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. 2026 Jan 22;16:5983. doi: 10.1038/s41598-026-36160-7

Morphological evaluation and ınheritance of powdery mildew resistance in garden pea (Pisum sativum L.)

Hanuman Ram 1,✉, Shri Dhar 2, H Choudhary 2, L Prasad 3, B S Tomar 2, G S Jat 2,✉
PMCID: PMC12901017  PMID: 41571761

Abstract

In this study, eleven diverse germplasm lines and their 55 F1s were morphologically evaluated for 10 economically important quantitative traits in garden pea. These germplasms were also screened to identify resistant sources against the powdery mildew disease. Among 11 parental lines evaluated, GP-17 was superior for earliness while pod length and width were highest in VP-233. Maximum number of seeds per plant, pod weight and shelling percentage were recorded in genotype GP-473. The maximum number of pods per plant were observed in IP-3 whereas, the highest yield per plant was recorded in VRP-7. Among F1 crosses, Arka Ajit × Pusa Pragati was the earliest and GP-17 × VRP-7 was superior for number of pods per plant. The maximum pod length, pod width, pod weight and number of seeds per pod were observed in GP-17 × VP-233, VRP-7 × IP-3, Arka Ajit × GP-6 and Arkel × VRP-7, respectively. Maximum shelling percentage was recorded in VP-233 × AP-3 while VP-233 × IP-3 gave maximum yield. Based on screening data, five genotypes (GP-6, GP-473, Arka Ajit, Pusa Pragati and VP-233) exhibited percent disease index (PDI) in the range of 2.54 to 7.78 and were considered resistant to powdery mildew and can be utilized as resistant sources in breeding programmes. The mendelian segregation ratio in six generations (P1, P2, F1, F2, BC1 and BC2) of five cross combinations using chi-square analysis showed that a monogenic recessive gene controls powdery mildew resistance in garden pea and it can be introgressed by backcross breeding in the popular and high yielding susceptible cultivars which will create a new pathway of resistance breeding in this crop.

Subject terms: Genetics, Plant sciences

Introduction

Garden pea (Pisum sativum L.) is an economically important annual herbaceous legume vegetable belongs to the family fabaceae grown for fresh green pods as vegetable throughout the world. It is used as pulse or in soup, canned, processed or dehydrated and green shelled forms. The green pods are rich source of proteins (5.42 g/100 g), carbohydrates (14.4 g/100 g), total dietary fibre (5.7 g/100 g), fats (0.4 g/100 g), vitamin A (765 I.U.), thiamine (0.266 mg/100 g), riboflavin (0.132 mg/100 g), ascorbic acid (40 mg/100 g), calcium (25 mg/100 g), magnesium (33 mg/100 g), phosphorus (108 mg/100 g), copper (0.176 mg/100 g), potassium (244 mg/100 g) and iron (1.47 mg/100 g)1. Generally, garden pea is grown in winter season (mid-October to November) in the North Indian plains but as an important summer (off-season) crop in the high hills2,3. In breeding of high-yielding and disease resistant varieties of vegetable crops, the breeder regularly faces problem in selection of parental lines and F1 crosses which is essential for genetic improvement of garden pea and can be easily and efficiently done through morphological evaluation, characterization and screening for disease resistance4. In garden pea, several cultivars have been developed for commercial cultivation through selection or hybridization for increasing the yield but still yield potential is not up to the desired level, hence, there is always a strong need to characterize superior genotypes or lines for utilizing them as a variety or as a parental line in the breeding programme. Therefore, emphasis should be given for characterization and development of high yielding disease resistant vegetable pea varieties with quality traits. For the identification and development of superior genotypes, the morphological characterization for economic traits is the most important step in breeding programme for genetic improvement of yield traits.

Garden pea is vulnerable to several diseases, viz., Powdery mildew, Fusarium wilt, Ascochyta blight, Bacterial blight, white rot and rust. Among these, powdery mildew caused by an obligate biotrophic fungus, Erysiphe pisi Syd., is one of the most troublesome diseases over the world4 and causes severe decline in yield as well as deteriorate the quality of pods5,6. The disease become prominent during warm, dry days and cool nights4,7, results in the decreases in total yield, number of pods per plant, number of seeds per pod, plant height and number of nodes8. The losses due to high incidence of powdery mildew leads to 24–27% reduction in pod weight, 21–31% in pod number and 25–86% in total yield8–11. The magnitude of yield losses can increase (up to 80%) many times under favourable conditions for disease development because plants fail to reach at reproductive phase12. The principal method of controlling powdery mildew in garden pea is the use of protective fungicides (particularly sulphur containing) at regular intervals but the air borne nature and fast conidial multiplication render fungicidal control ineffective and also enhance cost of cultivation.

Therefore, the development of cultivars with inherent resistance to powdery mildew is one of the most effective and economical means of controlling the disease. Earlier attempts were also made to locate the source of resistance to powdery mildew and different sources with complete or incomplete resistance to E. pisi reported worldwide10,13–16. Three genes er1, er213 and Er317 have been identified in distinct genomic locations on the pea genome, specifically on linkage groups VI, III, and IV, respectively18–21 carrying resistance to powdery mildew. The presence of gene er1 is reported in many field pea accessions22 but even with the absence of specific virulent races of E. pisi, breakdown in resistance against er1 gene had also been reported23,24 while, the expression of er2 gene had found to be dependent on temperature, growing conditions (field vs. green house) and leaf age22,24. The gene Er3 is present in wild Pisum species, Pisum fulvum and reported to be monogenic dominant in resistance, expressed as post penetration hypersensitive response, however the interspecific crossing led to sterile F1 hybrids17,25. The phenomenon of resistance in cultivated pea is specific to agro-climatological zones or environmental conditions, therefore, new sources of host resistance to E. pisi are needed to control the powdery mildew. Understanding inheritance patterns of disease resistance is crucial for effective disease management and developing resistant genotypes, which can help in reducing disease outbreaks and their impact on production as well as quality harvest. Therefore, keeping in view above all facts and realizing the importance of powdery mildew disease in garden pea cultivation, present investigation was undertaken to study the inheritance pattern of powdery mildew resistance along with morphological evaluation in garden pea with the hypothesis of powdery mildew resistance in selected pea genotypes is expected to be governed by a recessive gene.

