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Research article · 2025 · Full text

International Journal of Agriculture and Nutrition

Host plant resistance mechanisms in chickpea against pod borer Helicoverpa armigera

Alok Thorat

Volume 7(6) · Pages 84–86 · DOI: 10.33545/26646064.2025.v7.i6b.423 ↗

Research areasInsect-Plant Interactions and ControlGenetic and Environmental Crop StudiesAgricultural pest management studies

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Abstract

Author-written

Across South Asia, Helicoverpa armigera destroys 20-40% of chickpea pods each season despite heavy insecticide use. Resistant varieties could break this cycle, but the specific mechanisms need mapping before breeders can target them efficiently. This research screened five chickpea genotypes for antibiosis, antixenosis, and tolerance to H. armigera under laboratory and field conditions at Bhubaneswar, Odisha, during Rabi 2022-2023. ICC-506 showed the strongest antibiosis: larval survival on detached pods fell to 11% by day 14, and mean larval weight was 48% lower than on the susceptible ICC-3137. Trichome density on pods of ICC-506 was 8.7 per mm2 versus 3.1 in ICC-3137, and oxalic acid content was 3.82 mg g-1 compared with 1.68 mg g-1. Pod damage under open-field conditions was 12.4% on ICC-506 versus 38.7% on ICC-3137. These results identify ICC-506 as a strong antibiosis source linked to trichome and organic acid traits suitable for marker-assisted breeding programmes.

Summary

Source-checked

Plain-language reading of the full paper, prepared with AI and checked against the text. Each point links to the authors' own words.

  1. Why it matters

    • The pod borer Helicoverpa armigera is a major pest in South Asia, causing 20-40% pod loss in chickpeas despite the use of insecticides. View evidence ↗

      Why it matters

      The pod borer Helicoverpa armigera is a major pest in South Asia, causing 20-40% pod loss in chickpeas despite the use of insecticides.

      In the paper's own words

      “Across South Asia, Helicoverpa armigera destroys 20-40% of chickpea pods each season despite heavy insecticide use.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • Chemical control of H. armigera is difficult because the larvae bore into pods, which physically shields them from insecticides. View evidence ↗

      Why it matters

      Chemical control of H. armigera is difficult because the larvae bore into pods, which physically shields them from insecticides.

      In the paper's own words

      “The polyphagous caterpillar feeds on leaves during early instars and then bores into developing pods, consuming the grain inside—a behaviour that makes chemical control difficult because the larva is physically shielded once it enters the pod”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • Chickpea plants can defend themselves using glandular trichomes that secrete organic acids, such as oxalic and malic acid, which deter or kill young larvae. View evidence ↗

      Why it matters

      Chickpea plants can defend themselves using glandular trichomes that secrete organic acids, such as oxalic and malic acid, which deter or kill young larvae.

      In the paper's own words

      “Physical barriers like glandular trichomes secrete sticky exudates containing organic acids—chiefly oxalic and malic acid—that are toxic or deterrent to young larvae”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
  2. The question

    • The study aimed to rank five chickpea genotypes by their resistance mechanisms, link this resistance to pod-surface traits, and identify suitable genotypes for breeding in eastern India. View evidence ↗

      The question

      The study aimed to rank five chickpea genotypes by their resistance mechanisms, link this resistance to pod-surface traits, and identify suitable genotypes for breeding in eastern India.

      In the paper's own words

      “This research aimed to (i) rank five genotypes for antibiosis, antixenosis, and tolerance, (ii) link resistance to measurable pod-surface traits, and (iii) identify genotypes suitable for breeding in eastern India.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
  3. What they did

    • The researchers tested five genotypes: ICC-506, ICC-12475, ICCL-87322, ICC-3137, and the local cultivar 'Radhey'. View evidence ↗

      What they did

      The researchers tested five genotypes: ICC-506, ICC-12475, ICCL-87322, ICC-3137, and the local cultivar 'Radhey'.

      In the paper's own words

      “Five chickpea genotypes with varying resistance reports were obtained from the ICRISAT gene bank: ICC-506 (resistant), ICC-12475 (moderately resistant), ICCL-87322 (intermediate), ICC-3137 (susceptible), and a local cultivar ‘Radhey’ as field check.”

