Abiotic stressors, including drought, salt, cold, and heat, profoundly impact plant growth and development, forcing elaborate cellular responses for adaptation and resilience. Among the crucial orchestrators of these responses is the CBL-CIPK pathway, comprising calcineurin B-like proteins (CBLs) and CBL-interacting protein kinases (CIPKs). While CIPKs act as serine/threonine protein kinases, transmitting calcium signals, CBLs function as calcium sensors, influencing the plant’s response to abiotic stress. This review explores the intricate interactions between the CBL-CIPK pathway and plant hormones such as ABA, auxin, ethylene, and jasmonic acid (JA). It highlights their role in fine-tuning stress responses for optimal survival and acclimatization. Building on previous studies that demonstrated the enhanced stress tolerance achieved by upregulating CBL and CIPK genes, we explore the regulatory mechanisms involving post-translational modifications and protein–protein interactions. Despite significant contributions from prior research, gaps persist in understanding the nuanced interplay between the CBL-CIPK system and plant hormone signaling under diverse abiotic stress conditions. In contrast to broader perspectives, our review focuses on the interaction of the pathway with crucial plant hormones and its implications for genetic engineering interventions to enhance crop stress resilience. This specialized perspective aims to contribute novel insights to advance our understanding of the potential of the CBL-CIPK pathway to mitigate crops’ abiotic stress.

Salvia sclarea L. is a Cd²⁺ tolerant medicinal herb with antifungal and antimicrobial properties cultivated for its pharmacological properties. However, accumulation of high Cd²⁺ content in its tissues increases the adverse health effects of Cd²⁺ in humans. Therefore, there is a serious demand to lower human Cd²⁺ intake. The purpose of our study was to evaluate the mitigative role of excess Zn²⁺ supply to Cd²⁺ uptake/translocation and toxicity in clary sage. Salvia plants were treated with excess Cd²⁺ (100 μM CdSO₄) alone, and in combination with Zn²⁺ (900 μM ZnSO₄), in modified Hoagland nutrient solution. The results demonstrate that S. sclarea plants exposed to Cd²⁺ toxicity accumulated a significant amount of Cd²⁺ in their tissues, with higher concentrations in roots than in leaves. Cadmium exposure enhanced total Zn²⁺ uptake but also decreased its translocation to leaves. The accumulated Cd²⁺ led to a substantial decrease in photosystem II (PSII) photochemistry and disrupted the chloroplast ultrastructure, which coincided with an increased lipid peroxidation. Zinc application decreased Cd²⁺ uptake and translocation to leaves, while it mitigated oxidative stress, restoring chloroplast ultrastructure. Excess Zn²⁺ ameliorated the adverse effects of Cd²⁺ on PSII photochemistry, increasing the fraction of energy used for photochemistry (Φ_PSII) and restoring PSII redox state and maximum PSII efficiency (Fv/Fm), while decreasing excess excitation energy at PSII (EXC). We conclude that excess Zn²⁺ application eliminated the adverse effects of Cd²⁺ toxicity, reducing Cd²⁺ uptake and translocation and restoring chloroplast ultrastructure and PSII photochemical efficiency. Thus, excess Zn²⁺ application can be used as an important method for low Cd²⁺-accumulating crops, limiting Cd²⁺ entry into the food chain.

Salicylic acid (SA), an essential plant hormone, has received much attention due to its role in modulating the adverse effects of biotic and abiotic stresses, acting as an antioxidant and plant growth regulator. However, its role in photosynthesis under non stress conditions is controversial. By chlorophyll fluorescence imaging analysis, we evaluated the consequences of foliar applied 1 mM SA on photosystem II (PSII) efficiency of tomato (Solanum lycopersicum L.) plants and estimated the reactive oxygen species (ROS) generation. Tomato leaves sprayed with 1 mM SA displayed lower chlorophyll content, but the absorbed light energy was preferentially converted into photochemical energy rather than dissipated as thermal energy by non-photochemical quenching (NPQ), indicating photoprotective effects provided by the foliar applied SA. This decreased NPQ, after 72 h treatment by 1 mM SA, resulted in an increased electron transport rate (ETR). The molecular mechanism by which the absorbed light energy was more efficiently directed to photochemistry in the SA treated leaves was the increased fraction of the open PSII reaction centers (qp), and the increased efficiency of open reaction centers (Fv’/Fm’). SA induced a decrease in chlorophyll content, resulting in a decrease in non-regulated energy dissipated in PSII (Φ_NO) under high light (HL) treatment, suggesting a lower amount of triplet excited state chlorophyll (³Chl*) molecules available to produce singlet oxygen (¹O₂). Yet, the increased efficiency, compared to the control, of the oxygen evolving complex (OEC) on the donor side of PSII, associated with lower formation of hydrogen peroxide (H₂O₂), also contributed to less creation of ROS. We conclude that under non stress conditions, foliar applied SA decreased chlorophyll content and suppressed phototoxicity, offering PSII photoprotection; thus, it can be regarded as a mechanism that reduces photoinhibition and photodamage, improving PSII efficiency in crop plants.

