High temperatures and decreased rainfall are detrimental to yield in chickpea (Cicer arietinum L.), particularly during grain filling. This study aimed to (i) assess the individual and combined effects of drought and heat stress on biochemical seed-filling processes, (ii) determine genotypic differences in heat and drought tolerance, and (iii) determine any cross-tolerance. Plants were grown outdoors in the normal growing season when temperatures during seed filling were <32−20°C or were planted late (temperatures >32−20°C; heat stress). Half of the pots were kept adequately watered throughout, but water was withheld from the others from the initiation of seed filling until the relative leaf water content reached 50% of the irrigated plants (drought stress); all plants were rewatered thereafter until seed maturit. Water was withheld for 13 days (normal sowing) and 7 days (late sowing), so soil moisture decreased by 54–57%. Tests on leaves and seeds were performed after the stress. Individual and combined stress damaged membranes, and decreased cellular oxidising ability, stomatal conductance, PSII function and leaf chlorophyll content; damage was greater under combined stress. Leaf Rubisco activity increased with heat stress, decreased with drought stress and decreased severely with combined stress. Sucrose and starch concentrations decreased in all seeds through reductions in biosynthetic enzymes; reductions were greater under combined stress. These effects were more severe in heat- and drought-sensitive genotypes compared with drought-tolerant genotypes. Drought stress had a greater effect than heat stress on yield and the biochemical seed-filling mechanisms. Drought- and heat-tolerant genotypes showed partial cross-tolerance.
Chickpea (Cicer arietinum L.), in its reproductive stage, is sensitive to heat stress (32/20°C or higher as day/night temperatures) with consequent substantial loss of potential yields at high temperatures. The physiological mechanisms associated with reproductive failures have not been established: they constitute the basis of this study. Here, we initially screened a large core-collection of chickpea against heat stress and identified two heat-tolerant (ICC15614, ICCV. 92944) and two heat-sensitive (ICC10685, ICC5912) genotypes. These four genotypes were sown during the normal time of sowing (November–March) and also late (February–April) to expose them to heat stress during reproductive stage (>32/20°C). The genotypes were assessed for damage by heat stress to the leaves and reproductive organs using various indicators of stress injury and reproductive function. In the heat-stressed plants, phenology accelerated as days to flowering and podding, and biomass decreased significantly. The significant reduction in pod set (%) was associated with reduced pollen viability, pollen load, pollen germination (in vivo and in vitro) and stigma receptivity in all four genotypes. Heat stress inhibited pollen function more in the sensitive genotypes than in the tolerant ones, and consequently showed significantly less pod set. Heat stress significantly reduced stomatal conductance, leaf water content, chlorophyll, membrane integrity and photochemical efficiency with a larger effect on heat-sensitive genotypes. Rubisco (carbon-fixing enzyme) along with sucrose phosphate synthase (SPS) and sucrose synthase (SS) (sucrose-synthesising enzymes) decreased significantly in leaves due to heat stress leading to reduced sucrose content. Invertase, a sucrose-cleaving enzyme, was also inhibited along with SPS and SS. The inhibition of these enzymes was significantly greater in the heat-sensitive genotypes. Concurrently, the anthers of these genotypes had significantly less SPS and SS activity and thus, sucrose content. As a result, pollen had considerably lower sucrose levels, resulting in reduced pollen function, impaired fertilisation and poor pod set in heat-sensitive genotypes.
Plants are exposed to a wide range of environmental conditions and one of the major forces that shape the structure and function of plants are temperature stresses, which include low and high temperature stresses and considered as major abiotic stresses for crop plants. Due to global climate change, temperature stress is becoming the major area of concern for the researchers worldwide. The reactions of plants to these stresses are complex and have devastating effects on plant metabolism, disrupting cellular homeostasis and uncoupling major physiological and biochemical processes. Temperature stresses disrupt photosynthesis and increase photorespiration thereby altering the normal homeostasis of plant cells. The constancy of temperature, among different metabolic equilibria present in plant cells, depends to a certain extent on a homeostatically regulated ratio of redox components, which are present virtually in all plant cells. Several pathways, which are present in plant cells, enable correct equilibrium of the plant cellular redox state and balance fluctuations in plant cells caused by changes in environment due to stressful conditions. In temperature stresses, high temperature stress is considered to be one of the major abiotic stresses for restricting crop production worldwide. The responses of plants to heat stress vary with extent of temperature increase, its duration and the type of plant. On other hand, low temperature as major environmental factor often affects plant growth and crop productivity and leads to substantial crop loses. A direct result of stress-induced cellular changes is overproduction of reactive oxygen species (ROS) in plants which are produced in such a way that they are confined to a small area and also in specific pattern in biological responses. ROS (superoxide; O − 2 , hydroxyl radicals; OH − , alkoxyl radicals and non-radicals like hydrogen peroxide; H 2 O 2 and singlet oxygen; 1 O 2) are highly reactive and toxic and cause damage to proteins, lipids, carbohydrates which ultimately results in oxidative stress. ROS may also serve as signaling molecules in mediating important signal transduction pathways that coordinate an astonishing range of diverse plant processes under temperature stress. To counter temperature induced oxidative stress, plants upregulate a variety of enzymatic and non-enzymatic antioxidants which are also information-rich redox buffers and important components of redox signaling that interact with biomembrane-related compartments. They provide essential information on cellular redox state, and regulate gene expression associated with stress responses to optimize defense and survival, stress acclimation and tolerance. The work done by various researchers has explored a direct link between ROS scavenging and plant tolerance under temperature extremes in various crops which include legumes, cereals, oil crops and vegetables. There is ample need to develop temperature tolerance in crop plants by exploring suitable strategies to manage oxidative stress and maintain cel...
