Journal of Phytopathology and Pest Management 7(1): 1-13, 2020

pISSN:2356-8577 eISSN: 2356-6507

Journal homepage: http://ppmj.net/

∗

Corresponding author:

Abdelrazek S. Abdelrhim,

E-mail: abdelrazek.sharawy@mu.edu.eg

1

Molecular detection of stripe rust resistance

genes in some Egyptian bread wheat cultivars

and promising spring wheat lines

Samar M. Esmail

1

, Abdelrazek S. Abdelrhim

2*

1

Wheat Diseases Research Department, Sakha Agriculture research station, Institute of Plant Pathology,

Agriculture Research Centre (ARC), Sakha, Kafrelsheikh, 33717, Egypt

2

Department of Plant Pathology, Faculty of Agriculture, Minia University, Minia, Egypt

Abstract

Keywords: Puccinia striiformis f.sp. tritici, Yr genes, spring bread wheat cultivars.

2

1. Introduction

Wheat is one of the most important cereal

crops. It comes directly after corn and

rice in terms of world production. It is the

most important staple food for 30% of

the world population. It occupies more

than 220 million hectares of land with

725 million tons production

(Anonymous, 2015). More than 65% of

world population is mainly relying on

wheat as an important source for

nutrition. Which provides 21% of the

food calories consumed globally, and

20% of the protein to more than 4.5

billion people in 94 developing countries

(Braun et al., 2010). In Egypt, wheat is an

important staple food with an annual

consumption of about 12.4 million tons,

of which about 9.4 million tons are

grown locally (Rashed et al., 2016).

Wheat is liable to be infected by three

types of rusts; stem, stripe, and leaf rust.

Stripe rust considers the most important

diseases in wheat around the world such

as Asia, Europe, the Middle East, and

Africa (Chen, 2005; McIntosh & Brown

1997). Stripe rust may cause high yield

loss, that could reach up to 70% in usual

epidemic years and 100% in severe

epidemic years (Li & Zeng, 2002). In

Egypt, the first epidemic of stripe rust

was recognized on wheat varieties Giza

144 at Manzala during 1967/68 (Abdel-

Hak et al., 1972). Then starting from

1995 till 1998, it caused a sever loss on

wheat cultivars Sakha-69, Giza-163,

Gemmeiza-1, and most of the

commercial varieties, especially the long-

spiked ones, at the Northern governorates

in particular. A little bit cases of

epiphytotics were recorded during

1999/2000 (Abu El-Naga et al., 2001;

1999; 1997; El-Daoudi et al., 1996).

Among all rust management strategies,

breeding for resistant cultivars is the

most effective, economical, and

environment friendly strategy (Line &

Chen, 1995). Recently, 70 officially

designated and 100 temporarily named

stripe rust resistance genes and

quantitative trait loci (QTLs) have been

reported (Dracatos et al., 2016; Cheng et

al., 2014; McIntosh et al., 2013).

However, emerging of new races has

overcome most of these resistance genes,

in different parts of the world, especially

when these genes start to be widely used

in wheat breeding and production (Chen

et al., 2003; Li & Zeng, 2002; He et al.,

2001).

In Egypt, the source of

Pts

spores

is not accurately known. Therefore,

pyramiding of various stripe rust

resistance genes is required to protect the

growing wheat cultivars against the

domestic and the new arrival races. On

the other hand, the genetic background of

most commercial Egyptian wheat

cultivars and advanced lines is not well

studied, in terms of resistance to stripe

rust. The aim of this study is to evaluate

the response of 32 wheat genotypes,

including Egyptian commercial wheat

cultivars and promising breeding lines

against three of the most prevalent

Pst

physiological races 224E132, 224E32,

and 151E80. Also detecting the presence

of six

Yr

resistance genes,

Yr5

,

Yr10

,

Yr15

,

Yr17

,

Yr18

, and

Yr26

in tested

wheat genotypes using STS markers and

SSR markers.