Material and methods

Experimental location and materials used

The present investigation was carried out at the experimental farm of Division of Vegetable Science, ICAR-Indian Agricultural Research Institute, New Delhi (Latitude of 28.35° N and longitude 77.12° E and at an elevation of 228.6 m above mean sea level) during winter season of 2014–15, 2015–16 and 2016–17. Delhi’s climate is semi-arid sub-temperate experiences very hot summers (above 40 °C) and cool winters (below 5 °C). Daily maximum and minimum temperatures and evapo-transpiration rate are in increasing trend from February to June and in decreasing trend from July onwards. The total annual rainfall was about 625.6 mm, of which 74% were received during a short span of July to September. Fifty-five F1 hybrids developed through crossing along with the eleven parental lines (Table 1) were evaluated in randomized block design (RBD) with three replications. The plot size was kept as two rows of 4 m length for each genotype keeping plant spacing 45 × 10 cm apart to ensure at least 50 plants per plot. The recommended fertilizer (40:60:40 as NPK) doses were applied at the time of final land preparation and standard cultural practices were followed to raise a healthy crop. The observations were recorded based on 10 sample plants per plot.

Table 1.

List of the garden pea genotypes along with their source.

Designated Code Genotypes Maturity group Collected from
P1 Arkel Early ICAR-IARI, New Delhi
P2 GP-17 Early ICAR-IARI, New Delhi
P3 Arka Ajit Medium ICAR-IIHR, Bangalore
P4 Pusa Pragati Early ICAR-IARI, New Delhi
P5 GP-473 Medium ICAR-IARI, New Delhi
P6 VP-233 Medium ICAR-VPKAS, Almora
P7 VRP-7 Medium ICAR-IIVR, Varanasi
P8 VRP-6 Early ICAR-IIVR, Varanasi
P9 AP-3 Early CSAUAT, Kanpur
P10 IP-3 Medium GBPUAT, Pantnagar
P11 GP-6 Medium ICAR-IARI, New Delhi
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Observations recorded

The observations were recorded for ten quantitative traits, viz. plant height (cm), node bearing first flower, days to 50% flowering, number of pods per plant, pod length (cm), pod width, number of seeds per pod, average pod weight (g), shelling percentage and pod yield per plant (g). The shelling percentage was estimated as follows:

graphic file with name d33e556.gif

Screening and percent disease index (PDI)

The mass multiplication by artificial inoculation of pathogen, 30-days-old pea plants were challenged with ~ 10 mg of E. pisi (New Delhi Isolate) conidia was multiplied as dusted with camel hair brush for uniform development of inoculum and maintained under controlled conditions at Vegetable Research Farm, ICAR-Indian Agricultural Research Institute, New Delhi, India, till its further used for field screening (Fig. 1). Such Powdery mildew infected leaves were collected and spores with mycelia were harvested to prepare conidial suspension using haemocytometer. Artificial inoculation for powdery mildew inoculum was done by spraying conidial solution (2 × 105 conidia/ml of water) in the field conditions during crop seasons at ICAR-IARI, New Delhi.

Fig. 1.

Fig. 1

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Mass multiplication of pure culture of powdery mildew on pea plants.

The observations on disease severity on 10 plants in each genotype for terminal disease reaction were recorded 12–14 days post inoculation using 0–9 scale based on percentage of leaf area affected: 0 = no infection,1 =  < 1%, 2 = 1%–5%, 3 = 5%–10%, 4 = 10%–20%, 5 = 20%–40%, 6 = 40%–60%, 7 = 60%–80%, 8 = 80%–90%, 9 =  > 90%, according to Warkentin et al.(1995)26. Plant disease severity was calculated in term of percent loss and rating were given 0 (Immune), 1–10 (Resistant), 11–25 (Moderately resistant) 26–50 (Moderately susceptible), 51–75 (Susceptible) and > 75 (Highly susceptible) to the different genotypes of garden pea. Percent Disease Index (PDI) was calculated as:

graphic file with name d33e584.gif

Based on the screening data of genotypes for powdery mildew resistance, these genotypes were confirmed for their resistance in the field. The resistant and susceptible parents were selected for developing F1, F2, BC1 and BC2 populations. The F1 s seed of different crosses and selfed seeds of parental lines were individually collected and stored. The F1 seeds were sown in next year. F2 seeds were produced by selfing in the F1 plants and simultaneously backcross generations were also developed by crossing F1 plants with both the parents (B1 and B2). Thus, the seeds of a complete set of six generations (P1, P2, F1, F2, B1 and B2) for each cross were developed for field evaluation and genetic inheritance study of PM resistance.

Data analysis

The data obtained in the experiment were subjected to the statistical analysis of ANOVA for partitioning the total variance due to replications and treatments for all the characters. Mean comparisons among treatments/genotypes were performed using Duncan’s multiple range test (DMRT) when the P value less than 0.05 was considered significant. Data were subjected to analysis according to Method 2 Model I of Griffing (1956)27 for GCA and SCA analysis. The goodness of fit to Mendelian segregation of resistant and susceptible plants in the segregating population was tested by Chi- square test (χ2) as given by Pearson28.

where Inline graphic.

oi = Observed frequency of plants.

ei = Expected frequency of plants.

n = Number of phenotypic classes.

Null hypothesis (H0) is no difference between observed and expected frequency and H0 was accepted when the calculated χ2 value was less than table value for (n − 1) degree of freedom (d.f.) at P = 0.05. Whenever the calculated value of χ2 was greater than the table value, the null hypothesis was rejected. The statistical software used for the analysis is Statistical Package for Agricultural Research (SPAR 2.0) developed by ICAR-Indian Agricultural Statistics Research Institute, New Delhi.

