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Genetic Resources and Crop Evolution 45: 9–17, 1998. 9c 1998 Kluwer Academic Publishers. Printed in the Netherlands.

Diversity for abiotic and biotic stress resistance in the wild annual Cicerspecies �

K.B. Singh1, B. Ocampo1 & L.D. Robertson2

1Germplasm Program & 2Genetic Resources Unit, International Center for Agricultural Research in the DryAreas (ICARDA), P.O. Box 5466, Aleppo, Syria

Received 24 February 1997; accepted 10 June 1997

Key words: chickpea, Cicer arietinum, disease, genetic diversity, genetic resources, multiple resistance, nematode,pest, wild species

Abstract

Data on 228 accessions of eight annual wild Cicer species and 20 cultivated chickpea check lines were evaluated fordiversity in response to six of the most serious biotic and abiotic stresses which reduce crop yield and productionstability of chickpea, i.e., ascochyta blight, fusarium wilt, leaf miner, bruchid, cyst nematode, and cold. Relativefrequencies of score reactions to the above six stresses were recorded from all the annual wild Cicer species andthe cultivated taxon. Patterns of distribution and amount of variation of the resistance reactions differed betweenstresses and species. C. bijugum, C. pinnatifidum and C. echinospermum showed accessions with at least one sourceof resistance (1 to 4 score reactions) to each stress. Overall, C. bijugum showed the highest frequencies of the highestcategories of resistance. Next in performance was C. pinnatifidum followed by C. judaicum, C. reticulatum and C.echinospermum. Furthermore, C. bijugum had the highest number of accessions with multiple resistance to the sixstresses; two accessions were resistant to five stresses and 16 to four. According to the Shannon-Weaver diversityindices (�H0), five species showed discrete mean diversity indices which varied from 0.649 in C. pinnatifidum to0.526 in C. judaicum, whereas C. chorassanicum, C. cuneatum and C. yamashitae showed the lowest �H0s, whichwere respectively 0.119, 0.174 and 0.216. Pair-wise correlations among the six biotic and abiotic stresses showedthe possibility of combining these resistances. Interestingly, multiple resistant accessions were predominantly ofTurkish origin.

Introduction

Despite large breeding efforts to improve chickpea(Cicer arietinum L.), lack of stable production contin-ues to be a major concern. This condition is aggravatedby the continuous shift of the crop to marginal landswhere it faces a multitude of stresses. Improvementin the current crop status requires resistance to bioticand abiotic stresses which are the fundamental caus-es for unstable production. The cultivated taxon lacksadequate sources of resistance to these stresses. Thesuccess of the large-scale adoption of winter sowing inthe Mediterranean region, which has shown the poten-

� Joint publication from ICARDA and ICRISAT (InternationalCrops Research Institute for the Semi-Arid Tropics), PatancheruP.O., A.P. 502324, India

tial to double chickpea yields (Hawtin & Singh, 1984),is crucially dependent on resistance to ascochyta blight(caused by Ascochyta rabiei [Pass.] Lab.). Worldwide,ascochyta blight and fusarium wilt (caused by Fusar-ium oxysporum Schlecht. emend. Synd. & Hans. f.sp.ciceri [Padwick] Synd. & Hans.) are the most seriousdiseases of chickpea. Leaf miner (Liriomyza cicerinaRond.) and cold are important stresses in the Mediter-ranean region. Wild relatives of chickpea have beenshown to be promising sources for genes for resistanceto the major biotic and abiotic stresses in chickpea(Singh et al., 1994). There are many examples of theexploitation of wild relatives for improvement of othercrops (Stalker, 1980; Prescott-Allen, 1988).

