8 8 7 8
215
217
Sheet 6 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Nowshera Charsadda
06-N-13 06-N-15 06-N-18 06-N-21 06-N-24 06-N-27 06-N-30 06-Cd-1 06-Cd-4 06-Cd-6
51 52 53 54 55 56 57 58 59 60
1 Yr1/6* Avocet(s) 1 3 4 4 5 5 6 4 6 5 6 4 5 7 6 7 7
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 7 6 7 7 6 7 7 6 6 7 6 7 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 7 6 7 6 7 6 6 7 7 6 7 7
5 Yr8/6* Avocet(s) 8 2 2 3 6 7 2 3 2 2 2 3 5 4 5 5
6 Yr9/6* Avocet(s) 9 6 7 6 7 7 6 7 7 6 7 7 7 6 7 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 7 3 4 6 7 4 6 7 7 4 5 6 7 3 4 3 4
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 7 6 7 7 7 6 7 7 7 8 8 7 8
12 Yr18/3* Avocet(s) 18 3 4 6 5 6 6 7 3 4 3 4 6 7 6 6 6 7
13 Yr24/3* Avocet(s) 24 5 6 6 5 6 6 5 6 6 6 7 6 7 6 6 7
14 Yr26/3* Avocet(s) 26 5 6 4 5 4 5 4 5 7 6 7 5 4 5 4 5 4
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 6 7 6 7 5 6 5 6 6 5 6 6 6
17 Jupateco R (Yr18) 18 3 4 6 7 4 5 6 3 4 4 6 4 6 6 7
18 Jupateco S - 7 8 7 7 8 7 8 7 7 8 7 7 8 7 7 8
19 Avocet + YrA A 8 7 8 8
8 7 8 8 8 8 9 8 9 8
216
218
Sheet 7 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Charsadda Peshawar
06-Cd-9 06-Cd-
11 06-Cd-
12 06-Cd-
14 06-Cd-
15 06-Cd-
17 06-Cd-
18 06-P-2 06-P-3 06-P-5
61 62 63 64 65 66 67 68 69 70
1 Yr1/6* Avocet(s) 1 6 7 6 7 7 7 7 6 7 7 6 5 6 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 7 7 6 7 6 7 6 7 7 6 7 6 7 6 6 7
4 Yr7/6* Avocet(s) 7 7 6 7 7 7 7 6 7 7 7 6 7 7
5 Yr8/6* Avocet(s) 8 4 5 4 5 5 4 5 5 4 5 4 5 0 0 0
6 Yr9/6* Avocet(s) 9 6 7 6 7 7 7 6 7 6 7 7 6 7 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 3 4 4 4 5 4 5 4 4 4 5 2 2 3 2
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 6 7 6 6 7 6 6 7 6 6 6 6 7
12 Yr18/3* Avocet(s) 18 6 6 7 7 6 7 6 6 7 6 7 6 6 7 6 7
13 Yr24/3* Avocet(s) 24 6 6 6 7 7 6 7 6 6 7 6 7 6 7 7
14 Yr26/3* Avocet(s) 26 4 5 4 5 4 4 5 4 5 4 5 4 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 5 6 6 6 6 5 6 6 6 6 6
17 Jupateco R (Yr18) 18 6 6 7 6 6 7 6 7 6 6 7 6 7 6 7 6
18 Jupateco S - 7 7 7 8 7 7 8 7 7 8 8 7 8 8
19 Avocet + YrA A 7 8 7 8 8
7 8 7 8 7 8 8 8 8 7 8
217
219
Sheet 8 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Peshawar
06-P-7 06-P-8 06-P-9 06-P-10 06-P-12 06-P-15 06-P-16 06-P-18 06-P-20 06-P-21
71 72 73 74 75 76 77 78 79 80
1 Yr1/6* Avocet(s) 1 5 6 6 7 6 5 6 6 7 6 6 6 7 7 6 7
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 7 6 7 6 7 6 7 6 7 6 7 6 7 6 7 6 7 7
4 Yr7/6* Avocet(s) 7 6 7 7 6 7 6 7 7 7 6 7 7 7 7
5 Yr8/6* Avocet(s) 8 0 0 0 0 0 0 0 0 0 0
6 Yr9/6* Avocet(s) 9 7 6 7 6 7 7 7 7 6 7 7 6 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 2 0 2 2 3 1 1 2 2 2 3 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 7 6 7 7 7 6 7 6 7 7 6 7 7
12 Yr18/3* Avocet(s) 18 6 6 7 6 6 7 6 6 7 6 6 6 7 6
13 Yr24/3* Avocet(s) 24 7 6 7 7 6 7 6 7 7 6 7 6 7 7 6 7
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 7 6 6 7 6 7 6 6 6 6 7 6
17 Jupateco R (Yr18) 18 6 6 6 7 6 7 6 6 7 6 6 7 6 6 7
18 Jupateco S - 7 8 8 8 7 8 8 8 7 8 8 7 8 8
19 Avocet + YrA A 8 7 8 8
7 8 8 7 8 8 8 7 8 7 8
218
220
Sheet 9 of 9
Appendix III Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2005-06
Sr. No.
Near Isogenic Lines with Avocet background
Yr Resistance Gene
Peshawar Mardan
06-P-23 06-M-1 06-M-2 06-M-5 06-M-6 06-M-8 06-M-9 06-M-11 06-M-12 06-M-13
81 82 83 84 85 86 87 88 89 90
1 Yr1/6* Avocet(s) 1 6 6 7 4 5 7 6 7 4 5 7 6 7 6 7 6 7
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 7 7 8 7 7 8 7 7 8 7 8 7 8 7
4 Yr7/6* Avocet(s) 7 7 6 7 6 7 6 7 6 7 7 7 6 7 7 7
5 Yr8/6* Avocet(s) 8 0 4 5 3 4 4 5 5 3 4 4 4 5 4 5 4
6 Yr9/6* Avocet(s) 9 6 7 6 7 7 6 7 7 7 7 6 7 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 2 4 4 4 5 5 4 5 4 5 4 4 5 4
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 7 6 7 6 7 7 7 6 7 7 7 6 7
12 Yr18/3* Avocet(s) 18 6 6 7 6 6 7 6 6 6 6 6 6 7
13 Yr24/3* Avocet(s) 24 6 6 7 6 7 7 6 7 6 6 7 6 7 6 6
14 Yr26/3* Avocet(s) 26 0 4 5 5 5 4 5 4 5 5 4 5 4 5 5
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 6 6 6 7 6 7 6 6 6 7 6 7
17 Jupateco R (Yr18) 18 6 6 7 6 7 6 7 6 6 6 7 6 6 7
18 Jupateco S - 8 7 7 8 7 8 8 7 7 8 7 8 7 7 8
19 Avocet + YrA A 8 7 8 8
8 7 8 8 7 8 7 8 8 7 8
219
221
Sheet 1 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Sialkot
07-S-22 07-S-25 07-S-26 07-S-28 07-S-30 07-S-32 07-S-34 07-S-36 07-S-37 07-S-38
1 2 3 4 5 6 7 8 9 10
1 Yr1/6* Avocet(s) 1 4 5 4 4 5 4 5 4 5 4 5 5 4 5 4 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 6 7 6 6 7 6 7 7 7 6 7 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 6 6 7 6 7 6 7 6 6 7 6 6 7 6 7
5 Yr8/6* Avocet(s) 8 0 0 0 0 0 0 0 0 0 0
