1 INTRODUCTION

Terahertz (THz) wave radiation, located in the frequency range between microwaves and infrared light in the electromagnetic spectrum, has attracted much attention in recent years (Balakin et al., 2019). Compared to other electromagnetic waves, terahertz waves are noted for their unique penetrating and non-thermal effects and have become an emerging source of stress, especially in the field of plant physiological regulation. The special effect of terahertz waves stems from their frequency range, which allows them to penetrate many common substances, including biological tissues (Li et al., 2023). At the same time, the non-thermal effect of terahertz waves allows them to affect living organisms without causing significant temperature increases. This opens new possibilities for the use of terahertz waves in plant research as an innovative source of biological stimulation (Liu et al., 2023). Pinellia ternata, as a herbaceous plant, is subject to a wide range of environmental factors that affect its growth and development. Recent studies have shown that terahertz wave radiation exerts unique effects on plant physiological processes, including, but not limited to, the regulation of growth rate, cellular structure, gene expression, enzyme activities, etc (Lu et al., 2020). In P. ternata, a specific plant, terahertz waves may trigger unique physiological responses, such as growth promotion and enhancement of adversity resistance (Hang et al., 2023). A deeper understanding of this novel regulatory mechanism will provide important information for the development of plant physiology, agricultural production, and biotechnology.

A constantly expanding global population presents a formidable challenge for traditional Chinese medicine (TCM), which is to increase production to stimulate biological resources (Xu and Xia, 2019; Zhao et al., 2021b). Biologic and abiotic stressors are significant barriers to improving crop output in a sustainable manner (Zhang et al., 2022). Temperature, length of light, and humidity are three abiotic stressors that have an impact on plant growth (Ferrante and Mariani, 2018; Zhang et al., 2022). Underground roots and the microbiota that accompany them are examples of a multifunctional ecosystem (Zhuang et al., 2020). According to Zhang et al. (2022), plants must adapt to constantly changing conditions, including frequent stressful situations that are detrimental to their growth and development. According to Zhu (2016), these unfavourable conditions include biotic stressors like pathogen infections and herbivore predation as well as abiotic stresses including heat, cold, drought, nutrient deprivation, salinity, and toxic metals like cadmium, arsenic, and aluminium in the soil. The primary environmental conditions that impact plant distribution geographically, restrict crop yields, and jeopardise food security are salinity, drought, and temperature stress (Santos et al., 2022; Ikan et al., 2023). As demonstrated by Kuypers et al. (2018), roots are essential for the cycling of minerals and the transformation of organic molecules; symbiotic nitrogen fixation is one such example.

Apart from obtaining water and nutrients, roots participate in signalling amongst plants by secreting volatiles such as jasmonic acid (JA), brassinosteroids (BRs), and other substances (Zhu et al., 2023). Plant species optimise their root architecture in response to environmental conditions and requirements for the sections that are above ground (Khan et al., 2016; Garbowski et al., 2023). Coordinated endogenous genetic programming and interactions with external biotic and abiotic variables lead to optimised root architecture (Jung and McCouch, 2013). Water molecule arrangement alterations, plant root petiole lengthening, branching, and higher uptake of nutrients and water are all caused by THz waves (Zhao et al., 2020). These waves alter plant cells in addition to their morphology and physiology. This results in a cellular organisation that is more favourable to faster plant growth (Hoshina et al., 2016; Cherkasova et al., 2021). There is a group of plant hormones called BRs that control many aspects of plant growth and development. These include stomatal development, flowering, stress tolerance, photomorphogenesis, and seed germination (Nolan et al., 2020; Waadt et al., 2022). According to Sun et al. (2013) and Han et al. (2023), the membrane receptor kinase BRI detects the presence of BR and releases the inhibitor Brassinosteroid Insensitive (BRI). This activates the intracellular kinase structural domain of BRI and binds BRI to its co-receptor, BRI1-Associated Receptor Kinase (BAK). Following this, Peroxisome Proliferator-Activated (PPA) releases the transcription factors Brassinosteroid Enhanced Expression (BES) and Brassinazole Resistant (BZR) from Brassinosteroid-Insensitive 1-Interacting Receptor Kinase (BIN)-induced phosphorylation or dephosphorylates them, and BSU dephosphorylates and inactivates the negative regulator BIN in a multistep cascade of phosphorylation events (Wang et al., 2018; Li et al., 2020). Dephosphorylated BES and BZR have the ability to bind and control the expression of their target genes, which in turn triggers a BR response. Low BR levels cause BIN to become active, phosphorylate BZR and BES, and prevent them from binding to their target genes (Li et al., 2018).