The genus Cicer comprises 43 species made upof 33 perennials, nine annuals including the culti-

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Table 1. Country of origin of the annual wild Cicer species accessions and C. arietinumlines

Cicer Country of origin

AFGa ETH JOR LBN PAL SYR TUR Total

no. of accessions

arietinum 4 2 3 1 2 7 1 20

bijugum – – – – – 4 33 37

chorassanicum 5 – – – – – – 5

cuneatum – 4 – – – – – 4

echinospermum – – – – – – 11 11

judaicum 2 2 8 17 9 22 7 67

pinnatifidum – 1 – 1 5 8 34 50b

reticulatum – – – – – – 51 51

yamashitae 3 – – – – – – 3

Total 14 9 11 19 16 41 137 248

a AFG= Afghanistan, ETH= Ethiopia, JOR= Jordan, LBN= Lebanon, PAL= Pales-tine, SYR= Syria, TUR = Turkey.b Country of origin is unknown for one accession.

vated chickpea and one unspecified (van der Mae-sen, 1987). The International Center for AgriculturalResearch in the Dry Areas (ICARDA), Aleppo, Syr-ia, maintains the world’s largest collection of annualwild Cicer species (270 accessions), and of the Kabulitype chickpea (9870 accessions). Extensive collectionsof wild Cicer are also maintained at the InternationalCrops Research Institute for the Semi-Arid Tropics(ICRISAT), India and at the USDA Regional PlantIntroduction Station at Pullman, Washington, USA.

The ICARDA world collection of annual wild Cicerspecies has been evaluated for ascochyta blight, fusari-um wilt, leaf miner, bruchid (Callosobruchus chinensisL.), cyst nematode (Heterodera ciceri Vovlas, Grecoet Di Vito), and cold (Singh et al., 1994). The morpho-physiological features of the annual wild Cicer speciesaccessions of ICARDA have been catalogued (Robert-son et al., 1995).

To date, no studies have been conducted on thecombined variability and diversity of responses to biot-ic and abiotic stresses in the annual wild Cicer species.This paper examines the patterns of distribution and thevariation in response to the above mentioned adverseconditions in a collection of 228 accessions of 8 annualwild Cicer species maintained by ICARDA. The objec-tive of this work was to estimate their relative diversity[Shannon-Weaver information index (�H0) based on fre-quency data] for response to six stresses. This proce-dure has been used earlier to measure genetic diversitybased on biotic and abiotic stress responses (Singh &

Jana, 1993) as well as on quantitative traits (Porceddu,1976; Holcomb et al., 1977; Peeters, 1988; Jana et al.,1990; Pecetti et al., 1992).

Material and methods

The materials of this study consisted of 228 accessionsof the eight annual wild Cicer species from the ICAR-DA genebank, C. bijugum K.H. Rech., C. chorassan-icum (Bunge) M. Popov, C. cuneatum A. Rich., C.echinospermum Davis, C. judaicum Boiss., C. pin-natifidum Jaub. & Spach, C. reticulatum Ladiz. andC. yamashitae Kitamura, and 20 cultivated chickpealines. The cultivated accessions were chosen in orderto cover the geographical distribution of the annualwild Cicer species. Table 1 shows the number of acces-sions per species used in this study and their countryof origin. The accessions were screened for resistanceto ascochyta blight (Ascochyta rabiei), fusarium wilt(Fusarium oxysporum f.sp. ciceri), leaf miner (Liri-omyza cicerina), bruchid (Callosobruchus chinensis),cyst nematode (Heterodera ciceri), and cold resistance.

All evaluations, except for fusarium wilt, were car-ried out at Tel Hadya (36� 350E, 36� 510N and 284 ma.s.l.), Syria, the main research station of ICARDA.This site has a semi-arid continental Mediterraneanclimate with cold and relatively wet winters, dry andhot summers, and an average of 16 seasons precipi-

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tation of about 330 mm. Evaluation for fusarium wiltwas done at Monterotondo, Rome, Italy.