Results and discussion

Morphological characterization of parental lines and their cross combinations

Eleven garden pea genotypes along with fifty-five F1 hybrids were assessed and characterized morphologically (Fig. 2). ANOVA for ten economically important quantitative traits like plant height (cm), node bearing first flower, days to 50% flowering, number of pods per plant, pod length (cm), pod width (cm), number of seeds per pod, pod weight (g), shelling percentage and pod yield per plant (g) is presented in Table 2. The mean squares due to genotypes were highly significant for all the characters that revealed significant variation exists among the genotypes for the traits studied.

Fig. 2.

Fig. 2

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Morphological representation of pod shape, size and colour of the parental lines used in the study [P1: Arkel; P2: GP 17; P3: Arka Ajit; P4: Pusa Pragati; P5: GP 473; P6: VP-233; P7: VRP-7; P8: VRP-6; P9: AP-3; P10: IP 3; P11: GP 6].

Table 2.

Analysis of variance for different quantitative traits in garden pea.

Source of variation d.f Plant height Node bearing first flower Days to 50% flowering Number of pods per plant Pod length Pod width Number of seeds per pod Pod weight Shelling percentage Yield per plant
Replication 2 10.00 0.98 12.10 5.10 0.05 0.02 0.41 0.21 10.31 10.56
Treatment 65 160.53** 3.32** 159.01** 43.16** 4.66** 0.32** 1.33** 1.60** 65.34** 636.94**
Parents 10 257.95** 3.44** 389.41** 31.78** 4.50** 0.21** 1.99** 2.60** 77.69** 834.78**
F1 54 144.10** 2.86** 107.56** 43.90** 4.66** 0.26** 1.17* 1.43** 62.95** 543.27**
PVF1 1 73.67** 27.12** 633.60** 117.00** 6.65** 4.46** 3.28** 0.40* 71.15** 3716.43**
Error 130 12.66 0.86 6.54 4.42 0.57 0.04 0.58 0.18 19.32 83.34
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*&

**Significant at 5% and 1% probability, respectively.

There was a wide range of variation for plant height (49.09–74.15 cm) among the parental lines with the mean value of 60.30 cm. The lowest plant height was observed in genotype P2 (49.09 cm) while, maximum plant height was recorded in P10 (74.15 cm). The plant height was ranged from 43.37 to 75.80 cm among crosses with the mean value of 59.66 cm. Among the F1 crosses, P7 × P11 was tallest (75.80 cm) followed by P6 × P8 (75.09 cm), P2 × P10 (69.92 cm) and P6 × P10 (68.89 cm). Among the parental lines, the number of node bearing first flower was ranged from 8.30 to 11.59 with the mean value of 10.30. The parent P11 (8.30) produced the first flower at lowest node whereas P7 (11.59) produced the first flower at the highest node. The number of node bearing first flower was ranged from 6.98 to 11.59 among the F1 crosses with the mean value of 9.31. Among the F1 crosses, P6 × P9 produced first flower at lowest node (6.98). Among the parental lines, the days to 50% flowering ranged from 43.67 to 72.00 days with general mean of 58.24 days. The parent, P2 (43.67) was earliest genotype whereas the parent P10 (72.00) took the maximum number of days to 50% flowering. The range for days to 50% flowering was 43.67 to 70.00 among the F1 crosses and mean was 53.71. Among the F1 crosses, P3 × P4 was earliest (43.67). The range of variation for number of pods per plant varied from 12.66 to 23.43 among the parental lines. Maximum number of pods per plant was recorded in genotype P10 (23.43) while, minimum number of pods per plant was recorded in P3 (12.66) with a mean of 19.17. The parents P2 (GP-17), P5 (GP-473) and P7 (VRP-7) had high GCA for most of the horticultural traits including yield per plant. Therefore, these were observed to be good combiners for these traits29. The range of variation for number of pods per plant varied from 14.88 to 31.07 among the crosses with mean value of 21.14. Among the F1 crosses, P2 × P7 had the highest number of pods per plant (31.07). Among the parental lines, the pod length was ranged from 6.47 to 9.73 cm with a mean of 7.95 cm. The pod length was highest in P6 (9.73 cm) whereas minimum pod length was recorded in P10 (6.47 cm). The range of variation for pod length varied from 5.09 to 10.46 cm among the crosses with mean value of 7.46 and cross P2 × P6 had the highest pod length (10.46 cm) followed by P3 × P10 (10.20 cm), P4 × P8 (9.94 cm) and P3 × P7 (9.83 cm) (Fig. 3). The parental lines varied from 1.08 to 1.92 cm with a mean of 1.55 cm for pod width. The pod width was highest in P6 (1.92 cm) whereas, minimum pod width was observed in P5 (1.08 cm). The range of variation for pod width varied from 1.47 to 2.46 cm among the crosses with mean value of 1.95 cm. Among the F1 crosses, P7 × P10 exhibited highest pod width (2.46 cm) (Fig. 4). The number of seeds per pod varied from 6.02 to 8.38 among the parental lines with a mean value of 7.38. Maximum number of seeds per pod was recorded in genotype P5 (8.38) while minimum number of seed per pod was found in P11 (6.02). The range of variation for number of seeds per pod varied from 6.25 to 9.08 among the crosses with mean value of 7.72. Among the F1 crosses, P1 × P7 had highest number of seeds per pod (9.08). The parental lines found to be varied from 2.58 to 6.14 g with mean value of 4.29 g for average pod weight. The highest average pod weight was recorded in P5 (6.14 g) while minimum pod weight was recorded in P10 (2.58 g). The range of variation for average pod weight varied from 2.15 to 5.70 g among the crosses with mean value of 4.17 g. Among the F1 crosses, P3 × P11 had highest pod weight (5.70 g). The shelling percentage of the parental lines was recorded to the tune of 33.10 to 50.37% with the mean value 42.81%. The highest shelling percentage was recorded in P5 (50.37%) while minimum shelling percentage was recorded in P3 (33.10%). The range of variation for shelling percentage varied from 37.81 to 54.09% among the crosses with mean value of 44.42% (Table 3). Among the F1 crosses, P6 × P9 was having highest shelling percentage (54.09%). The green pod yield per plant was found to be in a range of 64.46 to 107.90 g with the mean value of 81.15 g in the parental genotypes. The highest yield per plant was recorded in P7 (107.90 g) while minimum yield per plant was recorded in P10 (64.46 g) (Table 3). The range of variation for yield per plant varied from 69.28 to 136.51 g among the crosses with mean value of 92.77 g. Among the F1 crosses, P6 × P10 was highest yielder (136.51 g), followed by P2 × P8 (122.83 g), P8 × P10 (118.63 g) and P5 × P7 (114.32 g) (Table 4; Fig. 5). The crosses P6 × P10, P2 × P8 and P8 × P10 had higher SCA values for total yield per plant and were considered as most promising combinations for yield29. The results indicated the importance of recombinant breeding for effective utilization of non-additive genetic variance in garden pea. Ram et al.30 found that narrow sense heritability in garden pea was less than 50% for most of the yield contributing traits except plant height and days to 50% flowering, suggested the preponderance of non-additive gene action and commercial exploitation of recombinant breeding for improvement. Sharma et al.31 reported genetic diversity among 56 garden pea genotypes using 12 morphological descriptors out of which, eight descriptors were found polymorphic, and highest Shannon diversity index was recorded for pod curvature. Rana et al.32 studied genetic structure, diversity and inter-relationships in a worldwide collection of 151 pea accessions for 21 morphological descriptors.