2. Materials and methods

2.1 Wheat materials

A total of 32 wheat entries (Table 1)

3

including; 15 contemporary Egyptian

wheat cultivars; five of the most

promising, advanced breeding lines

which were selected upon their

performance in the wheat breeding

program, at Sakha Agriculture research

station, Egypt; and 12

Yr

isogenic lines

were tested for their seedling and adult-

plant responses to the most prevalent

Egyptian

Pst

races, 224E132, 224E32,

and 151E80. In addition to wheat cultivar

Avocet S, which is susceptible to most of

Pst

races, was selected as the susceptible

control.

Table 1: Pedigrees of Egyptian bread wheat genotypes tested for stripe rust resistance.

Accessions and original sources

Pedigree

Egyptian bread wheat cultivars

Sakha-08

Cultivar

CIANO-67(SIB)//SONORA-64/KLEIN-RENDIDOR/3/II-8156; INDUS-66/(SIB)NORTENO;

Sakha-93

Cultivar

Sakha 92/TR 810328 S 8871-1S-2S-1S-0S

Sakha-94

Cultivar

Opata/Rayon//Kauz CMBW9043180-OTOPM-3Y-010M-010M-010Y-10M-015Y-0Y

Sakha-95

Cultivar

PASTOR // SITE / MO /3/ CHEN / AEGILOPS SQUARROSA (TAUS) // BCN /4/ WBLL1

Giza-160

Cultivar

(Regent975-11Giza1392)MidaCadetHindi 62

Giza-168

Cultivar

MIL/BUC//Seri CM93046-8M-0Y-0M-2Y-0B

Giza-171

Cultivar

Gemmeiza-9 / Sakha-93

Sids-12

Cultivar

BUC//7C/ALD/5/MAYA74/ON//1160-147/3/BB/GLL/4/CHAT"S"/6/MAYA/VUL-4SD-1SD-1SD-0SD.

Sids-13

Cultivar

KAUZ "S"//TSI/SNB"S". ICW94-0375-4AP-2AP-030AP-0APS-3AP-0APS-050AP-0AP-0SD.

Sids-14

Cultivar

Bow''s''/Vee''s''//Bow's'/Tsi/3/BANI SUEF 1

Gemmeiza-11

Cultivar

BOW ‘‘S’’/KVZ ‘‘S’’//7C/SERI82/3/GIZA168/SKHA61

Gemmeiza-12

Cultivar

BUC//7C/ALD/5/MAYA74/ON//1160-147/3/BB/GLL/4/CHAT ‘‘S’’/6/MAYA/VUL//CMH74A.630//4*SX

Misr-1

Cultivar

OASIS/SKAUZ//4*BCN1312*PASTOR

Misr-2

Cultivar

SKAUZ/BAV92

Misr-3

Cultivar

ATTILA*2/PBW65*2/KACHU

Breeding lines

Line-1

Breeding Line

FLORKWA -2 /10/ MAYA /YD/6/HK/MDA38/4/4777/3/REI//Y/KT/5/YR/7/KOEL/8/MOR/BOW/9/SERI/11/ATTILA*2/GIZA 168

Line-2

Breeding Line

SIDS1/ ATTILA // GOUMRIA-17 (PBW 343 /6/ SHA7 / VEE#5 /5/ VEE#8 // JUP / BJY /3/ F3.71 / TRM/4 / 2*WEVER )

Line-3

Breeding Line

SW89.5181/KAUZ/4/MILAN/KAUZ//PRINIA/3/BABAX

Line-4

Breeding Line

CHEN/AEGILOPS SQUARROSA(TAUS)/ /BCN/3/2* KAUZ/4/GEN*2 //BUC/ FLK /3/ BUCHIN.

Line-5

Breeding Line

SIDS1/ ATTILA // GOUMRIA-17 (PBW 343 /6/ SHA7 / VEE#5 /5/ VEE#8 // JUP / BJY /3/ F3.71 / TRM/4 / 2*WEVER )

Table 1 (Supplementary): Differential genotypes

*

used to identify

pathotypes of stripe rust incited by Puccinia striiformis f.sp. tritici in

Egypt.