Screening started in the 1987/88 season. The fol-lowing procedures were used. Ascochyta blight resis-tance (AB); was scored on a scale of 1 to 9, where1 = plants free from any damage and 9 = all plantskilled, following the technique and the scale describedby Singh et al. (1981). Each accession was sown in asingle 2 m row plot with inter- and intra-row spacing of45 and 10 cm, respectively (20 plants). Fusarium wiltresistance (FW); was scored on a scale of 1 to 9, where1 = plants free from any damage and 9 = all plantskilled, following the technique described by Infantinoet al. (1996) and a scale developed by Singh (unpub-lished): 10 to 15 plants per accession were assayed.Leaf miner resistance (LM); was scored on a scale of1 to 9, where 1 = plants free from any damage and9 = leaves severely mined, many dried and droppedfrom the plant, following the technique and the scaledescribed by Singh & Weigand (1994). Each accessionwas sown in a single 2 m row plot with inter- and intra-row spacing of 45 and 10 cm, respectively (20 plants).Bruchid resistance (CC); was scored on a scale of 1 to9, where 1 = seeds free from any damage and 9 = allseeds infested by beetles, following the technique andthe scale described by Weigand & Tahhan (1990); 50seed were assayed per accession. Cyst nematode resis-tance (CN); was scored on a scale of 1 to 9, where 1=plants free from any damage and 9 = highly suscepti-ble, following the technique described by Di Vito et al.(1988) and a scale developed by Singh (unpublished).Two pots of five seed each were grown per accessionto provide two replications. Cold resistance (CO); wasscored on a scale of 1 to 9, where 1 = plants freefrom any damage and 9 = all plants killed, followingthe technique and the scale described by Singh et al.(1989). Each accession was sown in two replicates ina randomized complete block design with each plot asingle 2 m row using inter- and intra-row spacing of 45and 10 cm, respectively (20 plants per replicate).

The screening for ascochyta blight resistance wascarried out in an ascochyta blight nursery, wheredisease was created artificially through inoculationwith diseased plant debris and with spore-suspensionsprays. The screening for fusarium wilt resistance wascarried out in a greenhouse at the Istituto Sperimentaleper la Patologia Vegetale, Rome. The screening forleaf miner resistance was carried out by growing theaccessions during spring under natural insect infesta-tion. The screening for bruchid infestation was carriedout by artificially releasing the bruchid in the Ento-

mology Laboratory of ICARDA to seed samples. Cystnematode screening was conducted in the greenhouseby growing accessions in plastic pots containing cystnematode infested soils. The screening for cold toler-ance was conducted at the Tel Hadya farm by advanc-ing the sowing date to early October. Evaluations weredone in systematic screening nurseries with repeatedchecks, except for fusarium wilt. Frequency distribu-tion, mean, standard deviation, coefficient of variationand skewness were calculated for each stress for eachspecies, and for the overall annual wild Cicer speciescollection using SPSSr for WindowsTM, Release 6.0(SPSS, 1993).

The measurements of genetic diversity for biot-ic and abiotic stress reactions were assessed by theShannon-Weaver information index (H0) (Shannon &Weaver, 1949) given as:

H0= �

sX

i=1

pi ln pi

where s is the number of reaction scores showed byaccessions of a species to a given stress and pi is theproportion of the total number of accessions with theith score reaction. To confine the information index inthe range 0 to 1, each value of H0 was normalized bydividing it by its maximum value, i.e., log s (Tolbertet al., 1979). The mean information index (�H0) of aspecies was estimated as the arithmetic average of the�H0s values over all stresses.

Simple correlation coefficients were computedfrom stress scores for all pairwise combinations ofstresses for the entire wild Cicer germplasm collec-tion using SPSSr for WindowsTM, Release 6.0 (SPSS,1993).