      Page 2

      Found in: Materials

      Read it in the full textOpen PDF at page 2 ↗
    • Field experiments were conducted in Bhubaneswar, Odisha, during November 2022 and 2023 using a randomized block design with four replications. View evidence ↗

      What they did

      Field experiments were conducted in Bhubaneswar, Odisha, during November 2022 and 2023 using a randomized block design with four replications.

      In the paper's own words

      “Seeds were sown in November 2022 and 2023 at the Orissa University of Agriculture and Technology (OUAT) research farm, Bhubaneswar (20°15′ N, 85°50′ E, 25 m a.s.l.), in a randomised block design with four replications, plot size 4 m × 3 m”

      Page 2

      Found in: Materials

      Read it in the full textOpen PDF at page 2 ↗
    • Antibiosis was measured using a no-choice detached-pod method with ten replicates per genotype, introducing one neonate larva per pod. View evidence ↗

      What they did

      Antibiosis was measured using a no-choice detached-pod method with ten replicates per genotype, introducing one neonate larva per pod.

      In the paper's own words

      “Laboratory antibiosis assays used a no-choice detached-pod method. Fresh pods at the R5 stage were placed individually in ventilated plastic cups (250 mL) lined with moist filter paper. One neonate H. armigera larva... was introduced per cup. Ten replicates per genotype were set up”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • Pod-surface traits were measured by counting trichome density under a microscope and quantifying organic acids using HPLC. View evidence ↗

      What they did

      Pod-surface traits were measured by counting trichome density under a microscope and quantifying organic acids using HPLC.

      In the paper's own words

      “Trichome density was counted under a stereo microscope at 40× on five randomly selected pods per plot. Organic acids (oxalic and malic) in pod walls were extracted with 0.1 M HCl and quantified by HPLC.”

      Page 2

      Found in: Methods

      Read it in the full textOpen PDF at page 2 ↗
  4. What they found

    • ICC-506 showed the strongest antibiosis, with only 11% larval survival by day 14, compared to 48% survival on the susceptible ICC-3137. View evidence ↗

      What they found

      ICC-506 showed the strongest antibiosis, with only 11% larval survival by day 14, compared to 48% survival on the susceptible ICC-3137.

      In the paper's own words

      “By day 14, only 11% of larvae survived on ICC-506 versus 48% on ICC-3137.”

      Page 2

      Found in: Results

      Read it in the full textOpen PDF at page 2 ↗
    • Larvae feeding on ICC-506 had a significantly lower mean weight (87.3 mg) than those on ICC-3137 (168.4 mg) by day 14. View evidence ↗

      What they found

      Larvae feeding on ICC-506 had a significantly lower mean weight (87.3 mg) than those on ICC-3137 (168.4 mg) by day 14.

      In the paper's own words

      “Mean larval weight at day 14 was 87.3 mg on ICC-506 and 168.4 mg on ICC-3137.”

      Page 2

      Found in: Results

      Read it in the full textOpen PDF at page 2 ↗
    • Field results mirrored laboratory findings, with pod damage being lowest on ICC-506 (12.4%) and highest on ICC-3137 (38.7%). View evidence ↗

      What they found

      Field results mirrored laboratory findings, with pod damage being lowest on ICC-506 (12.4%) and highest on ICC-3137 (38.7%).

      In the paper's own words

      “Pod damage in the field ranged from 12.4% on ICC-506 to 38.7% on ICC-3137.”

      Page 2

      Found in: Results

      Read it in the full textOpen PDF at page 2 ↗
    • ICC-506 demonstrated higher tolerance than ICC-3137, with a tolerance ratio of 0.86 compared to 0.58. View evidence ↗

      What they found

      ICC-506 demonstrated higher tolerance than ICC-3137, with a tolerance ratio of 0.86 compared to 0.58.

      In the paper's own words

      “The tolerance ratio (yield under infestation/yield under protection) was 0.86 for ICC-506 and 0.58 for ICC-3137”

      Page 3

      Found in: Comprehensive Interpretation

      Read it in the full textOpen PDF at page 3 ↗
    • In antixenosis tests, 73% of larvae migrated away from ICC-506 pods within 12 hours, while only 41% migrated away from ICC-3137. View evidence ↗

      What they found

      In antixenosis tests, 73% of larvae migrated away from ICC-506 pods within 12 hours, while only 41% migrated away from ICC-3137.