Differential cultivars

Abbreviation

Decanery value

Resistance gene

Type

GI. World differential set

Chinese 166

Ch

(2

0

) =1

Yr1

Winter

Lee

(2

1

) = 2

Yr7

Spring

HeinesKolben

HK

(2

2

) = 4

Yr2Yr6

Spring

Vilmorin 23

V23

(2

3

) = 8

Yr3

Winter

Moro

Mo

(2

4

) = 16

Yr10

Winter

StrubesDickkopf

Std

(2

5

) = 32

SD

Winter

Suwon 92 × Omar

Su

(2

6

) = 64

SU

Winter

Clement

Cl

(2

7

) = 128

Yr2Yr9

Winter

Tirticumspelta Album

Sp

(2

8

) = 256

Yr5

Spring

GII. European Differential set

Hybrid 46

H46

(2

0

) =1

Yr4

Winter

Reichersberg 42

R42

(2

1

) = 2

Yr(7)

Winter

Heines Peko

Pe

(2

2

) = 4

Yr2 Yr(6)

Spring

Nord Desprez

No.D

(2

3

) = 8

Yr(3)

Winter

Compare

Com

(2

4

) = 16

Yr8

Spring

Carstens V

CV

(2

5

) = 32

YrCV

Winter

Spaldings Prolific

Spa

(2

6

) = 64

YrSP

Winter

Heines VII

HVII

(2

7

) = 128

Yr2

Winter

*

Johnson et al., (1972).

2.2

Puccinia striiformis

f.sp.

tritici

races

The most prevalent Egyptian

Pst

races

224E132 and 224E32, and the new race

151E80 were identified previously using

the World and European group of wheat

differential varieties as proposed by

Johnson et al. (1972) (Supplementary

Table 1). The selected races were

carrying different virulence genes that

make them a good material for testing

the major and adult plant resistance of

tested wheat genotypes.

2.3 Seedling evaluation

This experiment was conducted under

4

greenhouse condition, as described by

Li

and Zeng (2002).

Five seeds of each

genotype were planted in plastic pots,

with both a diameter and height 10 cm,

with five replicates. Seedlings of tested

wheat genotypes were inoculated at the

two-leaf stage, with the fresh

urediniospores of

Pst

races 224E132,

224E32 and 151E80. After inoculation,

the seedlings were incubated in a dew

chamber at 10-12°C and 100% relative

humidity under dark conditions for 12 h.

Then, infected seedlings were moved to a

separate greenhouse chamber maintained

at 12–17°C and seedlings were subjected

to 16h light and 8 h darkness. Seedling

infection types (IT) were recorded 15–18

days after inoculation, following 0–9

scale (Line & Qayoum, 1992). Plants

with IT 0-3 were considered resistant, ITs

of 4-6 were considered intermediate, and

ITs of 7-9 were considered susceptible.

2.4 Field evaluation

All the tested wheat genotypes were

evaluated in adult plant stage, at the

experimental farm of Sakha Agriculture

research station, Egypt during 2017/2018

and 2018/2019 growing season. The

experiment was designed as a

randomized complete block. Three

replicates were used for each genotype.

Each replicate included 3 rows.

Approximately 20 seeds, of each wheat

genotype, were sown in a 1-m row with

20 cm distance between rows. The

susceptible cultivar Morocco was planted

in one row for every 20 rows and around

the blocks as a stripe rust spreader.

During early March, Morocco was

inoculated with a mixture of

urediniospores of 224E132, 224E32 and

151E80 as described by Li and Zeng

(2002).