Results

Distributions of relative frequencies (%) of reactionsof the annual Cicer species to the six stresses: ascochy-ta blight, fusarium wilt, leaf miner, bruchid, cystnematode and cold, are given in Table 2. Overall,the six stresses differed in their patterns of distribu-tion and amount of variation. For instance, distributionfor ascochyta blight and bruchid were skewed towardsresistance; that for fusarium wilt was skewed towardssusceptibility; whereas distributions for leaf miner andcold were nearly normal. Species-wise distribution ofscore reactions were similar for ascochyta blight, fusar-

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Table 2. Distribution of relative frequencies (%) of reactions (1–9 scale) of annual wild Cicer species to six biotic andabiotic stresses

Cicer species

Scale aria ret ech jud pin bij cho yam cun

Ascochyta blight

1 (free from damage) – – – – – – – – –

2 (highly resistant) – – 9.1 3.5 8.9 5.4 – – –

3 (resistant) – – – – 2.2 13.5 – – –

4 (moderately resistant) – – – 3.5 – 5.4 – – –

5 (tolerant) 15.8 – 9.1 8.8 26.7 18.9 – – –

6 (moderately susceptible) 5.3 4.1 27.3 10.5 20.0 35.1 – – –

7 (susceptible) 15.8 10.2 18.2 12.3 13.3 13.5 – – 25.0

8 (highly susceptible) 15.8 30.6 9.1 8.8 15.6 2.7 – – –

9 (all plants killed) 47.4 55.1 27.3 52.6 13.3 5.4 100.0 100.0 75.0

Skewness �0.88 �1.24 �0.98 �1.27 �0.47 �0.17 – – �2.00

No. of accessions 19 49 11 57 45 37 1 3 4

Fusarium wilt

1 (free from damage) 10.0 23.4 44.4 43.9 18.8 81.0 – – –

2 (highly resistant) – 12.8 11.1 5.3 6.3 6.1 – – 66.7

3 (resistant) 10.0 10.6 – 1.8 – 3.0 – – 33.3

4 (moderately resistant) 10.0 31.9 22.2 26.3 15.6 3.0 – – –

5 (tolerant) – 6.4 11.1 12.3 6.3 – – – –


7 (susceptible) 10.0 – – – 15.6 – – – –

8 (highly susceptible) 20.0 2.1 – – 12.2 – – – –

9 (all plants killed) 30.0 – – 1.8 15.6 – – – –

Skewness �0.89 0.29 0.52 0.58 �0.31 2.92 – – 1.73

No. of accessions 10 47 9 57 32 33 0 0 3

Leaf miner

1 (free from damage) – – – – – – – – –

2 (highly resistant) – – 9.1 29.0 8.5 2.8 100.0 – 100.0

3 (resistant) – 8.3 27.3 35.5 8.5 5.6 – – –

4 (moderately resistant) 18.8 14.6 27.3 11.3 19.1 16.7 – – –

5 (tolerant) 6.3 35.4 – 17.7 38.3 33.3 – 33.3 –


7 (susceptible) 18.8 20.8 9.1 3.2 8.5 30.6 – 33.3 –

8 (highly susceptible) 6.3 – – – – – – – –

9 (very highly susceptible) 43.8 – – – – 5.6 – 33.3 –

Skewness �0.60 �0.18 0.30 0.88 �0.39 0.14 – 0.00 –

No. of accessions 16 48 11 62 47 36 1 3 4

ium wilt and leaf miner, and different for bruchid, cystnematode and cold.

C. bijugum, C. pinnatifidum and C. echinospermumshowed accessions with at least one source of resis-tance (1 to 4 score reactions) to each stress (Table 2).The highest levels of resistance within this collectionwere found for fusarium wilt, bruchid and cyst nema-tode. The least number of resistance sources werefound for ascochyta blight. The highest relative as well

as absolute number of accessions resistant to ascochy-ta blight, fusarium wilt, cyst nematode and cold (1to 2 score reactions) were found in C. bijugum. C.chorassanicum and C. cuneatum showed the highestpercentage (%) of accessions highly resistant to leafminer but they numbered merely one and four acces-sions respectively. On the other hand, 18 (29%) ofthe accessions of C. judaicum were highly resistant toleaf miner. Similarly, for bruchids, all accessions of