More complicated side effects of salt stress and drought include oxidative stress, breakdown of cellular constituents such as proteins, lipids in membranes, and nucleic acids, as well as metabolic problems (Krasensky and Jonak, 2012; Cao et al., 2023). As a result, while primary stress signals trigger some cellular reactions, secondary signals trigger others (Galluzzi et al., 2018). There are separate and some shared signal transduction pathways between salt stress and drought stress. One significant characteristic of both salt and drought stress is that the plant hormone abscisic acid (ABA) can accumulate in response to osmotic stress signals, triggering adaptive responses in plants (Waadt et al., 2022).

Moreover, phytohormones such as ethylene, gibberellic acid (GA), ABA, salicylic acid, cytokinins, oleuropein steroids, and growth hormones control the THz-wave response (Trifunović-Momčilov et al., 2021). According to Neill et al. (2019), these plant growth regulators integrate endogenous signals and external inputs to modify plant defensive responses to heat stress. Additionally, plants can be made to tolerate heat using hormones applied to their roots or foliar sprays. Furthermore, plant hormones that confer tolerance to abiotic stimuli can be transported from roots to shoots (cytokinin, ABA, and growth hormone) or from shoots to roots (cytokinin, monogluconolactone, and ABA) (Wu et al., 2021). On the other hand, little is known about the long-distance movement of phytohormones under heat stress. Plants can withstand modest stress conditions by activating molecular pathways and exhibiting basal stress tolerance (Lacombe and Achard, 2016). It is interesting to note that they can also gain tolerance by resisting extreme stress levels by developing “molecular stress memory” from prior exposures. Different cellular responses can be reprogrammed in comparison to naive plants by brief exposure to non-destructive stressors or exogenously administered metabolites, chemicals, and microorganisms (Sharma et al., 2022). To lessen the consequences of heat stress, foliar or root, chemical and biological initiating events become significant.

For plants to adapt to environmental changes, they must achieve a new state of cellular homeostasis by carefully balancing a number of pathways that are located in various cellular compartments (Anderson and Song, 2020). However, under THz wave stress, this coordination could be upset, particularly if the plant as a whole or its cells are subjected to abrupt drops in water potential or if other environmental factors come into play (Zhao et al., 2020). Reactive oxygen species are created when electrons in high-energy states are transferred to molecular oxygen as a result of distinct pathways uncoupling. According to studies conducted with corresponding mutant lines of Arabidopsis, the mutants' stems displayed a slight rupture as a result of weakened intercellular adhesion. This suggests that the absence of BRs increases the binding of the plant's outer cells to its interior, which ultimately results in the dwarf mutation (Walsh et al., 2018). BRs functioned as “relaxants” for the surface cells, successfully alleviating the mechanical constraints on the inside cells, as demonstrated by cell wall reconstruction models. It has been demonstrated that THz increases BRI synthesis and causes mutations linked to BRIs. Consequently, it has been demonstrated that the BRs scavenging mechanism plays a significant role in shielding plants from osmotic stress as well as a combination of high light and temperature stress. High light levels can also cause cell damage under various abiotic stressors. Plant resistance may increase under high light-intensity stress, as well as the cellular production of BRs (Gyamerah et al., 2022).

Plants can act as stress signals to activate defence and adaptive mechanisms, thereby counteracting stress-related oxidative stress, in addition to actively producing BRs signals, such as NADPH oxidase, when metabolic imbalances occur during stressful times. (Mittler et al., 2022). The need to regulate the homeostatic level of BRs in cells during normal metabolism and in response to various stresses is highlighted by the two somewhat opposing “faces” of BRs, namely a beneficial signalling molecule on the one hand and a destructive toxic molecule on the other (D'Autréaux and Toledano, 2007). To help plants handle these environmental stresses better, we might be able to figure out how phytoalexin lactone signalling works when THz abiotic stressors happen.