Fig. 3.

Fig. 3

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Promising cross-combinations for pod length.

Fig. 4.

Fig. 4

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Promising cross-combinations for pod width.

Table 3.

Mean performance of the parents for quantitative traits in garden pea.

Parents Plant height (cm) Node bearing first flower Days to 50% flowering Number of pods per plant Pod length (cm) Pod width (cm) Number of seeds per pod Pod weight (g) Shelling percentage (%) Yield per plant (g)
P1 54.73de 10.30a-c 48.67d 21.14a-c 8.13a-c 1.48 cd 8.13a 3.75d 42.15b-d 70.71d
P2 49.09e 10.57ab 43.67d 21.08a-c 7.38bc 1.23de 7.25a-c 4.80b 45.12a-c 96.16ab
P3 66.60b 9.56b-d 64.33bc 12.66e 6.73bc 1.62bc 8.04a 4.99b 33.10e 65.18d
P4 51.08e 10.59ab 46.33d 17.56 cd 9.32a 1.23de 7.60ab 4.62bc 43.06b-d 86.81bc
P5 60.39 cd 10.23a-c 60.00c 17.70b-d 9.71a 1.08e 8.38a 6.14a 50.37a 106.34a
P6 62.79bc 11.29ab 71.67a 21.44ab 9.73a 1.92a 7.20a-c 4.15 cd 47.67ab 86.70bc
P7 73.84a 11.59a 69.67ab 21.48ab 6.57c 1.62bc 6.13c 4.79b 47.51ab 107.90a
P8 55.25de 10.62ab 47.67d 19.67a-c 8.00a-c 1.75ab 8.21a 3.70d 38.23de 64.89d
P9 72.30a 8.73 cd 48.33d 20.21a-c 8.35ab 1.62bc 7.47ab 3.58d 45.48a-c 77.22 cd
P10 74.15a 11.55a 72.00a 23.43a 6.47c 1.67a-c 6.71bc 2.58e 38.70de 64.46d
P11 54.09e 8.30d 68.33ab 14.55de 7.08bc 1.82ab 6.02c 4.07 cd 39.51c-e 66.26d
Mean 60.30 10.30 58.24 19.17 7.95 1.55 7.38 4.29 42.81 81.15
SE(d) 2.68 0.79 2.91 1.64 0.75 0.10 0.53 0.28 2.73 6.26
CD (p = 0.05) 5.64 1.67 6.12 3.44 1.57 0.21 1.10 0.59 5.73 13.16
C.V 5.36 9.44 6.121 10.46 11.51 8.03 8.72 8.06 7.80 9.45
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Table 4.

Mean performance of F1 crosses for quantitative traits in garden pea.