6 Yr9/6* Avocet(s) 9 6 6 6 6 7 6 6 7 6 7 6 6 6
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 6 6 7 6 7 7 7 6 7 7 6 7 6 7
12 Yr18/3* Avocet(s) 18 6 7 3 4 3 4 6 7 7 3 4 6 6 7 6 4
13 Yr24/3* Avocet(s) 24 6 7 2 2 3 6 6 7 2 3 6 6 7 6 2
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 7 6 6 6 6 7 6 6 7 6 7 6 7
17 Jupateco R (Yr18) 18 6 4 3 4 6 6 3 4 6 6 6 7 3 4
18 Jupateco S - 6 6 6 7 6 6 7 6 6 6 6 6
19 Avocet + YrA A 6 7 6 7 6
6 7 6 6 6 6 7 6 6
220
222
Sheet 2 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Sialkot Rawalpindi
07-S-42 07-S-44 07-R-38 07-R-39 07-R-40 07-R-42 07-R-43 07-R-46 07-R-47 07-R-48
11 12 13 14 15 16 17 18 19 20
1 Yr1/6* Avocet(s) 1 4 5 4 5 5 4 5 4 5 5 5 4 5 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 7 6 7 6 6 6 7 6 7 6 6 6 7
4 Yr7/6* Avocet(s) 7 6 7 6 7 6 7 6 7 7 6 7 7 7 6 7
5 Yr8/6* Avocet(s) 8 0 0 5 6 5 5 6 4 5 4 5 4 4 5 5
6 Yr9/6* Avocet(s) 9 6 6 6 7 6 6 7 6 6 6 7 6 7 6
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 7 7 6 7 7 6 7 7 6 7 6 7 7 6 7
12 Yr18/3* Avocet(s) 18 3 4 4 6 7 6 6 7 6 7 5 4 5 4 5 6 7
13 Yr24/3* Avocet(s) 24 2 2 3 6 7 6 6 6 7 4 5 4 5 5 6
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 7 6 7 6 6 6 6 7 6 7 6 7 6 7
17 Jupateco R (Yr18) 18 3 4 3 4 7 7 6 7 6 7 4 4 5 4 7
18 Jupateco S - 6 7 6 6 7 7 6 7 7 7 6 7 6 7 7
19 Avocet + YrA A 6 5 6 7 8
8 8 7 8 8 8 7 8 8
221
223
Sheet 3 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Rawalpindi Chakwal
07-R-49 07-R-51 07-R-54 07-R-57 07-R-59 07-R-63 07-R-64 07-R-67 07-Ck-13 07-Ck-14
21 22 23 24 25 26 27 28 29 30
1 Yr1/6* Avocet(s) 1 5 4 5 5 4 5 4 5 5 4 5 4 5 5 4 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 6 7 6 7 6 7 6 7 6 7 6
4 Yr7/6* Avocet(s) 7 7 6 7 7 6 7 6 7 6 6 7 6 7 6 7 7
5 Yr8/6* Avocet(s) 8 5 4 5 4 5 4 4 5 4 3 4 5 4 5 4 5
6 Yr9/6* Avocet(s) 9 6 6 6 7 6 6 6 6 7 6 6 6
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 7 6 7 7 6 7 7 6 7 6 6 7 7 7
12 Yr18/3* Avocet(s) 18 6 7 6 6 6 7 6 6 7 6 7 6 7 2 3 2
13 Yr24/3* Avocet(s) 24 6 6 7 6 7 6 6 6 7 6 6 7 4 5 4
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 0 0 0
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 6 7 6 7 6 7 6 7 6 6 6 7 6 7
17 Jupateco R (Yr18) 18 7 8 8 7 8 8 7 8 8 7 8 7 8 3 2 3
18 Jupateco S - 6 7 6 6 7 6 7 7 8 7 6 7 6 7
19 Avocet + YrA A 8 7 8 8
7 8 8 8 7 8 7 8 7 7 8
222
224
Sheet 4 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Chakwal Attock 07-Ck-
16 07-Ck-
17 07-Ck-20 07-Ck-23 07-Ck-24 07-Ck-27 07-Ck-28 07-A-20 07-A-22 07-A-24
31 32 33 34 35 36 37 38 39 40
1 Yr1/6* Avocet(s) 1 4 5 4 5 5 6 5 4 5 5 5 6 5 6 7 5 6
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 6 6 7 6 7 6 7 6 6 7 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 6 7 7 6 6 7 6 7 7 6 7 7 6 7
5 Yr8/6* Avocet(s) 8 5 6 5 6 4 5 5 6 5 6 4 5 6 5 6 5 6 5
6 Yr9/6* Avocet(s) 9 6 6 7 6 7 6 6 6 6 7 6 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 0 0 0 0 0 0 0 0 0 0
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 7 6 7 7 6 7 7 7 6 7 6 7 7
12 Yr18/3* Avocet(s) 18 2 3 3 2 3 2 3 2 3 2 2 3 6 7 6 6 7
13 Yr24/3* Avocet(s) 24 4 4 5 4 5 4 5 4 5 5 6 7 6 7 6 7
14 Yr26/3* Avocet(s) 26 0 0 0 0 0 0 0 3 4 4 3
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 7 6 6 7 6 6 6 6 6 7 0
17 Jupateco R (Yr18) 18 3 4 4 3 3 4 2 3 2 3 3 6 7 6 6 7
18 Jupateco S - 7 7 6 7 7 6 7 7 6 7 8 7 8 7 8
19 Avocet + YrA A 7 8 7 8
7 8 8 7 7 8 7 8 7 7
223
225
Sheet 5 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Attock
07-A-25 07-A-26 07-A-27 07-A-30 07-A-31 07-A-32 07-A-34 07-A-37 07-A-38 07-A-41
41 42 43 44 45 46 47 48 49 50
1 Yr1/6* Avocet(s) 1 5 6 6 7 7 7 6 7 6 7 7 7 5 6 7
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 7 7 6 7 6 7 7 7 7 6 7
4 Yr7/6* Avocet(s) 7 7 7 6 7 7 6 7 7 6 7 7 7 6 7
5 Yr8/6* Avocet(s) 8 5 6 6 7 5 6 7 6 7 6 7 5 6 6 7 5 6 7
6 Yr9/6* Avocet(s) 9 7 7 7 7 7 7 7 7 6 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 3 4 6 3 4 4 6 7 3 3 4 6 7 4 6 7
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 7 8 8 7 8 7 8 7 8 7 8 7 8 7 8 8 7 8
12 Yr18/3* Avocet(s) 18 8 8 7 8 7 8 7 8 7 8 8 8 7 8
13 Yr24/3* Avocet(s) 24 6 7 6 7 7 6 7 6 7 6 7 7 6 7 6 7 6 7
14 Yr26/3* Avocet(s) 26 3 4 6 4 3 4 6 7 3 3 4 6 3 4 6 7
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 7 6 6 6 7 7 6 7 6 7 7
17 Jupateco R (Yr18) 18 6 7 6 7 7 6 7 6 7 6 7 7 6 7 6 6 7
18 Jupateco S - 7 8 7 7 8 8 8 7 8 7 8 8 7 8 7 8
19 Avocet + YrA A 7 8 7 7 8
7 7 8 7 8 7 8 7 7 7
224
226
Sheet 6 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Nowshera
07-N-32 07-N-35 07-N-36 07-N-38 07-N-39 07-N-44 07-N-45 07-N-47 07-N-48 07-N-49
51 52 53 54 55 56 57 58 59 60