The effects of environmentally relevant bisphenol A (BPA) concentrations (0.3, 1 and 3 μg L⁻¹) were tested at 2, 4, 6 and 8 days, on intermediate leaves, of the seagrass Cymodocea nodosa. Hydrogen peroxide (H₂O₂) production, lipid peroxidation, protein, phenolic content and antioxidant enzyme activities were investigated. Increased H₂O₂ formation was detected even at the lowest BPA treatments from the beginning of the experiment and both the enzymatic and non-enzymatic antioxidant defense mechanisms were activated upon application of BPA. Elevated H₂O₂ levels that were detected as a response to increasing BPA concentrations and incubation time, led to the decrease of protein content on the 4th day even at the two lower BPA concentrations, and to the increase of the lipid peroxidation at the highest concentration. However, on the 6th day of BPA exposure, protein content did not differ from the control, indicating the ability of both the enzymatic and non-enzymatic mechanisms (such as superoxide dismutase (SOD) and phenolics) to counteract the BPA-derived oxidative stress. The early response of the protein content determined that the Low Effect Concentration (LOEC) of BPA is 0.3 μg L⁻¹ and that the protein content meets the requirements to be considered as a possible early warning “biomarker” for C. nodosa against BPA toxicity.

Five-day exposure of clary sage (Salvia sclarea L.) to 100 μM cadmium (Cd) in hydroponics was sufficient to increase Cd concentrations significantly in roots and aboveground parts and affect negatively whole plant levels of calcium (Ca) and magnesium (Mg), since Cd competes for Ca channels, while reduced Mg concentrations are associated with increased Cd tolerance. Total zinc (Zn), copper (Cu), and iron (Fe) uptake increased but their translocation to the aboveground parts decreased. Despite the substantial levels of Cd in leaves, without any observed defects on chloroplast ultrastructure, an enhanced photosystem II (PSII) efficiency was observed, with a higher fraction of absorbed light energy to be directed to photochemistry (Φ_PSΙΙ). The concomitant increase in the photoprotective mechanism of non-photochemical quenching of photosynthesis (NPQ) resulted in an important decrease in the dissipated non-regulated energy (Φ_NO), modifying the homeostasis of reactive oxygen species (ROS), through a decreased singlet oxygen (¹O₂) formation. A basal ROS level was detected in control plant leaves for optimal growth, while a low increased level of ROS under 5 days Cd exposure seemed to be beneficial for triggering defense responses, and a high level of ROS out of the boundaries (8 days Cd exposure), was harmful to plants. Thus, when clary sage was exposed to Cd for a short period, tolerance mechanisms were triggered. However, exposure to a combination of Cd and high light or to Cd alone (8 days) resulted in an inhibition of PSII functionality, indicating Cd toxicity. Thus, the rapid activation of PSII functionality at short time exposure and the inhibition at longer duration suggests a hormetic response and describes these effects in terms of “adaptive response” and “toxicity”, respectively.

In recent years, due to the development of industrial and agricultural production, heavy metal contamination has attracted increasing attention. Aromatic and medicinal plant Salvia sclarea L. (clary sage) is classified to zinc (Zn) accumulators and considered as a potential plant for the phytoremediation of heavy metal polluted soils. In this study, an adaptation of clary sage to 900 µM (excess) Zn exposure for eight days in a hydroponic culture was investigated. The tolerance mechanisms under excess Zn exposure were assessed by evaluating changes in the nutrient uptake, leaf pigment and phenolic content, photosynthetic activity and leaf structural characteristics. The uptake and the distribution of Zn, as well as some essential elements such as: Ca, Mg, Fe, Mn and Cu, were examined by inductively coupled plasma mass spectrometry. The results revealed that Salvia sclarea is a Zn-accumulator plant that tolerates significantly high toxic levels of Zn in the leaves by increasing the leaf contents of Fe, Ca and Mn ions to protect the photosynthetic function and to stimulate the photosystem I (PSI) and photosystem II (PSII) activities. The exposure of clary sage to excess Zn significantly increased the synthesis of total phenolics and anthocyanins in the leaves; these play an important role in Zn detoxification and protection against oxidative stress. The lipid peroxidation and electrolyte leakage in leaves, used as clear indicators for heavy metal damage, were slightly increased. All these data highlight that Salvia sclarea is an economically interesting plant for the phytoextraction and/or phytostabilization of Zn-contaminated soils.