      In the paper's own words

      “The dual-choice antixenosis assay showed 73% of larvae migrated away from ICC-506 pods within 12 hours, compared with 41% away from ICC-3137”

      Page 3

      Found in: Comprehensive Interpretation

      Read it in the full textOpen PDF at page 3 ↗
  5. What it means

    • The authors conclude that ICC-506 is the most resistant genotype, with its resistance driven by high trichome density and oxalic acid levels. View evidence ↗

      What it means

      The authors conclude that ICC-506 is the most resistant genotype, with its resistance driven by high trichome density and oxalic acid levels.

      In the paper's own words

      “ICC-506 displayed the strongest resistance to H. armigera, driven by high trichome density (8.7 per mm2) and elevated oxalic acid (3.82 mg g-1) in pod walls.”

      Page 3

      Found in: Conclusion

      Read it in the full textOpen PDF at page 3 ↗
    • The authors recommend that breeders in eastern India use trichome density and oxalic acid content as markers for developing resistant chickpea varieties. View evidence ↗

      What it means

      The authors recommend that breeders in eastern India use trichome density and oxalic acid content as markers for developing resistant chickpea varieties.

      In the paper's own words

      “Breeders targeting eastern India should prioritise trichome density and oxalic acid content as selection markers, ideally linked to molecular markers for marker-assisted introgression.”

      Page 3

      Found in: Conclusion

      Read it in the full textOpen PDF at page 3 ↗
    • The authors state that multi-location validation across different seasons is needed to strengthen their recommendations. View evidence ↗

      What it means

      The authors state that multi-location validation across different seasons is needed to strengthen their recommendations.

      In the paper's own words

      “Multi-location validation across seasons will strengthen these recommendations”

      Page 3

      Found in: Conclusion

      Read it in the full textOpen PDF at page 3 ↗

Visual summary

2 visuals

Illustrations explain the paper; charts are replotted from the paper's own table values.

Replotted from source data
Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.↗ Enlarge
Larval survival 14d (%): ICC-506 vs ICC-12475 vs ICCL-87322 vs ICC-3137 vs Radhey
Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.

Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.

Open full-size image ↗

Larval survival 14d (%): ICC-506 vs ICC-12475 vs ICCL-87322 vs ICC-3137 vs Radhey

Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.

View source evidence ↗
Replotted from source data
Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.↗ Enlarge
Larval wt (mg): ICC-506 vs ICC-12475 vs ICCL-87322 vs ICC-3137 vs Radhey
Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.

Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.

Open full-size image ↗

Larval wt (mg): ICC-506 vs ICC-12475 vs ICCL-87322 vs ICC-3137 vs Radhey

Replotted from Table 1 (page 2) of the original paper. Every bar is the paper's own table value.

View source evidence ↗

Figures & tables

From the original paper
Antibiosis parameters and field pod damage across five chickpea genotypesTable 1Page 2Larval survival curves of H. armigera on detached pods of resistant (ICC-506), intermediate (ICCL-87322), and susceptible (ICC-Figure 1Page 2Heatmap of resistance mechanism traits across five chickpea genotypes. Darker shading indicates higher trait expression.Figure 2Page 3
FULL TEXT Original article · reflowed for reading

Introduction

According to ICRISAT's long-term monitoring programme, Helicoverpa armigera (Hübner) accounts for more economic loss in chickpea than all other insect pests combined [1] . The polyphagous caterpillar feeds on leaves during early instars and then bores into developing pods, consuming the grain inside-a behaviour that makes chemical control difficult because the larva is physically shielded once it enters the pod [2] . Host plant resistance (HPR) offers a preventive strategy that costs the farmer nothing beyond the seed price. Resistance in chickpea operates through three recognised categories: antibiosis (traits that harm or retard the insect after feeding begins), antixenosis (traits that deter feeding or oviposition), and tolerance (the plant's ability to compensate for damage without yield loss) [3] . Physical barriers like glandular trichomes secrete sticky exudates containing organic acids-chiefly oxalic and malic acid-that are toxic or deterrent to young larvae [4,5] . Despite two decades of screening, the number of genotypes with confirmed, field-validated resistance remains small, and most data come from peninsular India rather than the eastern coastal belt where pest pressure peaks during mild winters [6] . This research aimed to (i) rank five genotypes for antibiosis, antixenosis, and tolerance, (ii) link resistance to measurable pod-surface traits, and (iii) identify genotypes suitable for breeding in eastern India.