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Table 2. Continued

Cicer species

Scale aria ret ech jud pin bij cho yam cun

Bruchid

1 (free from damage) – 5.0 100.0 12.8 – 42.9 – – –

2 (highly resistant) – 5.0 – 8.5 6.9 23.8 – – 33.3

3 (resistant) – 5.0 – – – 14.3 – – –

4 (moderately resistant) – 5.0 – – 3.4 4.8 – – 33.3

5 (tolerant) – 5.0 – 2.1 – – – – –

6 (moderately susceptible) – 15.0 – 4.3 – 14.3 – – –

7 (susceptible) 10.0 45.0 – 12.8 – – 33.3 – 33.3

8 (highly susceptible) 30.0 15.0 – 51.1 72.4 – 66.6 100.0 –

9 (very highly susceptible) 60.0 – – 8.5 17.2 – – – –

Skewness �1.18 �1.40 – �1.24 �2.64 1.28 �1.73 – �0.94

No. of accessions 10 20 3 47 29 21 3 1 3

Cyst nematode

1 (free from damage) – – – – 2.3 – – – –

2 (highly resistant) – 2.0 – – 6.8 37.8 – – –

3 (resistant) – 11.8 – – – 51.4 – 33.3 –

4 (moderately resistant) – 13.7 9.1 33.9 9.1 10.8 40.0 66.7 –

5 (tolerant) – 72.6 90.9 66.1 81.8 – 60.0 – 100.0

6 (moderately susceptible) 6.7 – – – – – – – –

7 (susceptible) – – – – – – – – –

8 (highly susceptible) 20.0 – – – – – – – –

9 (all plants killed) 73.3 – – – – – – – –

Skewness �1.18 �1.68 �3.32 �0.70 �2.66 0.33 �0.61 �1.73 –

No. of accessions 15 51 11 62 44 37 5 3 4

Cold

1 (free from damage) – – – – – – – – –

2 (highly resistant) – 4.7 – – – 63.6 – – –

3 (resistant) – 60.5 – – – 30.3 – – –

4 (moderately resistant) – 30.2 100.0 – 20.5 6.1 – – –

5 (tolerant) 6.7 4.7 – – 28.2 – – – –

6 (moderately susceptible) 6.7 – – – 18.0 – – – –

7 (susceptible) 26.7 – – 1.9 20.5 – – – –

8 (highly susceptible) 13.3 – – 5.8 10.3 – – 33.3 –

9 (all plants killed) 46.7 – – 92.3 2.6 – 100.0 66.7 100.0

Skewness �0.84 0.60 – �4.05 0.39 1.17 – �1.73 –

No. of accessions 15 43 8 52 39 33 3 3 4

a ari = arietinum (check); ret = reticulatum; ech = echinospermum; jud = judaicum; pin = pinnatifidum; bij =bijugum; cho = chorassanicum; yam = yamashitae; cun = cuneatum.

C. echinospermum were found free from damage butthere were only three accessions. However, C. bijugumshowed nine accessions (42.9%) free from damage andfive (23.8%) highly resistant.

C. bijugum was the only annual wild Cicer specieswhich showed accessions free from damage and highlyresistant (1–2 score reactions) to all the six stresses.Next in performance was C. pinnatifidum with a 1–2

score to all six stresses, except cold. These specieswere followed by C. judaicum, C. reticulatum and C.echinospermum. C. yamashitae, C. chorassanicum andC. cuneatum, in that order, showed the least number ofaccessions with desirable resistance ratings (scores of1–4).

