Exploring the synergistic effects of drought and heat stress on chickpea seed development: Insights into nutritional quality and seed yield

Abstract

Growing chickpea (Cicer arietinum L.) faces significant challenges due to rising temperatures and drought stress, particularly during the reproductive and seed-filling phases. This study investigated the single and joint impacts of drought and heat stress on seed development, focusing on the responses of drought-tolerant (DT) and drought-sensitive (DS) chickpea genotypes. Initially raised in an outdoor environment (mean day and night temperature of 27 and 16±1 °C, respectively, light intensity of 1230–1440 µmol m⁻² s⁻¹, relative humidity of 70/43 %) until seed filling (around 110–113 days after sowing) commenced. The plants were subsequently exposed to single or combined heat and drought stress under controlled conditions until maturity. Control pots were maintained at day and night temperature of 25 and 15 °C, respectively with 500 µmol m⁻² s⁻¹ light, 60–65 % RH, and regular irrigation, and drought-stressed pots were kept at 50 % field capacity under the same conditions of light and humidity. Heat stress in pots was gradually increased to 32(day)/20 °C (night) under regular irrigation, while combined stress pots experienced both drought (50 % field capacity) and heat stress conditions 32(day)/20 °C (night) under the same light and humidity conditions with irrigation. All stress treatments adversely affected cell membranes, photosynthesis, and water regulation, with more pronounced effects under combined stress. While heat stress increased stomatal conductance, drought and combined stress significantly reduced it. Seed filling rate and duration decreased under all stress conditions, especially combined stress. The stresses in combination severely reduced seed weight and pod numbers compared to individual stresses. Enzyme activities involved in starch and sucrose synthesis and hydrolysis substantially decreased under the combined stress. Seed composition elements (starch, storage proteins, sugars, fat, crude fiber, and ash) exhibited significant reductions across all stress treatments, particularly for the combined stress. Thus, under combined stresses, starch, proteins, and soulube sugars were markedly decreased to 13–20 %, 6.4–12.4 %, and 3–5 % in seeds, compared to 37–39 %, 21–24 %, and 6 % in control seeds. The DT genotype outperformed the DS genotype for all traits under individual and combined stress conditions. Principal component analysis revealed a complex interplay among various physiological responses (membrane damage, chlorophyll, chlorophyll fluorescence, relative leaf water content, and stomatal conductance), seed yield, and seed composition under the combined stress. This study highlighted that combined heat and drought stress severely impacted chickpea yield and nutritional traits, such as seed starch and protein content, compared to individual stresses underscoring the need to develop cultivars tolerant to this stress combination.