Crosses (F1) Plant height (cm) Node bearing first flower Days to 50% flowering Number of pods per plant Pod length (cm) Pod width (cm) Number of seeds per pod Pod weight (g) Shelling percentage (%) Yield per plant (g)
P1 × P2 52.37p-t 9.57b-k 52.33c-g 23.56d-j 8.63c-h 2.43ab 7.48b-j 4.36d-o 50.91a-f 93.40e-o
P1 × P3 57.26 h-r 9.93a-i 53.67c-f 23.52d-j 6.92j-o 1.47i 8.24a-g 2.99 s-v 47.18a-j 76.27o-s
P1 × P4 55.06 l-s 8.91e-l 47.67 g-k 17.38o-s 6.6j-o 2.16a-f 7.72a-j 4.46d-n 41.11 g-k 81.78j-s
P1 × P5 57.46 g-r 9.27c-k 48.33f.-k 21.39 h-p 6.9j-o 1.47i 8.32a-g 4.80b-h 40.95 g-k 91.99e-q
P1 × P6 60.35c-p 10.89a-d 52.67c-g 21.24 h-p 7.14i-o 2.16a-f 7.23e-j 4.05 g-q 40.51 h-k 82.84j-s
P1 × P7 59.70c-q 10.59a-e 46.00i-k 18.97 l-s 7.23 h-o 1.47i 9.08a 4.77bc-j 39.96 h-k 98.03d-l
P1 × P8 49.96q-t 9.24c-k 54.67c-e 17.65n-s 7.55 g-m 2.16a-f 7.45b-j 3.95i-q 51.34a-d 81.78j-s
P1 × P9 55.36 k-s 8.31 h-m 51.67c-h 18.92 l-s 7.67f.-l 2.19a-e 8.21a-g 4.01 h-q 39.75 h-k 72.17rs
P1 × P10 56.95 h-r 8.91e-l 53.67c-f 22.76e-l 6.73j-o 1.62hi 8.36a-g 3.86 l-r 41.08 g-k 95.58e-m
P1 × P11 57.75f.-r 9.97a-h 68.00ab 17.80n-s 6.82j-o 2.19a-e 7.21e-j 3.90 l-r 41.94 g-k 75.69o-s
P2 × P3 53.16n-t 8.61f.-m 46.67 h-k 17.68n-s 6.54j-p 1.97c-h 7.42c-j 4.27e-p 38.84jk 77.39 m-s
P2 × P4 52.88o-t 8.97e-l 52.33c-g 21.55 g-o 7.67f.-l 2.19a-e 7.34c-j 4.01 h-q 37.81 k 87.00f.-s
P2 × P5 52.64o-t 10.23a-g 51.00d-i 19.61j-q 6.73j-o 2.16a-f 7.40c-j 4.77b-k 42.21f.-k 103.6c-g
P2 × P6 53.08n-t 8.25 h-m 52.00c-h 23.55d-j 10.46a 1.62hi 8.29a-g 3.87 l-r 51.66a-c 97.41d-l
P2 × P7 63.50c-m 9.27c-k 52.00c-h 31.07a 7.09j-o 1.97c-h 7.19e-j 3.12r-u 52.23ab 108.83b-e
P2 × P8 49.47q-t 7.92j-m 52.00c-h 20.13i-q 7.14i-o 1.99c-h 8.02a-h 5.45ab 42.47e-k 122.83ab
P2 × P9 55.42j-s 11.59a 52.00c-h 19.35j-r 6.54j-p 1.47i 8.03a-h 3.95i-q 38.84jk 83.50i-s
P2 × P10 69.92a-c 10.89a-d 54.67c-e 19.68j-q 6.06n-p 1.62hi 7.16e-j 3.50p-t 40.85 g-k 76.43n-s
P2 × P11 58.88d-q 10.93a-c 67.33ab 19.99j-q 5.93op 1.62hi 7.79a-i 5.10a-d 39.30i-k 97.01d-l
P3 × P4 62.75c-o 8.58f.-m 43.67 k 15.96q-s 9.05b-f 1.51i 8.80a-c 4.08 g-q 50.99a-f 74.45q-s
P3 × P5 64.51c-m 8.58f.-m 56.00 cd 19.19 k-s 6.94jk-o 2.27a-c 7.41c-j 5.15a-d 43.27c-k 95.27e-m
P3 × P6 58.58e-q 9.90a-i 51.33c-i 22.76e-l 8.63c-h 2.43ab 8.38a-g 3.52p-t 41.12 g-k 86.41 g-s
P3 × P7 68.19a-e 10.93a-c 48.67f.-k 17.64n-s 9.83a-d 2.26a-d 7.47b-j 4.94a-f 43.26c-k 97.25d-l
P3 × P8 61.71c-p 8.25 h-m 52.33c-g 19.48j-r 9.71a-d 2.16a-f 7.37c-j 3.97i-q 48.63a-h 87.65f.-s
P3 × P9 56.47i-r 11.26ab 46.67 h-k 14.88 s 6.88j-o 2.07a-f 7.87a-h 4.51c-n 43.17c-k 74.80p-s
P3 × P10 55.26 k-s 9.24c-k 45.33jk 21.81f.-n 10.2ab 2.14a-f 7.19e-j 4.63c-m 41.03 g-k 104.74c-g
P3 × P11 65.62b-j 9.60b-k 65.00ab 15.33rs 6.25 l-p 1.97c-h 7.71a-j 5.70a 48.08a-i 89.73f.-r
P4 × P5 54.52 m-s 10.38a-f 54.67c-e 19.07 l-s 6.2 m-p 2.04b-g 8.32a-g 4.45d-n 46.38a-k 99.71d-k
P4 × P6 52.82o-t 9.73b-j 47.67 g-k 19.48j-r 8.86b-g 1.77f.-i 8.66a-e 4.78b-i 48.07a-i 105.09c-f
P4 × P7 66.18a-i 9.08d-l 48.00 g-k 18.18 m-s 7.97e-j 1.86d-i 6.25j 4.58c-m 41.49 g-k 94.84e-n
P4 × P8 58.59e-q 9.41c-k 52.33c-g 18.01 m-s 9.94a-c 1.77f.-i 8.11a-g 4.43d-o 45.05b-k 84.57i-s
P4 × P9 57.45 g-r 10.06a-h 51.67c-h 17.56n-s 5.09p 1.77f.-i 7.44b-j 4.03 h-q 40.70 g-k 81.60 k-s
P4 × P10 46.00st 9.41c-k 51.33c-i 24.3d-i 5.99n-p 1.86d-i 6.94 g-j 2.89t-v 39.39i-k 76.77n-s
P4 × P11 57.90f.-r 7.79 k-m 55.33c-e 21.60 g-o 9.34a-e 1.99c-h 8.26a-g 4.41d-o 43.22c-k 103.13c-h
P5 × P6 48.18r-t 9.41c-k 55.67c-e 19.80j-q 6.58j-o 1.77f.-i 7.86a-i 3.42qrst 52.66ab 88.69f.-r
P5 × P7 66.73a-h 8.11i-m 52.00c-h 27.00b-d 8.53d-i 1.77f.-i 7.27d-j 2.15v 51.13a-e 114.32b-d
P5 × P8 43.37t 9.73b-j 54.67c-e 18.14 m-s 6.20 m-p 1.99c-h 7.29c-j 4.77b-k 40.41 h-k 105.20c-f
P5 × P9 57.47 g-r 8.76e-m 55.33c-e 17.10p-s 5.98op 1.77f.-i 7.46b-j 5.32a-c 48.35a-h 90.57e-r
P5 × P10 60.59c-p 8.44 g-m 52.67c-g 19.80j-q 7.73f.-k 1.95c-h 7.15e-j 4.97a-e 40.51 h-k 100.16d-j
P5 × P11 67.65a-g 9.57b-k 64.00b 25.63b-g 6.81j-o 1.66 g-i 7.69a-j 3.87 l-r 50.94a-f 97.76d-l
P6 × P7 67.96a-f 8.31 h-m 54.67c-e 22.12e-m 7.67f.-l 2.19a-e 8.18a-g 4.01 h-q 42.29f.-k 99.26d-k
P6 × P8 75.09ab 9.90a-i 52.33c-g 24.72c-h 9.31a-e 1.50i 8.94ab 2.62uv 51.32a-d 69.28 s
P6 × P9 53.18n-t 6.98 m 57.00c 25.80b-f 8.97b-f 2.19a-e 7.84a-i 3.61o-t 54.09a 83.04j-s
P6 × P10 68.89a-d 9.93a-i 52.67c-g 29.39ab 7.44 h-n 1.47i 8.09a-g 4.16e-q 49.43a-g 136.51a
P6 × P11 62.63c-p 8.64f.-m 63.00b 28.98ab 7.01j-o 2.19a-e 8.47a-f 4.00 h-q 51.31a-d 93.36e-o
P7 × P8 64.79c-l 10.23a-g 50.33d-j 21.49 g-o 8.63c-h 2.43ab 8.78a-d 4.14f.-q 41.15 g-k 101.93c-i
P7 × P9 63.27c-n 9.24c-k 50.67d-j 20.93 h-p 5.94op 2.16a-f 6.56 h-j 3.83 m-r 40.31 h-k 93.20e-p
P7 × P10 62.13c-p 9.60b-k 55.67c-e 28.43a-c 6.88j-o 2.46a 7.62a-j 3.97i-q 41.97 g-k 99.63d-k
P7 × P11 75.80a 8.25 h-m 69.33a 15.99q-s 7.55 g-m 2.43ab 6.98f.-j 4.86b-g 42.36e-k 92.67e-q
P8 × P9 66.44a-i 9.27c-k 50.33d-j 27.44a-d 7.38 h-o 1.97c-h 7.20e-j 3.74n-s 47.61a-j 90.09f.-r
P8 × P10 66.87a-h 8.91e-l 54.33c-e 26.17b-e 6.47 k-p 1.83e-i 7.52b-j 4.36d-o 42.95c-k 118.63bc
P8 × P11 65.47b-k 7.29 lm 50.00e-j 25.82b-f 7.07j-o 1.79e-i 7.44b-j 4.69b-l 42.20f.-k 114.13b-d
P9 × P10 60.30c-p 9.24c-k 55.67c-e 23.46d-k 6.81j-o 1.96c-h 8.42a-g 3.48p-t 46.59a-k 85.01 h-s
P9 × P11 58.68d-q 8.91e-l 44.00 k 17.68n-s 7.55 g-m 2.43ab 6.35ij 4.02 h-q 39.90 h-k 80.55-s
P10 × P11 68.34a-e 8.91e-l 70.00a 21.12 h-p 6.47 k-p 1.62hi 7.40c-j 3.94j l-q 42.71d-k 89.04f.-r
Mean 59.66 9.31 53.51 21.24 7.46 1.95 7.72 4.17 44.42 92.77
SE(d) 4.15 0.75 2.29 1.743 0.59 0.16 0.68 0.34 3.59 7.48
CD (p = 0.05) 8.26 1.48 4.56 3.46 1.17 0.32 1.23 0.68 7.11 14.84
C.V 8.53 9.82 5.25 10.05 9.70 10.34 9.69 10.02 9.87 9.87
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Values within columns with different letters (superscript value) are significantly different according to Duncan’s test at P = 0.05.