Overall, C. bijugum showed the best mean reactionperformances followed by C. judaicum, C. reticula-

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Table 3. Means, ranges, standard deviations and coefficients of variation of biotic and abiotic stressesdescriptors (1–9 scale) in the ICARDA annual wild Cicer species collection

State Cicer species

ari ret ech jud pin bij cho yam cun Totalb

Ascochyta blight

Mean 7.7 8.4 6.7 7.6 6.2 5.4 9.0 9.0 8.5 7.1

Range 5–9 6–9 2–9 2–9 2–9 2–9 9–9 9–9 7–9 2–9

S.D. 1.52 0.84 2.10 1.89 1.98 1.72 0.00 0.00 1.00 1.99

C.V. 19.67 9.97 31.25 24.92 32.11 31.76 0.00 0.00 11.76 23.71

Fusarium wilt

Mean 6.4 3.3 2.8 2.9 5.3 1.5 – – 2.3 3.2

Range 1–9 1–8 1–6 1–9 1–9 1–6 – – 2–3 1–9

S.D. 2.94 1.81 1.98 2.01 2.87 1.33 – – 0.58 2.32

C.V. 44.85 54.16 71.49 68.31 53.96 87.52 – – 24.73 63.66

Leaf miner

Mean 7.2 5.3 4.4 3.4 4.7 5.5 2.0 7.0 2.0 4.6

Range 4–9 3–7 2–7 2–7 2–7 2–9 2–2 5–9 2–2 2–9

S.D. 2.00 1.20 1.63 1.34 1.33 1.61 0.00 2.00 0.00 1.66

C.V. 26.41 22.70 37.33 39.26 28.16 29.16 0.00 28.57 0.00 30.12

Bruchid

Mean 8.6 6.0 1.0 6.4 7.6 2.4 7.7 8.0 4.3 5.8

Range 7–9 1–8 1–1 1–9 2–9 1–6 7–8 8–8 1–7 1–9

S.D. 0.71 2.00 0.00 2.74 1.78 1.75 0.58 0.00 3.06 2.87

C.V. 8.32 33.33 0.00 42.78 23.37 73.33 7.52 0.00 70.50 38.28

Cyst nematode

Mean 8.6 4.6 4.9 4.7 4.6 2.7 4.6 3.7 5.0 4.3

Range 6–9 2–5 4–5 4–5 1–5 2–4 4–5 3–4 5–5 1–5

S.D. 0.83 0.78 0.30 0.48 0.97 0.65 0.55 0.57 0.00 1.00

C.V. 5.24 17.09 6.15 10.23 21.02 23.88 11.91 15.73 0.00 16.30

Cold

Mean 7.9 3.3 4.0 8.9 5.8 2.4 9.0 8.7 9.0 5.6

Range 5–9 2–5 4–4 7–9 4–9 2–4 9–9 8–9 9–9 2–9

S.D. 1.30 0.65 0.00 0.36 1.40 0.61 0.00 0.58 0.00 2.72

C.V. 16.55 19.41 0.00 4.02 24.14 25.33 0.00 6.66 0.00 14.19

a ari= arietinum (check); ret= reticulatum; ech= echinospermum; jud= judaicum; pin= pinnatifidum;bij = bijugum; cho = chorassanicum; yam = yamashitae; and cun = cuneatum.b calculated considering only the wild Cicer species.

tum, and C. pinnatifidum (Table 3). Also, C. bijugumhad the highest number of accessions with multipleresistance (scores 1–3) to the six stresses, followedby C. reticulatum and C. judaicum (Table 4). Two C.bijugum accessions were resistant to five stresses and16 to four; among these, three accessions were moder-ately resistant (score 4) to an additional stress.

A negative association (r = �0.34��, P < 0.01)was found between cold and leaf miner resistances,but significant positive associations were observedfor: ascochyta blight and cold, ascochyta blight and

cyst nematode, ascochyta blight and bruchid, fusariumwilt and cyst nematode, fusarium wilt and bruchid, leafminer and cyst nematode, bruchid and cyst nematode,bruchid and cold, and cyst nematode and cold (Table 5).This is a good indication that combining resistancesto these stresses, except cold and leaf miner, can bepossible using the wild Cicer species.

Comparative assessment of mean genetic diversi-ty (�H0) among the species showed considerable vari-ation (Table 6). The annual wild Cicer species withthe highest mean diversity over all stresses were: C.