Drought and heat stress are two major constraints that limit chickpea (Cicer arietinum L.) yield, particularly during seed filling. The present study aimed (i) to assess the individual and combined effects of drought and heat stress on oxidative metabolism during seed filling, and (ii) to determine any genetic variation in oxidative metabolism among genotypes differing in drought and heat tolerance and sensitivity. The plants were raised in outdoor conditions with two different times of sowing, one in November (normal-sown, temperatures <32°C−20°C (day–night) during seed filling), and the other in February (late-sown, temperatures >32°C−20°C (day–night) during seed filling). Plants were regularly irrigated to prevent any water shortage until the water treatments were applied. At both sowing times, the drought treatment was applied during seed filling (at ~75% podding) by withholding water from half of the pots until the relative leaf water content (RLWC) of leaves on the top three branches reached 42–45%, whereas leaves in the fully irrigated control plants were maintained at RLWC 85–90%. Drought-stressed plants were then rewatered and maintained under fully irrigated conditions until maturity. Several biochemical parameters were measured on the leaves and seeds at the end of the stress treatments, and seed yield and aboveground biomass were measured at maturity. Individual and combined stresses damaged membranes, and decreased PSII function and leaf chlorophyll content, more so under the combined stress treatment. The levels of oxidative molecules (malondialdehyde (MDA) and H2O2) markedly increased compared with the control plants in all stress treatments, especially across genotypes in the combined heat + drought stress treatment (increases in leaves: MDA 5.4–8.4-fold and H2O2 5.1–7.1-fold; in seeds: MDA 1.9–3.3-fold and H2O2 3.8–7.9-fold). The enzymatic and non-enzymatic antioxidants related to oxidative metabolism increased under individual stress treatments but decreased in the combined heat + drought stress treatment. Leaves had higher oxidative damage than seeds, and this likely inhibited their photosynthetic efficiency. Yields were reduced more by drought stress than by heat stress, with the lowest yields in the combined heat + drought stress treatment. Heat- and drought-tolerant genotypes suffered less damage and had higher yields than the heat- and drought-sensitive genotypes under the individual and combined stress treatments, suggesting partial cross-tolerance in these genotypes. A drought-tolerant genotype ICC8950 produced more seed yield under the combined heat + drought stress than other genotypes, and this was associated with low oxidative damage in leaves and seeds.
Green house study was aimed to investigate the effect of seed biopriming with drought tolerant isolates of Trichoderma harzianum, viz. Th 56, 69, 75, 82 and 89 on growth of wheat under drought stress and to explore the mechanism underlying plant water stress resilience in response to Trichoderma inoculation. Measurements of relative water content, osmotic potential, osmotic adjustment, leaf gas exchange, chlorophyll fluorescence and membrane stability index were performed. In addition, analysis of the phenolics, proline, lipid peroxidation and measurements of phenylalanine ammonia-lyase activity were carried out. Seed biopriming enhanced drought tolerance of wheat as drought induced changes like stomatal conductance, net photosynthesis and chlorophyll fluorescence were delayed. Drought stress from 4 to 13 days of withholding water induced an increase in the concentration of stress induced metabolites in leaves, while Trichoderma colonisation caused decrease in proline, malondialdehyde (MDA) and hydrogen peroxide (H 2 O 2 ), and an increase in total phenolics. A common factor that negatively affects plants under drought stress conditions is accumulation of toxic reactive oxygen species (ROS), and we tested the hypothesis that seed biopriming reduced damages resulting from accumulation of ROS in stressed plants. The enhanced redox state of colonised plants could be explained by higher L-phenylalanine ammonia-lyase (PAL) activity in leaves after 13 days of drought stress in Trichoderma treated plants. Similar activity was induced in untreated plants in response to drought stress but to a lower extent in comparison to treated plants. Our results support the hypothesis that seed biopriming in wheat with drought tolerant T. harzianum strains increased root vigour besides performing the process of osmoregulation. It ameliorates drought stress by inducing physiological protection in plants against oxidative damage, due to enhanced capacity to scavenge ROS and increased level of PAL, a mechanism that is expected to augment tolerance to abiotic stresses.
Components of disease reaction, including incubation period, pustule types, inoculum production and disease index (DI); and contents of protein, phenols, soluble sugars and reducing and non‐reducing sugars were investigated in cotyledonary and true leaves of six genotypes of Brassica juncea: Varuna, Kranti, EC‐399296, EC‐399299, EC‐399313 and EC‐399301, inoculated with Albugo candida. Cotyledonary leaves were examined 14 days after inoculation (d.a.i.), whereas true leaves were scored 14 and 21 d.a.i. Disease indices were assessed on a 0% (resistant) to 100% (susceptible) scale. DIs at the cotyledonary leaf stage in the above six genotypes were 67, 65, 32, 31, 31 and 38%, respectively, whereas at the true‐leaf stage they were 21, 28, 12, 17, 9 and 4%, respectively at 14 d.a.i., and 35, 45, 17, 19, 20 and 6%, respectively at 21 d.a.i. Protein contents were highest in the genotypes with the highest DIs, such as Varuna at the cotyledonary leaf stage and Kranti at the true‐leaf stage, and lowest in the genotypes with the lowest DIs, such as EC‐399299 at the cotyledonary stage and EC‐399301 at the true‐leaf stage. Total phenols, total sugars, reducing sugars and non‐reducing sugars were generally negatively correlated with DI, but were not always consistent, particularly when differences in DI were small. The results indicated that factors conditioning the response of host genotypes to A. candida may differ or operate in different ways at different growth stages.
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