In agricultural production applications, THz can promote the growth of cash crops, according to THz can change the arrangement of water molecules to affect plant nutrient absorption. We analyzed the possible interactions between P. ternata and THz through the development of P. ternata growth(Figure 1).

This review article's goal is to thoroughly investigate and clarify how terahertz radiation affects photosynthesis and stress tolerance, two vital functions of P. ternata. In order to better understand how THz radiation affects photosynthesis rates, chlorophyll content, and other relevant factors, this study will examine how it affects P. ternata. The article also aims to explore the ways in which THz radiation orchestrates and adjusts stress-induced reactive oxygen species (ROS) signalling in plant cells (Zheng et al., 2023). Through an analysis of these aspects, the review seeks to shed light on the complex mechanisms that underlie THz-induced stress responses in P. ternata. It will also go over the difficulties encountered in this field of study and suggest possible avenues for future research, all of which will come together to provide a comprehensive overview of what is now known about the effects of THz radiation on P. ternata photosynthesis, stress tolerance, and signalling (Wang et al., 2023).

2 IMPORTANCE OF TERAHERTZ WAVE COERCION ON PLANT RESPONSES IN THE CONTEXT OF TRADITIONAL CHINESE MEDICINE (TCM)

The traditional theory of Chinese medicine holds that there is a subtle and close association between plants and the human body, and their herbal applications occupy an important position in the field of Chinese medicine (Sun et al., 2018). Understanding the response of terahertz wave stress on plants is not only important in modern biology and agronomy but also carries far-reaching medical value in the context of TCM (Wang et al., 2023).

3 THz TECHNOLOGY IN AGRICULTURE, INDUSTRY, AND OTHER SCIENTIFIC FIELDS

THz technology, which refers to technology applied in the terahertz band (frequencies between approximately 0.1 and 10 THz), has found a variety of applications in agriculture, industry, and other scientific fields. Located between microwaves and infrared, terahertz waves exhibit unique properties in several ways, giving them the potential for a wide range of applications in different fields (Anitha et al., 2023).

THz technology finds diverse applications across various fields. In agriculture, it facilitates plant growth monitoring by assessing parameters like water content and biochemical changes, aiding in the optimisation of irrigation and fertilisation strategies (Afsah-Hejri et al., 2020). Additionally, its penetrating nature enables early detection of crop pests and diseases, enabling timely control measures. THz waves are also employed in seed quality assessment, evaluating moisture content and structural integrity. THz technology enables non-destructive testing of materials, quality control on production lines, and security inspections for non-metallic substances in industrial settings. In the realm of medical imaging, it plays a role in cancer detection and dermatological diagnostics due to its minimal impact on biological tissues and high contrast capabilities (Hlosta et al., 2022). In astronomy and atmospheric research, THz waves are utilised for observing galaxies, nebulae, and planetary atmospheres, as well as analysing atmospheric moisture and pollutants in the earth sciences. Furthermore, the high frequency of terahertz bands holds promise for future high-speed wireless communications technology in the field of communications technology (Gurvits et al., 2021).

4 OTHER CROP PLANT STRESS EFFECTS UNDER THREE-WAY INTERACTIONS

Studies on the response of grapes to salt stress described the physiological responses of grapes under different salt stress durations (0, 24, 48, and 72 hours) (Haider et al., 2019). It was found that grapes mitigate the negative effects of salinity by accumulating compounds such as glycine betaine (GB) and alginate. These compounds help to maintain higher potassium (K) levels, thereby reducing the potassium-to-sodium (K/Na) ratio. This study also observed that grapes were able to maintain better cellular water status and photosynthetic performance under salt stress. In addition, plants deal with oxidative stress by turning on the antioxidant scavenging system. This stopped the buildup of peroxides and lowered the production of reactive oxygen species (ROS) (Río Segade et al., 2017). Principal component analysis (PCA) was used to distinguish between different physiological and transcriptional responses, providing potential markers for salt stress tolerance studies in grapes (Jogaiah et al., 2013). Dandelion was found to exhibit higher total chlorophyll content, specific leaf area (SLA) (Rawat et al., 2021), nitrogen, and phosphorus content in leaves, as well as higher water use efficiency and nitrogen uptake following P fertiliser application (Haider et al., 2018). For shortleaf poplar, P fertiliser treatments reduced specific root length (SRL) and ectomycorrhizal infestation, as well as specific root tip density, but had no significant effect on mean root diameter (Zhu et al., 2023). The results suggest that the effects of soil nutrient availability and competition on plant functional traits are important for understanding plant adaptation during succession (Hortal et al., 2017).