Fig. 5.

Fig. 5

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Promising cross-combinations for yield.

Screening for powdery mildew resistance

The screening data viz., per cent disease index (PDI) and performance of 11 genotypes of garden pea for Powdery mildew reaction at harvesting stage is given in Tables 5, 6 and Fig. 6. The results revealed that GP-6 was found to be the most resistant genotype to powdery mildew as it exhibited minimum percent disease index (1.85). Four genotype Arka Ajit, Pusa Pragati, GP-473 and VP-233 exhibited percent disease index (PDI) in the range of 2.54 to 7.78 and were considered resistant. Similar results were also recorded by Sharma et al.33. The genotypes like Arkel, GP-17, VRP-7 and AP-3 were susceptible, with a PDI range of 65.34 to 74.71. Similarly, Pandey et al.34 evaluated 116 pea germplasm lines in Uttar Pradesh, India, and observed that most of the lines (43) were highly susceptible, with PDI values from 76.0—96.0. Azmat et al.35 also evaluated 146 pea accessions, collected from different countries, were screened against powdery mildew and observed that accessions 9057, 9370, 9375, 10,609, 10,612, 18,293, 18,412, 19,598, 19,611, 19,616, 19,727, 19,750, 19,782, 20,126, 20,152, 20,171, It-96, No. 267, and No. 380 were resistant and accessions It-96 and No. 267 were highly resistant. Parent IP-3 was highly susceptible to powdery mildew at scoring reaction of 8, which exhibited maximum PDI value of 75.92. The genotype VRP-6 was found moderately susceptible, exhibiting score of 6 with PDI value of 48.45. These results were in agreement with the results reported by Singh et al.36. VP-233 (Vivek Matar 11) was also highly resistant genotype as reported by Hooda et al. 37, Sharma and Sharma38 and Hedau et al.39.

Table 5.

Reaction of garden pea genotypes for powdery mildew disease at harvesting stage.

Genotypes 0–9 scale PDI Reaction
Arkel 7 73.16 S
GP-17 7 68.76 S
Arka Ajit 2 5.55 R
Pusa Pragati 2 7.78 R
GP-473 2 3.40 R
VP-233 2 2.54 R
VRP-7 7 65.34 S
VRP-6 6 48.45 MS
AP-3 7 74.41 S
IP-3 8 75.92 HS
GP-6 1 1.85 R
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R, Resistant, MR, Moderately Resistant, MS, Moderately Susceptible, S, Susceptible and HS, Highly Susceptible, PDI, Percent disease index.