At late tilleringstage all tested

wheat genotypes were uniformly dusted

with urediniospores of three races’

mixture and talcum powder at the rate of

1:25 (v/v).Infection response was scored

as described by Roelfs et al. (1992),

resistant (R), moderately resistant (MR),

moderately susceptible (MS) and

susceptible (S). Disease severities were

assessed based on the percentage of leaf

area affected (0, 1, 5, 10, 20, 30, 40, 60,

80 and 100%) (Li & Zeng, 2002). The

Coefficient of Infection (CI) for stripe

rust were calculated according to Akhtar

et al. (2002),

by multiplying the response

value with the intensity of infection in

percent. The response value was

determined as following: 0.0 =no

disease, 0.2 =R, 0.3 =R to MR, 0.4 =MR,

0.6 =MR-MS, 0.8 =MS, and 0.9 MS-S.

Average coefficient of infection (ACI)

was derived from the sum of CI values of

each entry divided by the number of

tested years. The highest ACI of a

candidate line is set at 100 and all other

lines are adjusted accordingly. This gives

the Country Average Relative Percentage

Attack (CARPA). The ‘0’ to ‘9’ scale

previously designated as Resistance

Index (R.I) has been re-designated as

RRI (Relative Resistance Index). RRI

was calculated from CARPA, on a 0 to 9

scale, where 0 denotes most susceptible

and 9 denotes highly resistant (Akhtar et

al., 2002). The RRI was calculated

according to the following formula:

 

  



 

2.5 DNA extraction and molecular

marker detection of

Yr

genes

All the tested wheat genotypes were

5

grown in a rust-free greenhouse. Two

weeks later approximately 100 mg leaf

tissue of each wheat entry was collected.

DNA was extracted using a

cetyltrimethylammonium bromide

(CTAB) method (Saghai-Maroof et al.,

1984). The DNA stock solution was

adjusted to a concentration of 50 ng/µl

with sterilized ddH2O for use as the

working solution for the polymerase

chain reaction (PCR). The eluted DNA

was stored at -20ºC. PCR mixture for

Yr

genes detection. PCR reaction was

conducted in reaction volume of 25 µl;

contained (1 µl) of 25 ng nucleic acid, 1

µl of each primer (10 pmol), 12.5 µl of

Master Mix (Promega Corporation, USA)

and 9.5 µl of Nuclease free water

(Promega). Three Sequence-Tagged Sites

(STS) markers and three Simple

Sequence Repeat (SSR) markers were

used for detection of (

Yr5

,

Yr15

, and

Yr26

) and (

Yr10

,

Yr17

, and

Yr18

),

respectively (Table 2).

Then 15 µl of all

PCR products were analyzed by

electrophoresis through a 1.5% agarose

gel, stained with ethidium bromide. DNA

bands were visualized using a UV

transilluminator. The PCR and

electrophoresis of process were carried

out as described by Murphy et al. 2009,

Singh et al. 2009, Wang et al. 2008, Wen

et al. 2008 and Lagudah et al. 2006, 2009

with minor modifications.

Table 2: Sequences of primers, PCR conditions and references for Yr marker used to identify six Yr genes in wheat

cultivars and lines.

Wheat cultivars. Gene

Marker type

Marker name

Primer sequence

GC %

Ta (°C)

Fragment size (bp)

Reference

Yr5

STS

STS7

F5'-GTACAATTCACCTAGAGT-3'

39

45

478

Chen et al. (2003)

STS8

R5'-GCAAGTTTTCTCCCTAT-3'

41

Yr10

SSR

Yr10-F11

F5'-TTGGAATTGGCGACAAGCGT-3'

53

64

755

Singh et al. (2009)

Yr10-R11

R5'-GTGATGATTACCCACTTCCTC-3'

50

Yr15

STS

Xa1LRF

5´-CTCACTCTCCTGAGAAAATTAC-3´

41

56

700

Murphy et al. (2009)

PtoFen-S

5´- ATGGGAAGCAAGTATTCAAGGC-3

45

Yr 17/Sr38/Lr37

SSR

VENTRUIP

F5'-AGGGGCTACTGACCAAGGCT-3'

60

65

259

Helguera et al. (2003)

LN2

R5'-TGCAGCTACAGCAGTATGTACACAAAA-3'