The study of the response of different chloroplasts to light conditions was concerned with how two types of chloroplasts (M and BS) adjust their functions to different light conditions (Wietrzynski and Engel, 2021). Researchers found that BS chloroplasts can adjust to different light conditions by changing how much light they absorb and how it is distributed between the two photosystems (Fukuda et al., 2023). This was linked to a high rate of phosphorylation of LHCII proteins in BS-like vesicles. It was shown that different chloroplast types employ different mechanisms to optimise their functioning, which are associated with different light penetrations through the leaf and are essential for the regulation of chloroplast membrane flexibility and its functioning (Lu et al., 2021). In response to salt stress conditions, heightened transglutaminase (TGase) activity leads to a substantial increase in total endogenous polyamine levels within LHCII (Zhao et al., 2021). This upregulation suggests a crucial role for Putrescine (Put) in efficiently dissipating excess excitation energy by modulating the functionality of endogenous polyamines to influence LHCII adaptation (González-Hernández et al., 2022). These discoveries bear significant implications for comprehending plant responses to environmental stresses at the molecular and biochemical levels. The insights gained not only enhance our foundational knowledge of plant adaptation mechanisms but also hold practical significance for enhancing crop resilience and productivity (Xing et al., 2023).

5 EFFECT OF THz RADIATION ON THE GERMINATION OF PINELLIA TERNATA SEEDLINGS

One of the most important phases of plant growth is the germination of cluster seedlings (Reed et al., 2022). In this section, the impacts of THz radiation on the germination of P. ternata seedlings will be reviewed, and the processes underlying these effects will be examined. These mechanisms include the effects of THz radiation on the rate, speed, and vigour of the plant's bud tubers. After light signalling, SAUR17 is a crucial apical organ-specific dark morphogenetic effector. By competitively binding to the phosphatase PP2C-D1, SAUR17 prevents the inhibition of its phosphatase activity. Ultimately, this prevents the closure of dark-grown cotyledons and apical hooks by blocking the cell expansion mechanism (Alexandrov et al., 2013). In the apical region of seedlings grown in darkness, SAUR17 is strongly expressed, and its transcriptional level has been discovered to be modulated by ethylene and oleoresin lactone signals in addition to light signals. The transcriptional level of SAUR17 is found to be highly expressed in the apical portion of seedlings grown in darkness. It has been discovered that these signals, in addition to light signals, also regulate the transcriptional level of SAUR17. The transcriptional complex formed in vivo by PIFs, BZR1, and EIN3 is responsible for regulating the transcriptional level of SAUR17; additional analysis reveals that PIFs and EIN3/EIL1 enhance BZR1's ability to bind to the SAUR17 promoter and contribute to the transcriptional complex (Wang et al., 2020). Additional investigation showed that while the stability of PIF3 and EIN3 proteins was necessary for the upkeep of the oleuropein lactone pathway, PIFs and EIN3/EIL1 improved BZR1's capacity to bind to the SAUR17 promoter and triggered the transcriptional complex to activate SAUR17 expression (Zhao et al., 2021a). The accumulation of EIN3 and PIF3 proteins has been observed to be facilitated by oleuropein lactone's down-regulation of the transcript levels of the ubiquitin ligase genes EBF1 and EBF2 through BZR1. The fast emergence of seedlings was ultimately facilitated by the higher binding of the EIN3-PIFs-BZR1 transcriptional complex in promoter DNA and the stimulation of SAUR17 and HLS1 expression, which were in turn caused by the elevated amounts of EIN3 and PIF3 proteins (Badr et al., 2020). In summary, the mechanism by which THz influences the growth of P. ternata seedlings involves the involvement of EIN3 and PIFs (Figure 2).