Table 6.

Grouping of genotypes on the basis of disease reaction scale.

Immune (0) Resistant (0–10) Moderately Resistant (11–25) Moderately Susceptible (26–50) Susceptible (51–75) Highly Susceptible (> 75)
– Arka Ajit, Pusa Pragati, GP-473, VP-233, GP-6 – VRP-6 Arkel, GP-17, VRP-7, AP-3 IP-3
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Fig. 6.

Fig. 6

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Graphical representation (bar charts) for PDI of genotypes.

Genetic inheritance for powdery mildew resistance

The knowledge regarding inheritance pattern of traits is helpful in formulating appropriate breeding programme for developing need-based crop varieties. The overall results indicated that lines GP-6, GP-473, VP-233, Pusa Pragati and Arka Ajit were found resistant to Powdery mildew disease which could be further used in breeding programme for the development of resistant genotypes while Arkel, AP-3, VRP-7, VRP-6, &GP-17 were susceptible and IP-3 was highly susceptible to powdery mildew in the present study.

The reaction of plants to powdery mildew was analysed using different generations derived from susceptible × resistant cross combinations (Arkel × GP-6, VRP-6 × GP-6, Arkel × VP-233, AP-3 × GP-6, AP-3 × GP-473). On the perusal of data (Table 7), it showed that all plants of parental line VRP 6 were susceptible while plant populations of GP-6 were resistant. All the F1 plants of cross Arkel × GP-6 were susceptible. Out of 158 F2 plants screened for disease reaction, 116 plants were found susceptible and 42 plants showed resistance to powdery mildew. The segregation ratio of F2 plants fitted well in the expected ratio of 3:1 as susceptible and resistant respectively, which is evident from the non-significant χ2 value of 0.216 (P = 0.642). A total of 25 plants of BC1 generation segregated into 14 susceptible: 11 resistant and fitted well with expected ratio (1:1). However, all the 25 plants of BC2 were susceptible.

Table 7.

Segregation of garden pea genotype response against powdery mildew and their probability in the F2 and back-cross generations.

Cross No of plants Observed frequency Ratio S:R χ2 value (cal.)* P** value (d.f = 1)
S R
Arkel × GP-6
Arkel 42 42 0 – – –
GP-6 42 0 42 – – –
F1 35 35 0 – – –
F2 158 116 42 2.76:1 0.216 0.5–0.75 (0.642)
BC1 25 14 11 1:1 0.360 0.5–0.75 (0.548)
BC2 25 25 0 1:0  – –
VRP-6 × GP-6
VRP-6 45 45 0 – – –
GP-6 42 0 42 – – –
F1 50 50 0 – – –
F2 110 78 32 2.43:1 0.981 0.25–0.5(0.321)
BC1 35 18 17 1:1 0.014 0.9–0.95(0.905)
BC2 35 35 0 1:0 – –
Arkel × VP-233
Arkel 42 42 0 – – –
VP-233 40 0 40 – – –
F1 50 50 0 – – –
F2 120 86 34 2.52:1 0.710 0.25–0.5(0.399)
BC1 50 27 23 1:1 0.320 0.5–0.75(0.571)
BC2 50 50 0 1:0  – –
AP-3 × GP-473
AP-3 55 55 0 – – –
GP-473 55 0 55 – – –
F1 40 40 0 – – –
F2 130 101 29 3.64:1 0.402 0.5–0.75 (0.526)
BC1 45 23 22 1:1 0.011 0.9–0.95 (0.916)
BC2 45 45 0 1:0 – –
AP-3 × GP-6
AP-3 55 55 0 – – –
GP-6 42 0 42 – – –
F1 40 40 0 – – –
F2 90 69 21 3.28:1 0.133 0.50–0.75 (0.715)
BC1 30 13 17 1:1 0.266 0.5–0.75 (0.606)
BC2 30 30 0 1:0 – –
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*Tabulated value for the P = 0.05 significance level is 3.84. ** Figures in parentheses are exact probability values.

Similarly, in cross VRP-6 × GP-6, GP-6 is a highly resistant parent and free from any symptoms in field while all the plants of VRP-6 were susceptible. All the F1 plants of this cross were susceptible. Out of 110 F2 plants of cross VRP-6 × GP-6, 78 plants were susceptible and 32 plants were found resistant. The segregation of susceptible and resistant in the F2 population was tested for agreement with the Mendelian ratio of 3 (susceptible):1 (resistant) using the χ2 test (χ2:0.981; p = 0.321). The back-cross (BC1) population using ‘GP-6’ as the recurrent parent, fitted the expected 1:1 ratio of resistant: susceptible as out of 35 plants in BC1, 18 were susceptible and 17 were observed as resistant whereas the observed ratio 35:0 was recorded in BC2.

In cross Arkel × VP-233, VP-233 is resistant parent, and it’s all plants were free from disease symptoms while plant population of Arkel were highly susceptible. In the F1 generation, all the 50 plants were found susceptible. Among 120 F2 plants, 86 were susceptible and 34 plants showed resistant. On the comparison of observed segregation ratio of 2.52:1 with the expected ratio 3: 1 in a χ2 test, 3:1 segregation ratio was observed with a non-significant χ2 value of 0.710 (P = 0.399) that fitted well with the expected ratio (Table 7 and Fig. 7). A total of 50 plants of BC1 generation segregated as 27 susceptible: 23 resistant and fitted well with expected ratio (1:1). However, the 50:0 ratio of BC2 indicating the recessive gene present in VP 233.

Fig. 7.

Fig. 7

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Graphical representation (bar charts) for segregation ratios.

In cross AP-3 × GP-473, all 55 plants of AP-3 were susceptible while all 55 plants of GP-473 were resistant. A total of 40 plants in the F1 generation were found susceptible. Among 130 F2 plants, segregation pattern of observed frequency 101:29 demonstrated a good fit to one gene model with an expected ratio of 3:1 with estimated χ2 value of 0.402 (P = 0.526). The non-significance of the χ2 test where P value > 0.05, indicated close agreement between the observed and expected ratio of susceptible: resistant plants so null hypothesis is accepted.