Quality Control of Bioassays

Neonates were obtained from a single egg batch to minimise genetic variability. Temperature in the bioassay room was held at 27±1 °C and relative humidity at 65±5%. A positive control (susceptible ICC-3137) was included in every batch to verify insect vigour. Any replicate where the control larva died before day 3 was discarded and re-established. Pod moisture was standardised by harvesting all genotypes on the same morning and conducting assays within 4 hours of detachment [8] .

Materials and Methods

Materials and Methods

Materials

Five chickpea genotypes with varying resistance reports were obtained from the ICRISAT gene bank: ICC-506 (resistant), ICC-12475 (moderately resistant), ICCL-87322 (intermediate), ICC-3137 (susceptible), and a local cultivar 'Radhey' as field check. Seeds were sown in November 2022 and 2023 at the Orissa University of Agriculture and Technology (OUAT) research farm, Bhubaneswar (20°15′ N, 85°50′ E, 25 m a.s.l.), in a randomised block design with four replications, plot size 4 m × 3 m [9] .

Methods

Trichome density was counted under a stereo microscope at 40× on five randomly selected pods per plot. Organic acids (oxalic and malic) in pod walls were extracted with 0.1 M HCl and quantified by HPLC. Pod damage in the field was assessed at maturity by counting bored vs. total pods on 20 plants per plot. Tolerance was estimated as the ratio of yield under natural infestation to yield under insecticide-protected conditions [10] . Data were analysed by ANOVA in R (v4.3.1), means compared by Tukey's HSD (p = 0.05).

Results

Larval survival dropped sharply on resistant genotypes (Table 1). By day 14, only 11% of larvae survived on ICC-506 versus 48% on ICC-3137. Mean larval weight at day 14 was 87.3 mg on ICC-506 and 168.4 mg on ICC-3137. Trichome density and oxalic acid content showed strong negative correlations with larval survival (r =-0.93 and-0.89, respectively). Pod damage in the field ranged from 12.4% on ICC-506 to 38.7% on ICC-3137.

Table 1. Antibiosis parameters and field pod damage across five chickpea genotypes
GenotypeLarval survival 14d (%)Larval wt (mg)Trichomes (/mm²)Oxalic acid (mg/g)Pod damage (%)
ICC-50611.087.38.73.8212.4
ICC-1247518.0104.27.23.1417.8
ICCL-8732227.0131.65.42.4324.6
ICC-313748.0168.43.11.6838.7
Radhey36.0147.84.21.9731.2
CD (p = 0.05)6.314.70.90.384.2

Table text extracted from the original PDF; superscript letters are shown inline. The original table image is below.

View original table image
Antibiosis parameters and field pod damage across five chickpea genotypes↗ Enlarge
Table 1
Antibiosis parameters and field pod damage across five chickpea genotypes

Antibiosis parameters and field pod damage across five chickpea genotypes

Open full-size image ↗
Original paper tableSource PDF · page 2 ↗

Comprehensive Interpretation

The tolerance ratio (yield under infestation/yield under protection) was 0.86 for ICC-506 and 0.58 for ICC-3137, confirming that ICC-506 combines antibiosis with moderate tolerance-a dual mechanism that should slow pest adaptation. The dual-choice antixenosis assay showed 73% of larvae migrated away from ICC-506 pods within 12 hours, compared with 41% away from ICC-3137 [11] .

Discussion

The 11% larval survival on ICC-506 is among the lowest reported in chickpea antibiosis assays, comparable to the 13-15% recorded on ICCV-10 in peninsular India [3,4] . The tight correlation between trichome density and larval mortality (r =-0.93) aligns with the hypothesis that glandular trichomes physically trap early-instar larvae while delivering toxic oxalic acid directly to the cuticle [5] . Malic acid content also differed among genotypes but contributed less to resistance than oxalic acid, consistent with in-vitro toxicity tests showing that oxalic acid is 3-4 times more lethal to neonate Helicoverpa than malic acid at equivalent concentrations [12] . The field pod damage results validated the laboratory rankings, which is important because laboratory-to-field consistency is often poor when environmental conditions dilute biochemical defences [13] .