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Table 4. Accessions of annual wild Cicer species with overall best reactions to six biotic andabiotic stresses

Species Accession Origin Biotic and abiotic stresses

(ILWC) ABa FW LM CC CN CO

Scoresb

C. bijugum 73 TURc 3 1 7 1 2 2

C. bijugum 65 TUR 5 1 5 1 2 2

C. bijugum 74 TUR 5 1 5 1 2 2

C. bijugum 79 TUR 6 1 4 2 2 2

C. bijugum 62 TUR 4 1 7 2 2 2

C. bijugum 66 TUR 5 2 3 NTd 3 2

C. bijugum 72 TUR 5 1 3 3 3 NT

C. bijugum 76 TUR 5 1 5 2 3 2

C. bijugum 69 TUR 3 4 5 2 3 2

C. bijugum 70 TUR 3 1 9 1 2 3

C. cuneatum 187 ETH 9 2 2 1 5 9

C. cuneatum 232 ETH 7 3 2 NT 5 9

C. echinospermum 39 TUR 7 1 4 1 5 4

C. echinospermum 245 TUR 2 5 3 NT 5 NT

C. judaicum 255 JOR 2 2 3 NT 4 NT

C. judaicum 256 JOR 2 NT 3 NT 4 NT

C. judaicum 46 SYR 9 1 2 1 5 9

C. judaicum 189 LBN 9 1 3 1 4 9

C. pinnatifidum 250 TUR 2 2 3 NT 2 NT

C. pinnatifidum 251 TUR 2 1 4 NT 5 NT

C. pinnatifidum 171 TUR 5 1 5 2 5 5

C. reticulatum 113 TUR 2 6 5 3 NT NT

C. reticulatum 112 TUR 9 1 6 2 4 3






a AB = ascochyta blight, FW= fusarium wilt, LM = leaf miner, CC = bruchid, CN = cystnematode, CO = cold.b 1–9 scale.c TUR = Turkey, ETH = Ethiopia, JOR = Jordan, SYR = Syria, LBN = Lebanon.d Not tested.

Table 5. Linear correlation coefficients for six biotic and abioticstresses in annual wild Cicer species

Character FWa LM CC CN CO

AB .016 �.101 .190� .319�� .280��

FW �.090 .357�� .406�� .120

LM �.057 .147� �.335��

CC .456�� .385��

CN .419��

a AB= ascochyta blight, FW= fusarium wilt, LM= leaf miner,CC = bruchid, CN = cyst nematode, CO= cold.�,�� Significant association at the 5% and 1% level, respectively.

pinnatifidum (0.649), C. reticulatum (0.588), and C.bijugum (0.544), which did not differ statistically fromeach other. The least diverse species were C. choras-sanicum (0.199), C. cuneatum (0.174), C. yamashitae(0.216), and C. echinospermum (0.372), which alsodid not differ statistically from each other but differedsignificantly (P < 0.05) from the species cited above.

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Table 6. Estimates of the diversity indices (Shannon-Weaver) (�H0) for annual wild Cicer speciesand mean diversity (�H0) and its standard error over six biotic and abiotic characters

Species Biotic and abiotic stresses

ABa FW LM CC CN CO �H0 � SE

H0

C. arietinum 0.630 0.780 0.701 0.409 0.264 0.609 0.566� 0.079

C. reticulatum 0.480 0.785 0.686 0.763 0.380 0.432 0.588� 0.073

C. echinospermum 0.762 0.649 0.682 0.000 0.139 0.000 0.372� 0.148

C. judaicum 0.680 0.676 0.682 0.685 0.290 0.143 0.526� 0.100

C. pinnatifidum 0.819 0.915 0.735 0.380 0.297 0.748 0.649� 0.102

C. bijugum 0.816 0.329 0.734 0.640 0.432 0.373 0.554� 0.083

C. chorassanicum 0.000 NTb 0.000 0.290 0.306 0.000 0.119� 0.073

C. yamashitae 0.000 NT 0.500 0.000 0.290 0.290 0.216� 0.096

C. cuneatum 0.256 0.290 0.000 0.500 0.000 0.000 0.174� 0.085

a AB = ascochyta blight, FW = fusarium wilt, LM = leaf miner, CC = bruchid, CN = cystnematode, CO = cold.b Not tested.