6 THE EFFECT OF THz RADIATION ON THE GROWTH PROCESS OF PINELLIA TERNATA

Root development, stem and leaf development, and the production of flowers and fruits are all aspects of plant growth (David, 2017). The P. ternata root system, plant shoots, and petiole length tubers were all greatly impacted by THz waves. One crucial developmental stage in a plant's life cycle is the transition from juvenile to adult (or nutritional growth stage). Numerous significant physiological and biochemical processes, including plant architecture, the build-up of secondary metabolites, and the plant's reaction to biotic and abiotic stressors, are impacted by this developmental process (Fedorov and Tzortzakis, 2020).

Temporary immersion bio-reactor system (TIBS) culture is based on a liquid medium. It uses air pumps and air-filtered bacterial treatment membranes, which results in gas exchange between the reactor and the outside world and eventually cultivates a robust culture. Digital control devices are used to set the parameters of intermittent submergence, intermittent time, and time of submergence so as to keep the exoskeleton and the nutrient solution in intermittent contact. Reactor culture method advantages include: large inoculum quantity and a high degree of automation, which can lower production costs and enable high-throughput expansion of cash crops and endangered species; fast culture growth rate and stable secondary metabolism, which promote metabolite accumulation and the production of medicinally active substances; intermittent submerged culture, which regulates the reactor's gas exchange and plant-culture contact time, effectively controls the histoculture. The implementation of intermittent submerged culture has the potential to regulate the duration of contact between the plant and the culture medium, as well as facilitate the gas exchange process of P. ternata within the reactor. This approach can effectively regulate the vitrification status of the histocultured seedling and enable its robust growth. Additionally, during the reactor cultivation process, the culture medium composition can be altered, and bacteriostatic agents can be added, but the plant bioreactor lacks the ability to sample automatically and is unable to identify the materials present in the medium at any point (Fink et al., 2021). Through the study of Siraitia grosvenorii plant cultivation using plant bioreactor, proliferation rate, shoot length, fresh and dry weight Indications of proliferation rate, shoot length, fresh weight, and dry weight showed that the total biomass of roselle cultivated in plant bioreactor was significantly higher than in solid and liquid cultivation (Shivani et al., 2021). The reactor promoted the growth and quality of S. grosvenorii seedlings, and the natural-like plant S. grosvenorii obtained may have a positive effect on isolated rooting and transplanting in large-scale commercial production (Xie et al., 2023). Some researchers evaluated Vanilla planifolia in solid culture, partial submergence and plant bioreactor for shoot proliferation efficiency and found that compared to solid culture, the proliferation rate in the reactor increased by a factor of three (Yeh et al., 2021). Intermittent submerged bioreactors combine the advantages of solid and liquid cultures by reducing humidity in the vessel through increased gas exchange, enhancing the normal metabolism of plant tissues in relation to their surroundings, and preventing vitrification (Zhang et al., 2018). At the same time, plant bioreactor systems allow for intermittent and brief contact between the explants and the liquid medium to ensure a proper supply of nutrients. The plant bioreactor provides better control of culture conditions, optimal supply of nutrients and growth regulators, and the ability to control the growth of the plant according to its condition (Uma et al., 2021). substances and growth regulators, changing media composition during culture according to the developmental stage, filtering media to control secretions, and controlling contamination. We placed THz energy rings in TIBS to co-cultivate plants, and through the effect of THz waves on the liquid medium, the roots of the plants absorbed the THz-treated medium, thus indirectly promoting plant growth (Fedorov and Tzortzakis, 2020). P. ternata seedlings can grow and reach maturity with the help of THz waves in TIBs(Figure 3).

7 COORDINATION OF STRESS-INDUCED ROS SIGNALLING BY THz IN PLANT CELLS

In addition to their more well-known roles in cytoprotection, ROS-scavenging enzymes have been demonstrated in recent years to be involved in signalling (Mittler et al., 2022). When exposed to moderate levels of light stress, Arabidopsis plants lacking cytoplasmic APX1 stimulated the expression of numerous stress-responsive genes and had greater H levels and H₂O₂ than wild-type plants (Koussevitzky et al., 2008; Li et al., 2019b; Zhang et al., 2022). It has been demonstrated that knockout APX1 plants develop more rapidly than wild-type plants under high salinity or osmotic environments. The apx1 plants were more susceptible to photooxidation and oxidative stress caused by paraquat, which is why these results were unexpected (Suzuki et al., 2016; Saxena et al., 2020). Similar to this, decreased tylAPX expression in Arabidopsis enhanced resistance to salt and osmotic stressors but had no effect on growth in the presence of oxidative stress (Miller et al., 2010; Belgaroui et al., 2018). In contrast, the loss of both genes in the double mutant apx1/tylapx preserved salt tolerance but enhanced sensitivity to sorbitol treatment. Analogously, antisense APX1 and antisense CAT1 in tobacco were continuously exposed to oxidative stress; however, the double antisense line grew increasingly resilient (Miller et al., 2007; Pabuayon et al., 2021).