Similarly, in cross AP-3 × GP-6, all plants of AP-3 were susceptible, and all plants of GP-6 were resistant. All F1 plants were susceptible and F2 population was having ratio of 69: 21 (Susceptible: Resistant) as observed frequency with estimated χ2 value of 0.133 (P = 0.715). The non-significance of the χ2 test where P value > 0.05, indicated close agreement between the observed and expected ratio of susceptible: resistant plants so null hypothesis is accepted. Backcross generations were also evaluated for confirmation of segregation pattern in F2 population. The frequency distributions of resistance and susceptibility in plants of the two backcross generations i.e., BC1 14 Susceptible:11 Resistant and for BC2 25 Susceptible: 0 Resistant tested for their best-fit with classical Mendelian ratios i.e., 1:1 in backcross progenies with a resistant parent (BC1) and 1:0 backcross progenies with a susceptible parent (BC2). Verma et al.40 used chi-square analysis to know the inheritance pattern of downy mildew resistance in cauliflower and reported the segregation of F2 generations in the ratio 3:1 in DPCaY-3 × DPCaY-8, DPCaY-3 × DPCaY-6, and PU × DPCaY-6 which suggests the monogenic inheritance of resistance in these crosses. This was further confirmed by the test cross ratio of 1:1 in a backcross with the susceptible parent in all the three crosses. Rani et al.41 also reported segregation ratio of 3:1 and 1:1 ratio in F2 and BC2F1 generations for resistance and susceptibility to the isolates of Fusarium wilt race 3 and race 4, indicating monogenic resistance to each race in F2 (BGD72 × WR315) and backcross (BGD72 × WR315) × BGD72 generations of chickpea using chi-square analysis. Leon et al.42 reported that the crosses of pea resistant line ILS6527 with two susceptible pea varieties (Andina and San Isidro) segregated in a monogenic Mendelian ratio of 3 susceptible: 1 resistant in F2 generation, and 1 susceptible: 1 resistant ratio in B1 generation, which indicated a recessive control of resistance. The resistance gene in ILS6527 may be allelic with er-1. UN6651 exhibited PM resistance only in pods, the F2 and backcrosses from the crosses with UN6651 segregated in a manner indicative of recessive resistance. They also reported PM resistance only in pods of UN6651, the F2 and backcrosses from the crosses of UN6651 with two susceptible varieties (Andina and San Isidro) segregated in a manner indicative of recessive resistance.

Since Segregation analysis in all the cross combination (Arkel × GP-6, VRP-6 × GP-6, Arkel × VP-233, AP-3 × GP-6, AP-3 × GP-473) for powdery mildew resistance gene in three populations (F2, B1, B2) fitted well in expected ratios and no complementation was observed in all the crosses therefore, it may be suggested that the resistance against powdery mildew disease in VP-233, GP-6 and GP-473 genotypes is governed by monogenic and recessive in nature. This type of segregation and monogenic recessive gene also reported by Tiwari et al.24, Kaur et al.43, Kumar et al.44, Aghora et al.45, Kushwah and Sharma46 and Azmat and Khan47. Thus, it has been proved that powdery mildew resistance is controlled by single recessive gene in these genotypes. Previous studies that looked into the inheritance of powdery mildew resistance found that the pea accessions had single recessive42,48, single dominant17,49 and duplicate recessive gene actions50. Till date, three resistance genes have been reported to provide resistance to plants to differing degrees thus far. One of these three genes is dominant (Er3), while the other two are recessive (er1 and er2). The gene er1, provide resistance of moderate to complete level, is location-specific51. Furthermore, it has been discovered that only a few resistant pea accessions carry the resistance gene er219. Research on the Er3 gene is still in its early stages, and few information is available.

Conclusion

In our study the genotypes like Arkel, GP-17, VRP-7 and AP-3 were susceptible IP-3 was highly susceptible and VRP-6 was moderately susceptible to powdery mildew disease. The genotypes like GP-6, GP-473, Arka Ajit, Pusa Pragati and VP-233 were resistant to powdery mildew. This study also unfolds the inheritance pattern of powdery mildew resistance and the phenotype of F1 progenies confirms the recessive nature of powdery mildew resistance. The segregation pattern observed in the F2 and backcross generations aligned with expectations for a recessive gene governing powdery mildew resistance. Chi-square analysis further supported the proposed inheritance model. This type of segregation suggested that the resistance in GP-6, GP-473 and VP-233 to powdery mildew is governed by a single recessive gene. These resistant genotypes could be used in introgression of powdery mildew resistance genes in garden pea to develop high yielding and disease resistant garden pea varieties. In this study, the validation of powdery mildew resistance using molecular marker was not done which limits the precision of confirming the genetic basis of powdery mildew resistance. Furthermore, the evaluation was conducted only at a single location, which may not represent the full range of environmental conditions influencing disease pressure. Therefore, molecular marker assisted validation of powdery mildew resistance and multi-location with multi-season evaluations is essentially required in future study to assess the stability of resistance across diverse environmental conditions and pathogen populations.

Acknowledgements

The authors are thankful to the Director, ICAR-Indian Agricultural Research Institute, New Delhi for providing the facilities.

Author contributions

Hanuman Ram : Conceptualization, Methodology, Investigation, Original draft. Shri Dhar and H. Choudhary : Methodology, Investigation. L. Prasad : Artificial inoculum for disease screening. B. S. Tomar : Review & Editing. G. S. Jat : Writing, review and editing, Statistical analysis and Observations.

Funding

Not applicable.

Data availability

Data will be made available on request to first author (hramdhanari@gmail.com).

Declarations

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Contributor Information

Hanuman Ram, Email: hramdhanari@gmail.com.

G. S. Jat, Email: singhgograj@gmail.com

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Associated Data

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Data Availability Statement

Data will be made available on request to first author (hramdhanari@gmail.com).

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