Discussion

The (�H0) values showed that the difference in responseto the six biotic and abiotic stresses among the annu-al wild Cicer species is large, suggesting moderateto high polymorphism. The low mean diversity (�H0)values (proportion of monomorphic accessions) of C.yamashitae, C. cuneatum, C. chorassanicum and C.echinospermum collections were quite possibly associ-ated with their small collection size. They were respec-tively composed of three, four, five and 11 acces-sions. Collection size is crucial for proper exploita-tion in breeding activities. Large collections are morelikely to hold rare genes. This would explain the lowmean diversity (�H0) of reactions to the biotic and abi-otic stresses of C. chorassanicum, C. cuneatum andC. yamashitae. Particularly critical is the condition ofC. echinospermum. C. echinospermum and C. reticu-latum are the only wild species able to cross and pro-duce fertile offspring with chickpea (ICARDA, 1990;Singh & Ocampo, 1993; Ocampo, 1995). C. echi-nospermum accounts for only 11 accessions (4% ofthe annual wild Cicer species collection maintainedat ICARDA), which mostly share a common genotype(Robertson et al., 1995). The diversity data of this studyunderscores the need for further exploration efforts.This is particularly true for C. echinospermum and C.reticulatum, which are of potential use in the genet-ic improvement of the cultivated taxon. Resistancesof promising C. reticulatum accessions for ascochy-ta blight were ephemeral when introgressed into Desichickpea genotypes (van der Maesen & Pundir, 1984).

Nevertheless, attempts are undergoing at ICARDA totransfer cyst nematode resistance and cold tolerancegenes from C. reticulatum and C. echinospermum tochickpea (ICARDA, 1997). These two wild relativesare also being exploited at ICARDA for improvingagronomic traits in chickpea (ICARDA, 1994, 1995;Singh & Ocampo, 1997). Results of two years trialshave shown that a few test lines surpassed the culti-vated parent, ILC 482, for seed yield, seed mass andplant height. These lines were also free of any knownundesirable traits from the wild species.

Genotypes with multiple sources of stress resis-tance are of great importance in breeding. Chickpealacks accessions with genes for multiple-stress resis-tance to the stresses studied, therefore much effort isbeing made to merge individual sources of resistanceinto one adaptive genotype through convergent crosses.For this, annual wild Cicer species might be a panacea,as several accessions are endowed with genes resistantto diverse biotic and abiotic stresses. Unfortunately,the most promising accessions belong to species of thetertiary chickpea gene-pool, namely, C. bijugum, C.judaicum and C. pinnatifidum. Most wild annual Cicerspecies occur in Turkey, and most (multiple) resistancehas its origin in Turkish accessions.

Multiway frequency analysis of data collectedfrom large-scale chickpea germplasm evaluation (5672accessions) showed high associations between seedyield and stress responses to ascochyta blight, leafminer and cold (Singh & Jana, 1993). Therefore, seedyield improvement and production stability of chick-pea through enhancement of their stress response to

17

biotic and abiotic stresses are strategies that should befirmly pursued.

Acknowledgements

We wish to thank the assistance of Drs. N. Greco andM. Di Vito for screening for cyst nematode resistance;Drs. A Porta-Puglia and A Infantino for fusarium wiltscreening at the Istituto Sperimentale per la PatologiaVegetale in Rome, Italy; Dr. R.S. Malhotra for screen-ing to cold and Drs. S. Weigand and O. Tahhan forbruchid and leaf miner screening.

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