40

Yr18/Pm38/Lr34

SSR

L34DINT9-F

F5-’TTGATGAAACCAGTTTTTTTTCTA-3’

25

58

517

Lagudah et al. 2009

L34PLUS-R

R5’-GCCATTTAACATAATCATGATGGA- 3’

33

Yr26

STS

WE173

F5’-GGGACAAGGGGAGTTGAAGC-3’

60

55

259

Wang et al. (2008)

R5’-GAGAGTTCCAAGCAGAACAC-3’

50

3. Results

3.1 Seedling evaluation

Three

Pst

races 224E132, 224E32, and

151E80 were used for seedling

evaluation of 32 wheat genotypes under

greenhouse condition (Table 3).Among

all tested wheat cultivars only three

cultivars Sakha-93, Sakha-95, and Misr-3

were showed high to intermediated

resistance against

Pst

races. However,

two cultivars Sakh-94 and Gemmiza-12

were showed intimidated resistance to

races 224E32 and 151E80, but both

cultivars were susceptible to race

224E132. In addition, among five

breeding lines only two lines, Line-2 was

showed high resistance, while Line-5

was expressed intermediate resistance to

the tested

Pst

races. The tested

Yr

NILs

varied for their responses to the three

races.

Yr5

and

Yr15

were showed high

resistance against all tested races,

compared with other NILs. However,

Yr1

,

Yr7

,

Yr10

, and

Yr26

were expressed

low to intermediate infection types to

only two races 224E132 and 224E32,

and high infection types for race 151E80.

The susceptible cultivar Avocet S was

showed high infection types against all

races.

6

3.2 Adult plant stage evaluation

The field experiment was implemented to

study the response of wheat genotypes

against the mixture of three

Pst

races,

(224E132, 224E32, and 151E80), in adult

plant stage. According to the obtained

data two cultivars Sakha-95 and Misr-3

were showed high resistance to

Pst

races,

and low disease severity was observed on

both cultivars during 2017/18 and

2018/2019. Also, Sakha-95 and Misr-3

were expressed high RRI value 8.86.

Four cultivars Sakha-93, Sakha-94, Misr-

1, and Misr-2 were expressed moderated

resistance, with low disease severity less

than 20, and IRR values between 6.57

and 8.64. However, all other cultivars

were showed moderated to high

susceptibility to the

Pst

races in both

growing seasons 2017/18 and 2018/2019.

Four breeding lines, Line-2, Line-3,

Line-4, and Line-5, were showed high

resistance with disease severity 20 and

less, and IRR 8.86 for line-2 and Line-5

and 8.73 for Line-3 and Line-4.

However, Line-1 was moderated

resistance to the

Pst

races with 30%

disease severity, IRR values for both

lines were 7.92. Among twelve tested

isogenic lines only four lines,

Yr1

,

Yr5

,

Yr10

, and

Yr15,

were expressed high

resistance. The same lines were showed

low disease severity less than 10, ACI

less than 1.5, and IRR ranged from 8.86

to 8.91during 2017/2018 and 2018/2019

growing seasons. Moderated resistance

was observed on

Yr18

against

Pst

races

with disease severity less than 20 in

2017/2018 and IRR 8.46.

Table 3: Responses of wheat genotypes to three Puccinia striiformis f. sp. tritici races in seedling

and adult plant stages.