The overexpression of Zat1 in Apx7 knockout plants led to considerable tolerance to cold stress and salinity, as well as increased expression of defence transcripts such as WRKY70, AOX1, NHX1, and Cor78. On the other hand, Zat7 RNAi lines displayed decreased resistance to osmotic stress (Yu et al., 2016; Marowa et al., 2020). Zat7 OE plants showed heightened sensitivity to sorbitol-induced hyperosmotic conditions, in contrast to their increased tolerance to salt. It has recently been demonstrated that MKK9 activates MPK3 and MPK6, two ROS-responsive MAP kinases triggered by H (Wani et al., 2020; Hussain et al., 2021). On the other hand, the MKK9 mutant was more resistant to salt stress, while transgenic Arabidopsis expressing constitutively active MKK9 displayed enhanced activation of endogenous MPK3 and MPK6. It's interesting to note that MKK1 and MKK9 activated MPK6 in response to salt exposure; however, plants that overexpressed MKK1 showed more tolerance to salt stress compared to wild-type plants, whereas transgenic plants with MKK9 showed greater sensitivity (Luo et al., 2017; Kumar et al., 2020).

Proteins with NAC structural domains are plant-specific transcription factors involved in a variety of responses, including development, hormonal signalling, and abiotic stress responses like cold, dehydration, and salinity. These domains may be used to further explore the relationship between THz-regulated ROS and salinity or drought stress. According to Petutschnig et al. (2010), CERK1 is a receptor-like kinase that is necessary for chitin-evoked signalling in Arabidopsis.

ROS functions as signalling molecules that control growth and development in addition to stress reactions (Mittler et al., 2022). In addition to being viewed as a negative event that should be avoided or minimised, oxidative stress brought on by drought and salt stress can also be perceived as essential for plants to adapt appropriately and initiate the proper domestication mechanisms (Krasensky & Jonak, 2012). According to Rosa et al. (2010), cytosolic APX1 or tylAPX deficiency in Arabidopsis causes ROS buildup, which in turn produces signals that improve plant tolerance to salt stress and osmotic stress. Moreover, Li et al. (2021a) reported that salt-induced ROS buildup in AtVAMP7C antisense plant endosomes produced cytosolic signals that improved salt tolerance. These studies demonstrate how plants experience enhanced ROS generation in various tissues and subcellular compartments in response to decreased water availability and/or increased salt under light circumstances. Many evanescent ROS signals may be produced in this situation, necessitating a high level of coordination and control on the part of the plant cell (Jiao et al., 2021).

It has been demonstrated that ROS signals from various organelles cause significant transcriptional alterations and cellular reprogramming, which can either protect plant cells or cause plant cells to die according to a set schedule (Cheung and Vousden, 2022). These kinds of reprogramming imply that ROS signalling is mediated by at least partially involved organelle retrograde communication, which enables the ROS-producing organelle and the nucleus—and possibly the individual organelles—to coordinate the stress response. Retrograde signals can be divided into two primary groups: Two key mechanisms are (1) developmental control over the biogenesis of organelles and (2) operational control, which involves quick adaptation to environmental and developmental cues. It is believed that ROS, which are created by these organelles, are significant signalling molecules that are engaged in retrograde signalling during stressful situations(Figure 4).

8 CHALLENGES AND FUTURE DIRECTIONS

Addressing the methodological challenges in the study of terahertz (THz) wave stresses on P. ternata is paramount to advancing this research field. Challenges encompass limited access to THz sources and the need for precise characterisation; difficulties in sample preparation and beam uniformity; sensitivity and complexity in data acquisition and analysis; inherent biological variability; ethical considerations regarding plant health; the integration of multi-omics data; conducting long-term studies; standardising protocols; fostering interdisciplinary collaboration; and securing adequate funding and resources. Tackling these challenges collectively will be essential to pave the way for comprehensive investigations into the response surfaces and mechanisms of THz wave stresses on P. ternata, ultimately benefiting both plant science and THz technology applications.