1 Yr1/6* Avocet(s) 1 6 7 4 5 6 7 6 6 7 4 5 6 7 5 6 5 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 7 8 7 7 8 7 7 6 7 7 7 7 8 6 7
4 Yr7/6* Avocet(s) 7 6 7 7 6 7 6 7 7 6 6 6 6
5 Yr8/6* Avocet(s) 8 0 0 0 0 0 0 0 0 0 0
6 Yr9/6* Avocet(s) 9 7 6 7 6 7 7 6 7 7 6 7 7 6 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 6 7 6 7 6 7 7 6 7 6 7 3 4 6 7 4
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 6 7 7 6 7 6 7 6 7 7 6 7 7 6 7
12 Yr18/3* Avocet(s) 18 6 7 5 6 6 7 6 7 5 6 5 6 6 7 6 7 5 6 6 7
13 Yr24/3* Avocet(s) 24 5 6 5 6 6 5 6 5 6 5 6 6 6 5 6 6
14 Yr26/3* Avocet(s) 26 3 4 5 6 7 4 5 6 4 5 7 5 5 5 6
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 7 7 7 7 6 7 6 7 6 7 6 6
17 Jupateco R (Yr18) 18 6 7 5 6 6 6 7 5 6 5 6 6 7 6 5 6
18 Jupateco S - 8 7 7 8 8 8 7 8 8 8 7 8 8
19 Avocet + YrA A 6 7 7 8 8
7 8 8 7 8 8 7 8 8 7 8
225
227
Sheet 7 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Nowshera Charsadda
07-N-51 07-N-52 07-N-55 07-N-56 07-N-59 07-N-60 07-Cd-22 07-Cd-23 07-Cd-24 07-Cd-25
61 62 63 64 65 66 67 68 69 70
1 Yr1/6* Avocet(s) 1 4 5 5 5 6 5 6 5 5 6 6 7 7 6 7 6 7
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 6 6 7 7 6 6 6 6 7 6 7
4 Yr7/6* Avocet(s) 7 7 7 6 7 6 7 6 7 6 6 7 7 7 7
5 Yr8/6* Avocet(s) 8 6 6 7 0 0 6 7 0 4 5 4 5 5 4 5
6 Yr9/6* Avocet(s) 9 7 6 7 7 6 7 7 6 7 6 7 6 7 6 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 6 7 7 4 5 4 5 6 7 4 5 4 4 5 5 4 5
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 6 6 6 7 6 7 6 7 7 8 8 7 8 7 8
12 Yr18/3* Avocet(s) 18 6 5 6 7 6 6 7 6 6 6 7 6 6 7
13 Yr24/3* Avocet(s) 24 5 6 5 6 6 6 5 6 6 6 6 6 7 6 7
14 Yr26/3* Avocet(s) 26 5 6 4 5 5 6 5 6 5 5 4 5 4 5 4 4 5
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 6 7 6 6 7 6 7 6 7 6 6 7 6 7
17 Jupateco R (Yr18) 18 5 6 5 6 6 7 6 5 6 6 6 7 6 6 7 6 7
18 Jupateco S - 8 7 8 8 8 7 8 7 8 7 7 8 7 8 7 8
19 Avocet + YrA A 7 8 7 8 8
7 8 7 8 8 7 8 8 8 7 8
226
228
Sheet 8 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Charsadda Peshawar 07-Cd-
28 07-Cd-
30 07-Cd-33 07-Cd-34 07-P-24 07-P-24 07-P-25 07-P-26 07-P-27 07-P-29
71 72 73 74 75 76 77 78 79 80
1 Yr1/6* Avocet(s) 1 7 6 7 6 7 6 7 6 6 7 4 5 6 7 7 7 8
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 7 6 7 7 6 7 6 7 7 8 7 6 7 6 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 6 7 6 7 7 7 8 7 8 6 7 6 6 7
5 Yr8/6* Avocet(s) 8 4 5 5 5 4 5 4 5 5 6 5 6 4 5 5 6 5 6
6 Yr9/6* Avocet(s) 9 7 6 7 7 6 7 7 8 6 7 7 8 7 8 6 7 7 8
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 4 5 5 4 5 5 6 4 5 4 4 5 5 4 5 4 5
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 7 7 7 6 7 7 6 7 6 7 7 7 6 7
12 Yr18/3* Avocet(s) 18 6 7 6 6 7 6 6 7 6 7 6 7 6 7 6 7 6
13 Yr24/3* Avocet(s) 24 6 7 6 6 7 7 6 7 6 6 7 6 6 7 6 7
14 Yr26/3* Avocet(s) 26 4 4 5 4 4 5 4 3 4 4 5 4 4 3 4
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 6 6 6 7 6 7 6 6 7 6 6
17 Jupateco R (Yr18) 18 6 7 6 7 6 6 6 7 6 7 6 7 6 7 7 6 7
18 Jupateco S - 7 8 7 8 7 8 7 8 7 7 8 7 8 7 8 8
19 Avocet + YrA A 8 7 8 8
7 8 8 8 7 8 8 7 8 7 8
227
229
Sheet 9 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Peshawar
07-P-32 07-P-33 07-P-35 07-P-37 07-P-38 07-P-39 07-P-41 07-P-43 07-P-44 07-P-45
81 82 83 84 85 86 87 88 89 90
1 Yr1/6* Avocet(s) 1 6 5 6 6 7 6 4 4 5 4 5 5 4 5
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 6 6 7 6 7 6 7 6 7 7 6 7 6 7 6 7
4 Yr7/6* Avocet(s) 7 6 7 6 7 8 7 7 8 6 7 6 7 7 6 7 7
5 Yr8/6* Avocet(s) 8 5 4 5 4 5 6 5 5 4 4 5 4 5 5
6 Yr9/6* Avocet(s) 9 7 8 7 8 7 8 7 6 7 6 7 7 7 6 7 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 4 5 4 4 5 5 4 4 4 5 4 4 4 5
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 6 7 6 7 7 6 7 6 7 6 7 6 7 6 7 7
12 Yr18/3* Avocet(s) 18 6 7 6 6 7 6 6 6 7 6 7 6 7 6 7
13 Yr24/3* Avocet(s) 24 6 7 7 6 7 6 6 6 7 6 6 7 6 6 7
14 Yr26/3* Avocet(s) 26 3 4 4 3 3 4 3 3 3 4 4 4 3 4
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 6 7 6 7 6 7 6 7 6 6 6 6 7 6
17 Jupateco R (Yr18) 18 6 7 6 6 6 7 6 7 6 6 7 6 6 7 6 7
18 Jupateco S - 7 8 7 7 8 8 7 8 7 7 8 6 7 6 7 7
19 Avocet + YrA A 7 7 8 7
6 7 8 7 8 7 8 7 8 8 8
228
230
Sheet 10 of 10
Appendix IV Infection Types of Single Pustule Isolates of Pst on Stripe Rust Differentials (NILs) Based on Seedling Reactions during 2006-07
Sr. No.
Near Isogenic Lines with Avocet
background
Yr Resistance
Gene
Peshawar Mardan
07-P-48 07-P-49 07-M-16 07-M-17 07-M-19 07-M-21 07-M-24 07-M-27 07-M-28
91 92 93 94 95 96 97 98 99
1 Yr1/6* Avocet(s) 1 4 5 4 6 6 7 6 7 7 8 6 4 5 7 8
2 Yr5/6* Avocet(s) 5 0 0 0 0 0 0 0 0 0
3 Yr6/6* Avocet(s) 6 6 6 7 7 8 7 8 8 7 8 7 8 7 7 8
4 Yr7/6* Avocet(s) 7 6 7 7 6 7 6 7 7 8 6 7 6 7 7 8 6 7
5 Yr8/6* Avocet(s) 8 5 6 5 6 4 5 4 4 5 4 5 5 4 5 5