Wheat genotype

Seedling stage

Adult stage

CI

ACI

RRI

2017/18

2018/19

224E132

224E32

151E80

IT

DS

IT

DS

Egyptian wheat cultivars

Sakha-8

9

S

70

S

80

150

75

2.25

Sakha-93

5

4

3

MR

5

MS

30

26

13

7.83

Sakha-94

7

6

MR

10

MR

10

8

4

8.64

Sakha-95

4

3

4

R

5

R

10

3

1.5

8.86

Giza-160

9

S

80

S

90

170

85

1.35

Giza-168

6

7

8

MS

20

S

5

21

10.5

8.05

Giza-171

7

8

MS

10

S

30

38

19

7.29

Sids-12

9

S

80

S

80

160

80

1.8

Sids-13

9

8

S

70

S

70

140

70

2.7

Sids-14

9

7

MS

30

S

40

46

23

6.93

Gemmiza-11

9

S

70

S

80

150

75

2.25

Gemmiza-12

8

6

MS

30

S

30

54

27

6.57

Misr-1

7

8

MR

10

S

30

34

17

7.47

Misr-2

7

MR

20

MS

30

32

16

7.56

Misr-3

3

4

R

5

R

10

3

1.5

8.86

Wheat breeding lines

Line-1

8

MR

30

MR

30

24

12

7.92

Line-2

3

4

R

5

R

10

3

1.5

8.86

Line-3

8

7

8

R

10

R

20

6

3

8.73

Line-4

7

8

R

10

R

20

6

3

8.73

Line-5

6

R

5

R

10

3

1.5

8.86

Yr-NILs

Yr1

0;

8

R

5

R

5

2

1

8.91

Yr5

0;

1

0;

R

5

R

5

2

1

8.91

Yr6

9

3

8

S

80

S

80

160

80

1.80

Yr7

4

9

S

40

S

60

100

50

4.50

Yr9

8

9

8

S

30

S

70

100

50

4.50

Yr10

1

0;

8

R

5

R

10

3

1.5

8.86

Yr15

3

2

3

R

5

R

5

2

1

8.91

Yr17

8

6

7

MS

10

MS

30

32

16

7.56

Yr18

9

8

MR

10

MR

20

12

6

8.46

Yr26

3

7

MS

20

MS

30

40

20

7.20

Yr27

8

9

MS

30

S

20

44

22

7.02

Yr32

7

3

8

S

10

S

30

40

20

7.20

Avocet s

9

S

90

S

100

190

95

0.45

7

3.3 Detection of six

Yr

genes in wheat

genotypes using molecular markers

Six different markers were used for

detecting the presence of six

Yr

genes;

Yr5

,

Yr10

,

Yr15

,

Yr17

,

Yr18

and

Yr26,

in

a collection of 15 recent Egyptian wheat

cultivars and 5 breeding lines (Table 4)

and (Figure 1). The results indicated that

Yr5

was only detected in three wheat

genotypes including two cultivars Sakha-

95 and Misr-3, one breeding line Line-2.

Yr18

, and

Yr26

were detected in all

Egyptian wheat cultivars and breeding

lines. However,

Yr10

was detected in

only one cultivar Sakha-93.

Yr15

and

Yr17

were detected in some genotypes

but not all the tested genotypes.

Table 4: Detection of yellow rust (Yr) resistance genes using different molecular markers in Egyptian wheat

cultivars and breeding lines.

No.

Wheat accessions

Yr5

Yr10

Yr15

Yr17

Yr18

Yr26

Wheat genotypes

1

Sakha-8

-

+

2

Sakha-93

-

+

-

+

3

Sakha-94

-

+

4

Sakha-95

+

-

+

5

Giza-160

-

+

6

Giza-168

-

+

-

+

7

Giza-171

-

+

-

+

8

Sids-12

-

+

9

Sids-13

-

+

10

Sids-14

-

+

-

+

11

Gemmiza-11

-

+

-

+

12

Gemmiza-12

-

+

-

+

13

Misr-1

-

+

-

+

14

Misr-2

-

+

15

Misr-3

+

-

+

Breeding lines

16

Line-1

-

+

17

Line-2

+

-

+

18

Line-3

-

+

19

Line-4

-

+

20

Line-5

-

+

Figure 1: Electrophoretic amplified pattern of DNA extracted from 15 wheat cultivars and five breeding lines

using the specific primers of six Yr genes (a) Yr5, (b)Yr10, (c)Yr15, (d)Yr17, (e)Yr18, and (f)Yr26. In all

patterns L = DNA Ladder, 1 = positive samples, 2 = Sakha-8, 3 =Sakha-93, 4 = Sakha-94, 5 = Sakha-95, 6 =

Giza-160, 7 = Giza-168, 8 = Giza-171, 9 = Sids-12, 10 = Sids=13, 11 = Sids-14, 12 = Gemmiza-11, 13 =

Gemmiza-12, 14 = Misr-1, 15 = Misr-2, 16 = Misr-3, 17 = Line-1, 18 = Line-2, 19 = Line-3, 20 = Line-4, 21

= Line-5.