In the study of THz wave stresses on P. ternata, several unresolved questions persist, necessitating further investigation. Key among these are the precise molecular mechanisms underpinning the plant's response to THz radiation, including the role of specific genes, proteins, and signalling pathways. Additionally, understanding the long-term effects and potential ecological implications of THz exposure on P. ternata and surrounding ecosystems remains an open question. The extent to which THz-induced stress alters the plant's secondary metabolite production and how this may impact its medicinal properties requires deeper exploration. Moreover, the establishment of standardised methodologies and protocols for THz-plant interaction studies is crucial to enabling robust comparisons between different research efforts. Lastly, as THz technology advances, researchers must address the ethical considerations surrounding the use of THz radiation on plants and ecosystems. Resolving these unanswered questions will enhance our comprehension of THz-plant interactions and their broader implications for agriculture and biotechnology.

Exploring future research avenues in the study of THz wave stresses on P. ternata presents exciting prospects. To advance this field, researchers can delve deeper into the development of non-invasive and high-throughput THz imaging techniques to monitor plant responses over time precisely. Investigating the potential for THz radiation to modulate specific metabolic pathways and enhance the synthesis of valuable secondary metabolites in P. ternata holds promise for medicinal plant cultivation. Additionally, interdisciplinary collaborations between botanists, physicists, and engineers can lead to innovative THz technologies tailored for plant research. Harnessing artificial intelligence and machine learning algorithms for data analysis and predictive modelling may further elucidate complex THz-plant interactions. Long-term ecological studies can shed light on the sustainability and environmental impacts of THz technology in agriculture. Overall, future research avenues should focus on integrating cutting-edge technology, bioinformatics, and ecological considerations to unlock the full potential of THz wave stresses in optimising P. ternata and other plant species for various applications, from medicine to agriculture.

9 CONCLUSION

In conclusion, this review has delved into the multifaceted effects and intricate mechanisms governing THz wave stress on P. ternata, offering valuable insights into the plant's responses to this emerging stressor. Our exploration has underscored the significance of future research, which should prioritise a systemic examination of THz wave impacts, spanning from shoots to roots, and delve into the molecular intricacies that define the structure and function of the root system. We have highlighted how THz waves, in synergy with other environmental stressors, act as accelerators of plant growth while inducing alterations in cellular homeostasis, ultimately leading to the generation of reactive oxygen species (ROS) and augmentation in oleoresin lactone production. Notably, brassinosteroids have emerged as pivotal players in P.ternata response to abiotic stresses, serving both as toxic by-products of stress metabolism and as essential signal transduction molecules. This review has also shed light on the spatio-temporal regulation and long-distance transport of phytohormones, encompassing growth hormone, cytokinin, and abscisic acid (ABA), as pivotal determinants of growth and development in P. ternata under THz wave stress conditions. The examination of ROS and brassinosteroid insensitive (BRI) homeostasis and signalling under THz wave stress has expanded our understanding of BRI's role in stress perception, stress signalling, and the regulation of domestication responses. Additionally, we have highlighted the importance of considering the spatio-temporal regulation and long-distance transport of key plant hormones, including growth hormone, cytokinin, and ABA, in shaping root growth and development in response to THz stress. As we conclude, it is evident that deciphering how plants perceive and respond to environmental stresses is a fundamental biological question. Furthermore, enhancing plant stress tolerance is not only pivotal for bolstering agricultural production but is also indispensable for fostering environmental sustainability, particularly as low-tolerance crops consume excess water and fertilisers, thereby exacerbating the environmental burden. Considering these findings, future research endeavours should continue to unveil the intricate facets of THz wave stress on P. ternata, thereby contributing to our scientific understanding and promoting sustainable agricultural practices.

FUNDING INFORMATION

This work was supported by Jiangsu Synergetic Innovation Center for Advanced Bio-Manufacture (Grant Number: XTD1825). National Natural Science Foundation of China (82373981).