6 Yr9/6* Avocet(s) 9 7 6 7 6 7 7 7 6 7 6 7 7 6 7
7 Yr10/6* Avocet(s) 10 0 0 0 0 0 0 0 0 0
8 Yr11/3* Avocet(s) 11 0 0 0 0 0 0 0 0 0
9 Yr12/3* Avocet(s) 12 3 4 4 4 5 5 5 4 5 4 5 5 4
10 Yr15/6* Avocet(s) 15 0 0 0 0 0 0 0 0 0
11 Yr17/6* Avocet(s) 17 6 7 7 6 7 6 6 7 6 7 6 7 6 7
12 Yr18/3* Avocet(s) 18 6 7 7 8 6 7 6 6 6 6 7 6 6
13 Yr24/3* Avocet(s) 24 6 7 6 7 6 6 7 6 6 6 6 7 6 7
14 Yr26/3* Avocet(s) 26 0 0 4 5 4 5 4 5 5 4 5 5 4 5
15 YrSP/3* Avocet(s) Sp 0 0 0 0 0 0 0 0 0
16 Yr27/3*Avocet(CX 94.19.1.1) Sk 6 7 6 6 7 6 6 6 7 6 7 6 6
17 Jupateco R (Yr18) 18 6 7 7 6 6 7 6 7 6 6 7 6 7 6
18 Jupateco S - 7 8 8 7 7 8 6 7 6 7 6 7 6 7
19 Avocet + YrA A 7 8 7 8 7 7 8 7 7 8 7 8 7 7 8
229
230
Appendix V Stripe rust reactions of commercial varieties to Puccinia striiformis f. sp. tritici at different locations of Northern Punjab & NWFP during 2005-06 & 2006-07
Sr. No
Differential Sets
Yr Genes
Disease Reaction to Yr at Trap nurseries established at
PRS (Sialkot) PMAS-AAU (Rwp) NARC (Islamabad) CCRI (Pirsabak) AUP (Peshawar) NIFA (Peshawar)
05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 05-06 06-07 1 Morocco 80S 90S 80S 100S 100S 100S 90S 90S 90S 100S 100S 100S 2 Pavon 76 Yr29 APR 0 0 5MS 5MS 10MS 10MS 0 0 0 0 0 0 3 MH 97 0 0 60S 70S 70S 80S 30S 60S 80S 40S 60S 50S 4 Inquilab-91 Yr27 0 0 30S 40S 40S 50S 50S 50S 30S 40S 30S 30S 5 Kohistan 97 10R 5R 80S 80S 30S 40S 60S 30S 0 0 40S 50S 6 Punjab 85 0 0 5MRMS 10MS 10MS 20S 50S 40S 10S 40S 80S 70S 7 Bakhtawar 93 Yr9+ 0 0 20MS 30MS 30MRMS 40S 20S 20S 0 0 50MSS 50S 8 Blue Silver Yr6+A 0 0 50S 80S 60S Missed 30S 20S 0 10MRMS 50S 40S 9 Chakwal 86 0 0 30MRMS 40MRMS 20MRMS 20MRMS 30MRMS 40MRMS 0 5MRMS 40MRMS 20MRMS 10 Sindh-81 0 0 50S 70S 40S 50S 40MRMS 30S 20MSS 30MSS 60S 50S 11 Zarghoon Yr6+ 0 0 10MSS 10S 60S 40S 70S 40S 10S 20S 50S 40S 12 Faisalabad 83 Yr7+2 APR 0 0 50S 80S 60S 80S 30S 70S 10MSS 20MSS 60S 70S 13 Faisalabad 85 Yr9+Yr4 0 0 20S 30S 50S 50S 60S 60S 10MSS 10MSS 40MRMS 40MS 14 Kaghan 93 0 0 90S 60S 70S 40S 60S 30S 10MSS 10MSS 60S 50S 15 Kirin 95 0 0 90S 80S 80S 60S 70S 40S 20MS 30MS 80S 50S 16 Kohinoor 83 0 0 80S 60S 60S 50S 90S 40S 20MS 30MS 70S 60S 17 LU-26 Yr6 0 0 70S 60S 80S 40S 50S 40S 20MSS 40MSS 60S 70S 18 Nowshera 96 0 0 50MSS 30S 50S 40S 50S 20S 10MSS 20MSS 60S 60S 19 Parwaz 94 Yr6+Yr7 0 0 20MSS 20MSS 40MSS 10S 40MSS 20MS 0 TRMR 10MSS 20MSS 20 Pasban 90 0 0 50S 50S 30MSS 20S 70S 20S 0 TR 60S 60S 21 Pirsabak 85 0 0 80S 60S 80S 50S 60S 40S 0 TR 60S 40S 22 Punjab 96 0 0 80S 50S 60S 60S 80S 50S 10MSS 10MSS 50S 60S 23 Sariab-92 Yr6+ 0 0 80S 80S 80S 70S 50S 60S 20S 40S 60S 70S 24 Sarsabz Yr7 0 0 80S 70S 60S 60S 50S 70S 10S 20S 60S 50S 25 Shaheen 94 0 0 80S 60S 30MSS 30S 40S 40S 0 10MSS 50S 40S 26 Shahkar 95 Yr6+ 0 0 10MS 20MS 30MRMS 10S 40MRMS 30S 0 10MRMS 50MRMS 40MRMS 27 Soughat 90 Yr6+Yr7 0 0 10S 10S 10S 10S 10S 10S 10MS 10MS 40S 40S 28 Tandojam 83 Yr6+ 0 0 100S 60S 80S 70S 60S 70S 40S 50S 60S 50S 29 SH-2002 0 0 90S 60S 70S 60S 40MSS 40S 60S 50S 60MSS 50S 30 Pak 81 Yr9+Yr7 0 0 70S 50S 60S 50S 60S 40S 40S 60S 70S 50S 31 Bahawalpur-97 0 0 40S 30S 40S 30S 60S 20S 40S 40S 60MSS 40MSS 32 Kohsar 93 0 0 0 0 10MRMS 0 10MS 5MS 0 TR 10MS 5MSS 33 Rohtas 90 0 0 10MSS 30MSS 40S 50S 20MSS 40S 10S 20S 10MSS 10MSS 34 Suleman 96 0 0 50S 50S 70S 30S 60S 30S 10MSS 20MSS 70S 60S 35 WL 711 Yr2 0 0 80S 70S 60S 70S 80S 70S 10S 40S 70S 50S 36 Zardana Yr7 0 0 80S 80S 80S 40S 50S 30S 10S 40S 40S 50S 37 Abadgar 93 0 0 40S 50S 60S 30S 40S 30S 10MSS 30MSS 50S 60S 38 Anmol-91 Yr9 0 0 50S 50MS 60MSS 40S 30MSS 40S 0 20MSS 5MRMS 5MRMS 39 Bahawalpur-2000 0 0 20MSS 20MSS 30S 10S 50S 20S 5MSS 5MSS 50S 60S 40 Bahkhar-2002 0 0 70S 40MRMS 80S 30MRMS 0 5MRMS 0 20MSS 0 0 41 Fakhr-e-Sarhad 0 0 0 0 0 5MS 10MS 10MS 10MS 5MS 10MS 10MS 42 Marvi-2000 0 0 0 TR 0 10MS 20S 10S 20S 10S 30S 40S 43 Mehran-89 Yr9 0 0 70S 50S 40S 50S 0 30S 0 5MS 0 0 44 Soorab-96
(Barley) 0 0 TR TR 0 0 0 0 0 0 0 0
45 Tatara Yr3 0 0 0 0 TR 0 0 TMR 0 TMS 0 5RMR 46 Takbeer 0 0 70S 50MS 0 10S 0 10MS 0 40MS 0 40MSS 47 AS-2002 0 0 20MSS 20MS 0 10S 0 30MSS 0 5S 0 TR 48 Iqbal 2000 Yr9 0 0 0 0 0 0 10MS 10MS 0 TR 0 TR 49 Auqab-2000 Yr9 0 0 10MRMS 10MRMS 20MR 20MR 10MR 0 10MS 30MS 20MSS 30MSS 50 Chakwal-97 0 0 70S 50S 60MSS 50S 50MSS 30MSS 0 10MRMS 10MRMS 10MRMS 51 Durum-97 0 0 10MR 20MR 20MR TR 0 TR 0 5R 0 5R 52 Watan 94 0 0 20MSS 20MS 50MSS 20S 20MSS 30MSS 10MSS 20MSS 50S 50S 53 Moomal 2002 0 0 80S 80S 70S 60S 60S 20S 20S 40S 30MSS 40MSS 54 Zarlashta Yr9 0 0 10S 30S 40MSS 50S 30S 20MS 0 10MSS 0 10MR 55 GA-2002 0 0 TR TR 0 0 TR 0 0 0 TR TR 56 Wafaq-01 Yr9 0 0 20S 20S 20S 20S 40MS 40MS 0 10S 30S 30S 57 Margalla-99 0 0 60S 60S 70S 60S 70S 80S 30S 60S 30S 50S 58 Manthar-3 Yr9 0 0 20S 30S 30MSS 20S 10MSS 10S 0 5MRMS 20MRMS 20MSS