Conclusion

ICC-506 displayed the strongest resistance to H. armigera, driven by high trichome density (8.7 per mm 2 ) and elevated oxalic acid (3.82 mg g -1 ) in pod walls. Field pod damage was limited to 12.4%, less than a third of the susceptible check. ICC-12475 is a secondary source. Breeders targeting eastern India should prioritise trichome density and oxalic acid content as selection markers, ideally linked to molecular markers for marker-assisted introgression. Multilocation validation across seasons will strengthen these recommendations [14] .

Figures and tables from the paper

Figure 1. Larval survival curves of H. armigera on detached pods of resistant (ICC-506), intermediate (ICCL-87322), and susceptible (ICC-
Larval survival curves of H. armigera on detached pods of resistant (ICC-506), intermediate (ICCL-87322), and susceptible (ICC-↗ Enlarge
Figure 1
Larval survival curves of H. armigera on detached pods of resistant (ICC-506), intermediate (ICCL-87322), and susceptible (ICC-

Larval survival curves of H. armigera on detached pods of resistant (ICC-506), intermediate (ICCL-87322), and susceptible (ICC-

Open full-size image ↗
Original paper figureSource PDF · page 2 ↗
Figure 2. Heatmap of resistance mechanism traits across five chickpea genotypes. Darker shading indicates higher trait expression.
Heatmap of resistance mechanism traits across five chickpea genotypes. Darker shading indicates higher trait expression.↗ Enlarge
Figure 2
Heatmap of resistance mechanism traits across five chickpea genotypes. Darker shading indicates higher trait expression.

Heatmap of resistance mechanism traits across five chickpea genotypes. Darker shading indicates higher trait expression.

Open full-size image ↗
Original paper figureSource PDF · page 3 ↗

References

14 in the original paper
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  3. Sharma HC, Ortiz R. Host plant resistance to insects: an eco-friendly approach for pest management. Journal of Environmental Biology. 2002;23(2):111-135.
  4. Lateef SS. Gram pod borer and its management with special reference to host plant resistance. Indian Journal of Pulses Research. 1985;1(1):1-16.
  5. Yoshida M, Cowgill SE, Wightman JA. Roles of oxalic and malic acids in chickpea trichome exudate in hostplant resistance to Helicoverpa armigera. Journal of Chemical Ecology. 1997;23(4):1195-1210.
  6. Srinivasan S, Sridhar S, Vijayakumar J. Helicoverpa management in pulse crops of eastern India. Pest Management in Horticultural Ecosystems. 2016;22(1):1-10.
  7. Armes NJ, Bond GS, Cooter RJ. The laboratory culture and development of Helicoverpa armigera. Natural Resources Institute Bulletin. 1992;57:1-22.
  8. Narayanamma VL, Sharma HC, Gowda CLL, Sriramulu M. Mechanisms of resistance to Helicoverpa armigera in chickpea. Euphytica. 2007;154(1):117-126.
  9. ICRISAT. ICRISAT chickpea germplasm catalogue. International Crops Research Institute for the SemiArid Tropics. Patancheru; 2018.
  10. Smith CM. Plant resistance to arthropods: molecular and conventional approaches. Springer. Dordrecht; 2005.
  11. Green PWC, Stevenson PC, Simmonds MSJ, Sharma HC. Phenolic compounds on the pod-surface of pigeonpea mediate feeding behaviour of larvae of Helicoverpa armigera. Journal of Chemical Ecology. 2003;29(3):811-821.
  12. Bhatt NJ, Patel RK. Screening of chickpea cultivars for resistance to gram pod borer. Indian Journal of Entomology. 2001;63(3):277-280.
  13. Sharma HC, War AR, Pathania M, Munghate RS, Nikam PB, Sharma SP. Elevated CO 2 influences host plant defence response in chickpea against Helicoverpa armigera. Arthropod-Plant Interactions. 2016;10(2):171-181.
  14. Patil SB, Goyal A, Chitgupekar SS, Kumar S, ElBouhssini M. Sustainable management of chickpea pod borer. Agronomy. 2017;7(3):52.