8

Yr15

was found in seven cultivars,

Sakha-93, Giza-168, Giza-171, Sids-14,

Gemmiza-11, Gemmiza-12, and Misr-1.

However,

Yr15

was not found in any of

the tested breeding lines.

Yr17

was

detected only in one of the most recent

released wheat cultivars, Misr-3. All the

tested breeding lines appeared to have

Yr17

except Line-1.

4. Discussion

This study was aimed to evaluate the

recent cultivated Egyptian wheat

cultivars and some promising wheat

breeding lines against the most

predominant

Pst

races, 224E132,

224E32, and 151E80.The tested wheat

genotypes showed variation in their

responses to

Pst

races. Wheat cultivars,

Sakha-93, Sakha-95, and Misr-3 were the

only wheat cultivars that showed

moderated to high resistance against

tested

Pst

races in seedling and adult

plant stages. The resistance of these

cultivars could be due to the presences

ofeffective stripe rust genes such

as;

Yr5

which found in Sakha-95, and

Misr-3,

Yr15

in Sakha-93, and

Yr17

in

Misr-3.Stripe rust resistance genes,

Yr5

,

Yr15

, and

Yr17

, are still effective to

stripe rust in Egypt (Shahin et al., 2015;

Abu-Aly et al., 2014). Only breeding

Line-2 was showed high resistance to

Pst

races in seedling and adult plant stages

that could be due to the presence of

Yr-5

,

17, 18 and 26 in its genomic background.

Also, Line-5 was showed moderated

resistance in seedling and high resistance

with low disease severity in adult stage,

against

Pst

races. In general, all the

tested Lines were showed high to

moderated resistance against

Pst

races in

the field which could be due to presence

of adult plant resistance genes.

Interestingly, high infection type was

observed on

Yr1

which considered one

of the most durable

Yr

gene against

151E80 (author observation). However,

it expressed high resistance in the open

field at adult plant stage. Another study

in Iran by Afshari (2018) was reported

virulence against

Yr1

,

Yr3

, and

Yrsp

.

Therefore, Breeders who relaying on

Yr1

as a major resistance gene at

Mediterranean region should be

cautious.Sakha-94, Misr-1, Misr-2, and

Line-1 showed moderated resistance to

the race mixture in adult plant stage. The

molecular study indicated that

Yr5

was

detected in Sakha-95, Misr-3, and Line-

2. While,

Yr18

and

Yr26

were detected in

all tested wheat cultivars and

lines.

Yr5

was originally identified in

hexaploid

Triticum aestivum

ssp.

spelta

cv.

album

(TSA) (Zhang et al., 2009).

Yr5

is an important

Yr

gene. It confers

resistance to almost all isolates of

Pst

in

the world, except for Australia (Wellings

& McIntosh, 1990) and probably India

(Nagarajan et al., 1986).

Yr18

/

Lr34

/

Pm38

/

Sr57

is a slow-rusting

and mildewing gene that confers partial,

durable resistance to multiple fungal

pathogens (Wu et al., 2015; Liang et al.,

2006; Spielmeyer et al., 2005; McIntosh,

1992; Singh, 1992; Singh & Rajaram,

1992). Therefore, this multi-pathogen

resistance locus is a valuable source of

resistance in wheat breeding

(Urbanovich et al., 2006). The

appearance of

Lr34/Yr18/Pm38,

in

Egyptian cultivars goes back to the

1970s, when Egyptian breeders began

9

incorporating the Japanese cultivar

Akakomughi (grandparent of spring

wheat variety Frontana, used widely as a

source of

Lr34

) in the parentage of most

released Egyptian cultivars (Basent et al.,

2011). The Brazilian cultivar Frontana is

the source of

Lr34

/

Yr18

in a significant

proportion of CIMMYT Since Sakha-8 is

considered the donor of

Lr34

in

subsequent crosses which led to many

recent varieties such as Sakha-94 and

Sids-13(Fahmi et al., 2015).