a Field response reactions (0 to 100) represent the highest percentage infection recorded on the flag leaves for each entry during the growing season. + = Resistant, - = Susceptible, ** = Partial Resistant
230
231
Appendix VI Screening results of wheat germplasm based on Coefficient of Infection, Relative Resistance Index recorded during the year 2005-06 and 2006-07
Line / Variety
Disease Reaction Coefficient of
Infection CI
Total
ACI
RRI 2005-06 2006-07 2005-06 2006-07
RAWAL 87, IQBAL 2000, V 02172, 99FJ03, 1C020, 3C060, 3C062, 3C063, Asfoor – 3, Dajaj – 5, Husi – 2, Atris – 1, Girwill – 1, NR – 189, DHARWAR DRY, KLEIN CHAMACO, PBW343, KLEIN CACIQUE, KAUZ / PASTOR, WEAVER/4/NAC/TH.AC/3*PVN/3/MIRLO/BUC, CROC_1/AE.SQUARROSA(205)//KAUZ/3/SASIA, DHARWAR DRY / NESSER, SUJATA / SERI, PASTOR//SITE/MO/3/CHEN/AEGILOPS SQUARROSA (T…, FILIN/IRENA/5/CNDO/R143//ENTE/MEXI_2/3/…, CHIL/2*STAR/5/CANDO/R143//ENTE/MEXI_2/3/…, MUNIA//CHEN/ALTAR, 84/3/CHEN/,VIV90/91.28/5/CNDO/R143/ENTE/MEXI_2/3/…, PJN/BOW//OPATA/3/CROC_1/AE.SQUARROSA(224)//…, VEE/MJI//2*TUI/3/PFAU/BOW//VEE#9, CHEN/AEGILOPS SQUARROSA (TAUS)//BCN/3/BAV92, CNDO/R143/ENTE/MEXI_2/3/…, RABE/6/WRM/4/FN/3*TH//K58/2*N/3/AUS-6869/5/…, VEE/MJI//2*TUI/3/PASTOR, PASTOR/BAV92, IRENA/BABAX//PASTOR, ND/VG9144//KAL/BB/3/YACO/4/CHIL/5/PASTOR, CNO79//PF70354/MUS/3/URES/JUN/KAUZ, URES//BUC/PVN/3/KAUZ/4/FILIN, ND/VG9144//KAL/BB/3/YACO/4/CHIL/5/PASTOR, CROC_1/AE.SQUARROSA(224)//OPATA/3/PASTOR, BJY/COC//PRL/BOW/3/ATTILA, PARUS/PASTOR, FILIN/3/CROC_1/AE.SQUARROSA(205)//KAUZ/4/…, GEN*2//BUC/FLK/3/2*PASTOR, VIV90/91.28//2*PASTOR, FILIN/2*PASTOR, VEE/MJI//2*TUI/3/2*PASTOR, MILAN/KAUZ//BABAX/3/BABAX, PASTOR/3/ALTAR 84/AEGILOPS SQUARROSA (TAUS…, TUI/CLMS, NAI60/HEINE VII//BUC/3/PSN/BOW//TUI, ATTILA*2/HUITES, SNI/2*PASTOR, TRAP#1.R1/2*PASTOR, SERI.1B*2/4/COOK/VEE//DOVE/SERI/3/GEN, SERI.1B*2//TAM200/TUI, BAVIACORA M 92, ATTILA, ATTILA*2//CHIL/BUC, WEAVER/9/KT/BAGE//FN/U/3/BZA/4/TRM/5/…, SKAUZ/KS94U276//SKAUZ, BABAX/KS91WGRC11//BABAX, BABAX/KS94U276//BABAX
0
0
0
0
0
0
9
WEEBILL1 5 R 5 R 1 1 2 1 8.91
UQAB 2000 (AARI, FAISALABAD), SALEEM 2000 (CCRI, PIRSABAQ), KHYBER 87 (CCRI, PIRSABAQ), NR – 199, PJN/BOW//OPATA/3/CROC_1/AE.SQUARROSA(224)//…, TOB/ERA//TOB/CNO67/3/PLO/4/VEE#5/5/KUAZ/6/…, SERI M 82, SERI.1B*2/3/KAUZ*2/BOW//KAUZ, WEAVER/STAR//BORL95, SKAUZ/KS93U76//SKAUZ, SKAUZ/KS94U215//SKAUZ
TR
TR
0
0
0
0
9
SKAUZ/KS91WGRC11//SKAUZ 0 TR 0 0 0 0 9
SIREN/CLMS//SIREN TR R TR R 0 0 0 0 9
3C065 (B-LINE, BARI, CHAKWAL) 1 MR 5 MR 0.4 2 2.4 1.2 8.892
3C061 (B-LINE, BARI, CHAKWAL) 1 MR MS 1 MR MS 0.6 0.6 1.2 0.6 8.946
GA 2002 (BARI, CHAKWAL) 1 MR MS 5 MR MS 0.6 3 3.6 1.8 8.838
CHAKWAL 86 (BARI, CHAKWAL) 40 MR MS 50 MR MS 24 30 54 27 6.57
Chakwal 97 10 MR MS 10 MR MS 6 6 12 6 8.46
PASTOR/3/MUNIA//CHEN/ALTAR 84/5/CNDO/R143//… 20 MR MS 30 MR MS 12 18 30 15 7.65
CHAKWAL 86 5 MR MS 10 MR MS 3 6 9 4.5 8.595
CNDO/R143/ENTE/MEXI_2/3/… 10 MR MS 15 MR MS 6 9 15 7.5 8.325
CHEN/AEGILOPS SQUARROSA (TAUS)//BCN/3/BAV92 10 MR MS 15 MR MS 6 9 15 7.5 8.325
232
KAUZ//PRL/VEE#6/3/BAV92 5 MR MS 10 MR MS 3 6 9 4.5 8.595
URES/PRL//BAV92 10 MR MS 20 MR MS 6 12 18 9 8.19
ARIV92/BABAX//PASTOR 10 MR MS 10 MR MS 6 6 12 6 8.46
PASTOR//HXL7573/2*BAU 10 MR MS 20 MR MS 6 12 18 9 8.19
PASTOR/3/ALTAR 84/AEGILOPS SQUARROSA (TAUS… 10 MR MS 10 MR MS 6 6 12 6 8.46
DHARWAR DRY//HXL7573/2*BAU 20 MR MS 20 MR MS 12 12 24 12 7.92
ATTILA*2//CHIL/BUC 10 MR MS 10 MR MS 6 6 12 6 8.46
WEEBILL4 10 MR MS 10 MR MS 6 6 12 6 8.46
KAMBARA2 5 MR MS 10 MR MS 3 6 9 4.5 8.595
PBW65/2*SERI.1B 1 MR MS 5 MR MS 0.6 3 3.6 1.8 8.838
PFAU/SERI.1B//AMAD 1 MR MS 5 MR MS 0.6 3 3.6 1.8 8.838
SERI.1B//KAUZ/HEVO/3/AMAD 1 MR MS 5 MR MS 0.6 3 3.6 1.8 8.838
PASTOR 1 MR MS 10 MR MS 0.6 6 6.6 3.3 8.703
PBW65/2*PASTOR 5 MR MS 5 MR MS 3 3 6 3 8.73
HUW234+LR34/PRINIA 10 MR MS 10 MR MS 6 6 12 6 8.46
HUW234+LR34/3/KAUZ*2/TRAP//KAUZ 5 MR MS 10 MR MS 3 6 9 4.5 8.595
FRET2 20 MR MS 20 MR MS 12 12 24 12 7.92
ATTILA*2/PVN 10 MR MS 10 MR MS 6 6 12 6 8.46
ATTILA*2/STAR 5 MR MS 10 MR MS 3 6 9 4.5 8.595
BABAX/LR42/BABAX 5 MR MS 10 MR MS 3 6 9 4.5 8.595
BABAX/LR43/BABAX 20 MR MS 40 MR MS 12 24 36 18 7.38
3C069 (B-LINE, BARI, CHAKWAL) 10 MS S 10 MS S 9 9 18 9 8.19
SULEMAN 96 (CCRI, PIRSABAQ) 5 MS S 10 MS S 4.5 9 13.5 6.75 8.3925
00FJ03 (MICROLINE, FATEHJANG) 10 MS S 10 MS S 9 9 18 9 8.19
2495 (MICROLINE BAHAWALPUR) 20 MS S 30 MS S 18 27 45 22.5 6.975
116/10 5 MS S 10 MS S 4.5 9 13.5 6.75 8.3925
106/20 10 MS S 10 MS S 9 9 18 9 8.19
124/2 30 MS S 40 MS S 27 36 63 31.5 6.165
125/1 40 MS S 50 MS S 36 45 81 40.5 5.355
PRL/2*PASTOR 10 MS S 10 MS S 9 9 18 9 8.19
ATTILA*2/CROW 10 MS S 10 MS S 9 9 18 9 8.19
PAVON/SERI 20 MS S 20 MS S 18 18 36 18 7.38
BAVIACORA M 92 5 MS S 10 MS S 4.5 9 13.5 6.75 8.3925
HIDHAB 10 MS S 10 MS S 9 9 18 9 8.19
DHARWAR DRY / SITTA 10 MS S 20 MS S 9 18 27 13.5 7.785
CHAM 6 15 MS S 20 MS S 13.5 18 31.5 15.75 7.5825
PAVON/SERI 30 MS S 40 MS S 27 36 63 31.5 6.165
DHARWAR DRY / NESSER 20 MS S 30 MS S 18 27 45 22.5 6.975
MIRONOVSKA #2//CHIL/CHUM18 30 MS S 40 MS S 27 36 63 31.5 6.165
ATTILA*2/4/CAR//KAL/BB/3/NAC 5 MS S 10 MS S 4.5 9 13.5 6.75 8.3925
KAMBARA1 5 MS S 10 MS S 4.5 9 13.5 6.75 8.3925
ATTILA*2/PASTOR 10 MS S 20 MS S 9 18 27 13.5 7.785
HUW234+LR34*2/AMAD 5 MS S 10 MS S 4.5 9 13.5 6.75 8.3925
ATTILA*2/STAR 20 MS S 50 MS S 18 45 63 31.5 6.165
WEAVER/OCI//BORL95 10 MS S 30 MS S 9 27 36 18 7.38
BABAX/LR42/BABAX 30 MS S 50 MS S 27 45 72 36 5.76
98C017 (NATIONAL LINE) 10 MS 10 MS 8 8 16 8 8.28
2KC033 (NATIONAL LINE) 10 MS 20 MS 8 16 24 12 7.92
99FJ016 (MICROLINE, FATEHJANG) 10 MS 20 MS 8 16 24 12 7.92
3C068 (B-LINE, BARI, CHAKWAL) 30 MS 40 MS 24 32 56 28 6.48
233
TOB/ERA//TOB/CNO67/3/PLO/4/VEE#5/5/KUAZ/6/… 30 MS 40 MS 24 32 56 28 6.48
CHEN/AEGILOPS SQUARROSA (TAUS)//FCT/4/NAC/… 5 MS 10 MS 4 8 12 6 8.46
PJN/BOW//OPATA/3/PASTOR 10 MS 20 MS 8 16 24 12 7.92
CNDO/R143/ENTE/MEXI_2/3/… 5 MS 10 MS 4 8 12 6 8.46
URES/PRL//BAV92 20 MS 30 MS 16 24 40 20 7.2
URES/PRL//BAV92 5 MS 10 MS 4 8 12 6 8.46
TOB/ERA//TOB/CNO67/3/PLO/4/VEE#5/5/KAUZ/6/… 10 MS 20 MS 8 16 24 12 7.92
PASTOR/3/ALTAR 84/AEGILOPS SQUARROSA (TAUS… 20 MS 30 MS 16 24 40 20 7.2
SRMA/TUI//BUBAX 30 MS 40 MS 32 24 56 28 6.48
FN/2*PASTOR 5 MS 10 MS 4 8 12 6 8.46
ATTILA*2/STAR 30 MS 40 MS 24 32 56 28 6.48
BABAX/PASTOR//AMAD 5 MS 10 MS 4 8 12 6 8.46
ATTILA/2*PASTOR 30 MS 40 MS 24 32 56 28 6.48
KALYANSONA 20 MS 30 MS 16 24 40 20 7.2
HUW234+LR34 O 5 MS 0 4 4 2 8.82
TRAP#1.R1/2*SERI.1B 30 MS 40 MS 24 32 56 28 6.48
PRL/2*SERI.1B 20 MS 40 MS 16 32 48 24 6.84
HUW234+LR34*2//PRL/VEE#10 20 MS 20 MS 16 16 32 16 7.56
HUW234+LR34/PRINIA 5 MS 10 MS 4 8 12 6 8.46
HUW234+LR34*2/3/KAUZ*2/TRAP//KAUZ 5 MS 5 MS 4 4 8 4 8.64
ATTILA*2//CHIL/BUC 5 MS 10 MS 4 8 12 6 8.46
ATTILA*2/3/KAUZ*2/TRAP//KAUZ 5 MS 5 MS 4 4 8 4 8.64
ATTILA*2/AMAD 1 MS 1 MS 0.8 0.8 1.6 0.8 8.928
ATTILA*2/PBW65 20 MS 20 MS 16 16 32 16 7.56
ATTILA*2/STAR 20 MS 20 MS 16 16 32 16 7.56
WEAVER/OCI//BORL95 40 MS 40 MS 32 32 64 32 6.12
BABAX/LR42/BABAX 20 MS 40 MS 16 32 48 24 6.84
INQALAB 91 (AARI, FAISALABAD) 10 MS S 40 S 9 40 49 24.5 6.795
CHAKWAL 97 (BARI, CHAKWAL) 10 S 10 S 10 10 20 10 8.1
AS 2002 (AARI, FAISALABAD) 20 S 20 S 20 20 40 20 7.2
MAN 1 (NATIONAL LINE) 20 S 30 S 20 30 50 25 6.75
V 02169 (MICROLINE, FAISALABAD) 20 S 30 S 20 30 50 25 6.75
1C001 (MICROLINE, BARI, CHAKWAL) 20 S 30 S 20 30 50 25 6.75
1C002 (MICROLINE, BARI, CHAKWAL) 20 S 30 S 20 30 50 25 6.75
NR 234 (MICROLINE, NARC, ISLAMABAD) 20 S 30 S 20 30 50 25 6.75
1C007 (MICROLINE, BARI, CHAKWAL) 30 S 30 S 30 30 60 30 6.3
SH 2002 (AARI, FAISALABAD) 40 S 30 S 40 30 70 35 5.85
HAIDER 2000 (CCRI, PIRSABAQ) 50 S 40 S 50 40 90 45 4.95
00BT004 (MICROLINE, BIO.TECH.FSD) 50 S 50 S 50 50 100 50 4.5
MARGALA 99 (BARI, CHAKWAL) 50 S 60 S 50 60 110 55 4.05
V 02166 (MICROLINE, FAISALABAD) 70 S 80 S 70 80 150 75 2.25
3C067 (B-LINE, BARI, CHAKWAL) 90 S 100 S 90 100 190 95 0.45
00C010 (NATIONAL LINE) 100 S 100 S 100 100 200 100 0
Katila II 10 S 10 S 10 10 20 10 8.1
101/25 30 S 30 S 30 30 60 30 6.3
PASTOR/3/ALTAR 84/AEGILOPS SQUARROSA (TAUS… 60 S 80 S 60 80 140 70 2.7
URES/JUN//KAUZ/3/BABAX 50 S 70 S 50 70 120 60 3.6
LOCAL CHECK 50 S 60 S 50 60 110 55 4.05
BABAX/LR39/BABAX 40 S 50 S 40 50 90 45 4.95
KAMBARA1 20 S 40 S 20 40 60 30 6.3
234
Appendix VII A Wheat Stripe Rust Development Influenced by Environmental Factors
Prevailed at Rawalpindi during 2006
Months
Environmental Factors
Mean Temperature (oC) Rainfall
(mm) Humidity
(%) Minimum Maximum
January 3.52 10.94 62.20 70.35
February 9.69 17.35 25.60 61.41
March 11.44 18.81 45.50 62.10
April 15.33 24.05 20.30 37.32
May 20.05 31.07 62.90 36.00
Appendix VII B Wheat Stripe Rust Development Influenced by Environmental Factors
Prevailed at Rawalpindi during 2007
Months
Environmental Factors
Temperature (oC) Rainfall
(mm) Humidity
(%) Minimum Maximum
January 1.05 10.41 00.50 62.68
February 6.63 12.99 92.30 75.93
March 9.06 16.09 143.20 64.68
April 15.90 24.91 19.60 44.37
May 19.83 28.59 79.60 42.05
235
Appendix VIII PRECIPITATION RECEIVED AT RAWALPIN DI DURING 2006 AND 2007
RAINFALL (mm) 2006 RAINFALL (mm) 2007
Jan Feb March April May Days Jan Feb March April May
12.4 0 0 0 0 1 0 0 3.2 13.4 0
16.3 0 0 0 0 2 0 0 0 0 0
0 0 0 0 0 3 0 0 0.2 0 0
0 0 0 0 0 4 0 0 14.4 0 2.6
0 0 0 0 0 5 0 0 0 0 0
0 0 0 0 0 6 0 0 0 0 0