Yr26

was

alleged to be from the

T. turgidum

durum

line c80-1 (Ma et al., 2001) and was

located on wheat chromosome 1B

(Zhang et al., 2013; Wang et al., 2008;

Ma et al., 2001).

Yr10

was isolated from

Moro, it located in the short arm of

chromosome 1B (Singh et al., 2009;

Metzger & Silbaugh, 1970). In Egypt,

Yr10

consider an important

Yr

gene, in a

study by Abu-aly et al. (2014), it

exhibited high level of resistance against

198E56 and 128E28. Also,

Yr10

expressed seedling and adult plant

resistance to numerous races tested by

Shahin et al. (2015). Among all tested

cultivars,

Yr10

was detected in only

Sakha-93. This gene needs to be

introgressed intocommercial wheat

cultivars. The importance of

Yr10

is not

solely the response of the gene itself, but

also the expression of another genes that

combined to

Yr10

such as

Yr8

(Chen et

al., 2013).

Yr15

was detected in seven

Egyptian wheat cultivars but not in any

of the tested breeding lines. It derived

from

Triticum dicoccoides

, was located

in chromosome 1BS (McIntosh et al.,

1996).

It is an effective

Yr

gene against

most

Pst

races in Egypt (Shahin &

Ragab, 2015).

For pyramiding a durable

resistance to stripe rust in Egypt,

Yr5

and

Yr15

recommendedto be introgressed into

the most recent cultivars.

Yr17

is an

important

Yr

gene that confers durable

resistance. Also, it is linked to

Yr5

and

contribute to the resistance pathway of

Yr5

in wheat plants. In addition to

Yr17

,

Lr37

a leaf rust resistance gene, and

Sr38

a stem rust resistance gene is

located within a segment of

Triticum

ventricosum

chromosome 2NS

translocated to the short arm of wheat

chromosome 2A (Helguera et al., 2003).

Among all tested wheat cultivars only

the most recent cultivar Misr-3 has

Yr17

.

The low infection type of Misr-3 in

seedling stage and the high resistance of

adult plants could be due to the presence

of

Yr5

,

Yr17

,

Yr18

, and

Yr26

.

Yr17

was

found in four spring wheat breeding

lines, which could be used as a source of

Yr17

. Despite the detection of

Yr18

in all

the tested wheats, most of them

expressed high infection types against

tested

Pst

races. That could be due to the

absences of other

Yr

or functional genes

which involve in the final phenotype

response

Yr18

. There are a various

studies reported that resistances

controlled by NBS-LRR genes may not

be durable, and resistances controlled by

non-NBS-LRR genes are more likely to

be durable (Chen et al., 2013; Fu et al.,

2009; Krattinger et al., 2009). Where,

the final response phenotypes are not

only resulted as expression of one gene it

could result as a various defense which

regulates by numerous

Yr

and functional

genes. Therefore, breeders should be

cautious about what

Yr

genes could be

combined in particular cultivar to

express high level of resistance as a final

10

phenotypic response. In conclusion, six

Yr

genes were detected using different

markers.

Yr5

,

Yr18

,

Yr26

were detected

in all tested genotypes. The presences of

Yr5

in three wheat genotypes expressed

high resistance against all tested

Pst

races in all stages. However,

Yr18

was

not successfully achieved high resistance

in seedling and adult plant stages in some

cultivars due to the absences of

important

Yr

genes such as

Yr17

that has

a complementary resistance pathway to

Yr18

. There are four promising spring

wheat breeding lines could be used as a

source for

Yr5

,

Yr17

,

Yr18

, and

Yr26

.

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