0 0 0 0 0 7 0 0 0 0 0
0 0 0 0 0 8 0 0 0 0 1.8
0 0 0 20.3 0 9 0 0 0.1 0 0
0 0 0 0 0 10 0 27.7 3.4 0 0
0 0 0 0 0 11 0 33 1.1 0 0
0 0 0 0 0 12 0 10.4 31.9 0 0
0 0 16.6 0 0 13 0 7.3 7.9 0 0
0 0 9.3 0 0 14 0 1 0 0 0
1.2 3.6 0 0 0 15 0 0 0 0 0
13.2 0 0 0 0 16 0 0.8 0 0 36
20.1 0.3 0 0 14 17 0 0 0 0 0
0 0 0 0 0 18 0 0.2 0 0 0
0 0 6.3 0 27.8 19 0 0 0 0 37
0 0 5.9 0 9.2 20 0 0 59.3 0 2.2
0 0 0 0 10.5 21 0 2.4 21.7 0 0
0 0 0 0 0 22 0 0 0 0 0
0 0 0 0 0 23 0 0 0 6.2 0
0 0 0 0 0 24 0 0 0 0 0
0 13.6 0 0 1.2 25 0 0 0 0 0
0 8.1 7.3 0 0 26 0 0 0 0 0
0 0 0.1 0 0 27 0.5 6.7 0 0 0
0 0 0 0 0.2 28 0 2.8 0 0 0
0 0 0 0 0 29 0 0 0 0 0
0 0 0 0 0 30 0 0 0 0 0
0 0 0 0 0 31 0 0 0 0 0
63.2 25.6 45.5 20.3 62.9 Total 0.5 92.3 143.2 19.6 79.6
236
Appendix IX A TEMPERATURE RANGE IN RAWALPINDI D URING 2006
Temperature (0C) Min. 2006 Temperature (0C) Max. 2006
JAN FEB MARCH APRIL MAY Days JAN FEB MARCH APRIL MAY
4.5 8.8 8.3 15 23.1 1 8.9 16.6 18.2 23.1 30
6.5 5 7.6 14.5 24.5 2 8.6 14.4 18.3 22.5 30.1
4.8 5.5 8.6 14.7 18.4 3 9.9 15.6 18.2 23.3 27.4
2.2 5.7 10.5 14 18.5 4 8.7 16.1 20.6 22.6 29
0.2 6.5 10 13.7 20.9 5 9.6 16.9 17.1 23.5 30.7
-0.3 7.5 13.1 15 21.6 6 7.8 14.1 19.8 24.1 31.7
-1.2 7.6 11 14.8 23.2 7 9.2 17.2 18.8 24.9 32.7
0.5 7.3 9.3 14.4 25.1 8 10.9 17 18.5 23.5 32.9
1.7 8.1 10 14.8 25 9 8.8 18.3 19.1 20.4 31.6
3 7.2 10.5 9.5 22.8 10 11.9 17.5 20 16.8 31
0.5 7.5 13.5 13.5 20.3 11 11.1 17.8 19.4 19.1 30.6
2.5 10.8 14.8 10.5 22.3 12 12.1 18.7 20.8 18.5 31.9
-7 10.5 15.1 10.7 21.5 13 5.6 18.4 20.4 20.3 32
2.8 12 12.7 11.6 29.2 14 11.3 17 15.7 19.9 35.3
5.2 11.6 11.7 16.7 25.5 15 9.4 17.8 16.8 23.1 32.5
9.5 9.5 9.3 13.4 26.1 16 14.7 18 16.5 23.1 32.7
11.2 12 8.2 12.5 21 17 12.6 19.1 16.1 21.8 27.4
1.3 13.7 9.1 12.6 23.7 18 9.3 19.8 17.7 22.1 30.6
1 15.1 12 13.5 18.5 19 9.9 19.4 17.9 23 26
3.1 10.2 12.5 15 23 20 10.8 18.6 15.1 23.9 30.8
3.7 12.2 9 16.7 18 21 10.1 19.9 16.9 24.8 27.1
5 10.1 10.2 17.5 24.5 22 11.4 16.5 18.9 25.9 30.5
3.1 11 12.2 20.9 22.8 23 10.3 18.8 20.1 28.5 29.8
1.8 12.3 14.1 17.5 24.8 24 10.8 17.5 19.2 27 31.4
2 14.8 14 16.5 23.9 25 11.6 16.4 19.4 27.6 32
4.2 12 12.9 18.4 23.5 26 13.1 15.3 19.8 28.8 32.4
6.6 8.6 12 18.1 24.1 27 12.5 16.2 17.5 29.5 33.2
8.3 8.2 11.6 18.5 24.6 28 14.6 17 19.4 29.4 32.6
6.3 0 12 20.4 24.6 29 12.4 0 20.5 29.8 31.6
7.5 0 13.3 25 24 30 15.3 0 22.4 30.6 33
8.5 0 15.5 0 25.5 31 16 0 24.1 0 32.8
3.52 9.69 11.44 15.33 20.05 Mean 10.94 17.35 18.81 24.05 31.07
237
Appendix IX B TEMPERATURE RANGE AT RAWALPINDI DU RING 2007
Temperature (0C) Min. 2007 Temperature (0C) Max. 2007
JAN FEB MARCH APRIL MAY Days JAN FEB MARCH APRIL MAY
2.3 3.9 7.8 15.3 20.5 1 10 14.2 13.8 20.3 29.4
1.1 5.4 4.6 11 22.4 2 9.2 14.6 12.6 21.1 29.8
-1.5 7.1 7.9 10.5 23.6 3 7.8 15.4 12.1 18.7 29.5
-1.7 8.4 8.1 12.9 19.8 4 7.3 15.3 12.3 21.6 27.3
-1.6 6.1 2.7 13.5 17.6 5 7.3 13.5 11.4 21.6 26.7
-2.7 9.6 4.4 13.1 18.2 6 6.7 16.5 13.1 21.5 28
-2.3 6.3 4.8 13.5 19.5 7 7.1 14.6 14.3 22.1 29
-2.3 5.9 5.7 13.3 19.8 8 8.3 14 15.5 22.3 27
-0.5 6.7 9.1 13.5 20.7 9 9 13.2 14.2 22.5 28.2
0.9 10 10.1 12.5 18.7 10 10.1 12.6 13.3 22.2 27.8
-0.3 9.5 7.9 12.8 19.9 11 9 10.7 12 23.5 28
-1.3 8 7.1 14.5 19.9 12 9.1 10.7 10.3 25.1 28.4
-1.4 7.9 7.5 16.3 20.2 13 7.7 11.7 13.6 26.3 29
-1.5 8.3 5.8 17.1 23.4 14 7.8 11.8 12.5 26.5 31.3
-1.6 3.7 4.1 16.8 24.4 15 8.3 10.2 12.2 26.9 32.6
-0.6 4.7 7.7 17.5 17.3 16 9.2 12.3 15 27.6 26.1
-0.8 5.2 8.5 21.2 20.8 17 7.7 12.5 17.1 28.6 29.4
-0.5 5.3 12.1 21 21.7 18 6.4 9.5 18.5 26.9 31.2
2.8 4.1 14.5 20.2 18 19 10.6 11.2 20.7 27.5 26.3
1.2 5.2 13.2 16.4 19.5 20 10.4 13.2 16.9 25.1 26.9
-0.1 7.9 11.8 15.9 19.2 21 9.7 10.5 15.3 25.6 27.7
-0.6 4.4 7.4 16.2 21.6 22 10 12.3 16.3 26.2 29.4
-0.7 3.9 9.2 19.3 18.8 23 10.7 13.4 17.2 26.5 26.8
1.8 4.6 10.8 16.4 20.5 24 11.8 13.6 17.9 25.5 29.1
4.5 7.4 8.8 16.9 19.4 25 14.7 15.6 18.3 25.6 29.2
5.3 8.3 10.1 16.6 17.2 26 14.8 15 18.5 26.2 27.7
10.4 9.8 10 15.3 18 27 17.8 12 19.9 26.6 25.3
7.7 7.9 11.9 16.1 19 28 16.8 13.6 21.7 27.6 30
5.4 0 14.7 18.9 17.5 29 16 0 23.6 29.1 30.1
6.7 0 15.4 22.6 18.7 30 16.3 0 24.3 30.6 31.3
4.5 0 17.1 0 19 31 15 0 24.5 0 27.9
1.05 6.63 9.06 15.90 19.83 Mean 10.41 12.99 16.09 24.91 28.59
238
Appendix X HUMIDITY PERCENTAGE AT RAWALPINDI DUR ING EXPERIMENTATION YEARS
HUMIDITY (%) 2006 HUMIDITY (%) 2007
JAN FEB MARCH APRIL MAY Days JAN FEB MARCH APRIL MAY
83.5 63 54.5 53 30.5 1 77 56 66 60.5 38
92 63 54.5 43.5 33.5 2 69.5 65 57 45 43
85 56 51 43 29 3 62.5 60.5 77 61 42
71.5 49.5 54.5 40.5 30 4 63 71.5 85.5 65.5 59
68.5 53.5 59 41.5 23 5 61 79.5 68.5 56 37
69.5 68 46.5 34 25.5 6 62.5 66.5 57 57.5 29.5
65 51 52.5 32.5 28.5 7 66 64 56.5 51 41
63.5 57.5 58 37.5 31.5 8 69 73 61.5 50.5 47
82.5 55.5 57 85 32 9 63 80.5 68.5 41 37
65.5 50 53 27 28 10 65 86.5 79.5 37 36.5
65.5 49 68 36.5 27.5 11 62 94 84.5 44.5 35
62 50.5 65.5 38 26 12 56 91 93 44 33
70.5 58.5 74 32.5 26.5 13 64 89.5 70 44.5 38
65 59.5 87.5 40.5 32 14 55 88 67 46 46
84.5 71 78 43.5 31 15 55 81.5 56.5 44 37.5
82.5 63 66 40.5 36.5 16 60 73.5 55 40.5 51
90 64.5 57 37.5 46.5 17 63 72 61.5 43 38.5
73.5 66.5 54.5 37 44.5 18 75.5 83.5 69 46 48.5
67 68.5 82.5 36.5 56 19 71 71.5 66 46.5 54.5
67.5 66 86.5 38 70.5 20 63 75 85.5 41 38
68 55.5 73 38.5 49.5 21 58.5 91 75.5 42.5 44
71.5 61.5 57.5 36 49 22 52.5 78.5 66 43.5 39
66.5 61 54 35 48.5 23 64 66 59.5 39.5 39.5
65 67.5 72.5 33.5 43 24 64 64.5 58.5 36 29
56 91 64.5 29 37.5 25 61.5 64 57.5 36.5 45
55 74 75.5 23.5 33.5 26 62 73.5 48.5 37 56.5
71.5 68 70 27 32 27 61.5 88 47 32.5 45
55 57 55.5 22.5 34 28 58 78 45 31.5 46.5
73 0 50 28.5 40 29 63 0 54.5 36.5 47.5
64.5 0 50.5 28 29 30 60.5 0 57 30.5 42.5
60.5 0 42 0 31.5 31 54.5 0 51 0 39
70.35 61.41 62.10 37.32 36.00 Avg 62.68 75.93 64.68 44.37 42.05
239
Appendix XI Disease Severity Percentage using Cobb’s scale during 2005-06
Varieties Replicates
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 10 15 10 15 15 15 10 15 Inquilab 91 5 10 10 10 10 5 5 10 Wafaq-2001 25 30 20 25 25 25 25 25 Morocco 50 55 55 55 60 50 50 50
Appendix XII Disease Severity Percentage using Cobb’s scale during 2006-07
Varieties Replicates
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 20 20 25 20 25 20 25 20 Inquilab 91 35 30 40 40 35 35 40 30 Wafaq-2001 30 30 35 30 30 35 30 35 Morocco 50 60 55 80 55 50 50 50 Appendix XIII 1,000 grain weight of wheat in control plots during 2005-06
Varieties Control
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 41.98 43.52 42.75 43.30 42.80 41.70 42.80 43.20 Inquilab 91 44.30 43.80 45.70 45.50 44.70 43.00 43.35 45.60 Wafaq-2001 45.82 44.88 45.35 45.00 45.80 46.20 45.20 46.70 Morocco 42.10 42.80 41.70 42.30 42.90 44.20 42.80 41.70 Appendix XIV 1,000 grain weight of wheat in diseased plots during 2005-06
Varieties Replicates
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 34.00 30.95 31.96 30.20 34.50 34.30 32.19 32.22 Inquilab 91 36.20 32.19 34.30 33.30 34.70 35.60 36.10 34.20 Wafaq-2001 26.70 27.30 29.30 25.30 28.80 28.10 28.70 27.90 Morocco 18.30 13.72 15.60 14.30 13.72 14.81 14.38 18.00 Appendix XV 1,000 grain weight of wheat in control plots during 2006-07
Varieties Control
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 42.0 40.0 42.1 44.0 45.9 43.1 40.0 43.2 Inquilab 91 43.3 40.0 40.0 42.7 41.6 41.0 41.0 42.0 Wafaq-2001 44.0 41.1 41.8 46.0 45.5 49.2 43.2 48.7 Morocco 43.6 43.0 41.4 41.0 43.9 48.0 43.8 41.1
240
Appendix XVI 1,000 grain weight of wheat in diseased plots during 2006-07
Varieties Replicates
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 21.5 20.1 24.3 23.3 27.1 24.5 22.0 25.6 Inquilab 91 37.4 30.6 30.6 30.6 35.3 33.6 36.3 34.5 Wafaq-2001 22.7 22.0 25.1 24.3 25.3 28.5 23.2 25.8 Morocco 17.7 13.6 16.2 12.0 14.2 18.4 14.7 14.8 Appendix XVII Wheat yield (kg per hectare) in control plots during 2005-06
Varieties Yield in control plots
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 1970 1925 1889 1928 1915.2 1919.6 1949.2 1928 Inquilab 91 2540 2300 2600 2470.2 2469 2489.2 2495.8 2475.8 Wafaq-2001 1540 1456 1560 1518.3 1534 1506.8 1516.6 1517.7 Morocco 1756 1690 1745 1730.8 1733.6 1699.8 1745 1742
Appendix XVIII Wheat yield (kg per hectare) in diseased plots during 2005-06
Varieties Yield in diseased plots
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 1654 1623 1645 1636.5 1629.2 1647.9 1667.8 1622.2 Inquilab 91 2380 2430 2200 2338.1 2320.6 2359.5 2343.3 2322.3 Wafaq-2001 1434 1369 1451 1437.9 1411.4 1416.9 1418.3 1405.5 Morocco 1124 1078 923 1025.4 1008.6 1035.4 1077.4 1062.1 Appendix XIX Wheat yield (kg per hectare) in control plots during 2006-07
Varieties Yield in control plots
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 1924.7 1910.4 1916.5 1910.0 1917.1 1908.1 1935.2 1919.0 Inquilab-91 2347.1 2298.7 2289.2 2334.4 2292.1 2275.1 2297.9 2349.8 Wafaq-2001 1490.0 1498.0 1545.0 1503.0 1470.0 1514.2 1476.1 1485.5 Morocco 1765.0 1722.4 1775.0 1790.0 1775.9 1725.0 1746.0 1761.0 Appendix XX Wheat yield (kg per hectare) in diseased plots during 2006-07
Varieties Yield in diseased plots
R1 R2 R3 R4 R5 R6 R7 R8 Bakhtawar 1272.3 1241.1 1099.0 1238.1 1120.3 1196.2 1165.0 1167.3 Inquilab-91 2313.5 2305.4 2240.3 2238.5 2245.5 2254.7 2239.9 2336.7 Wafaq-2001 1276.5 1255.8 1236.4 1268.2 1276.2 1233.6 1284.1 1225.9 Morocco 1054.2 1004.4 1018.7 996.8 1039.6 1038.8 1061.9 1037.75
241
