In collaboration with Scientific Association of Iranian Medicinal Plants

Document Type : Reviewed scientific papers

Authors

1 Associate Professor, Department of Agriculture Payame Noor University

2 Instructor, Department of Agriculture, Payame Noor University, Tehran, Iran

10.22092/ijmapr.2026.372415.3620

Abstract

     Background and Objectives: Medicinal plants are important sources of bioactive compounds used in pharmaceutical, food, and health industries, but the quantity and quality of their active constituents are strongly affected by environmental conditions. Recent evidence indicates that abiotic stress does not merely restrict plant growth; they also reprogram secondary metabolism through physiological and molecular regulatory networks. Focusing on drought, salinity, temperature, and radiation, this review examines the roles of redox regulation, hormonal crosstalk, transcriptional control, and biosynthetic pathways in shaping secondary-metabolite production. The main objective was to clarify the relationships among stress intensity, physiological responses, metabolic reprogramming, and the final quality of medicinal compounds.
Methodology: This study was conducted as an analytical review. Scientific literature was retrieved from Web of Science, Scopus, and Google Scholar using combinations of English keywords related to medicinal plants, abiotic stress, secondary metabolism, redox regulation, hormonal signaling, and biosynthetic pathways. The review mainly focused on peer-reviewed articles published from 2018 to 2024 that investigated relationships among abiotic stresses, physiological and molecular responses, and secondary-metabolite production in medicinal plants. Studies were selected when they were directly relevant to the review topic and provided physiological, biochemical, genomic, or metabolomic evidence useful for explaining the regulatory mechanisms of secondary metabolism. Purely descriptive studies or papers without a direct link to the quality of bioactive compounds were excluded. The selected findings were classified and compared according to stress type, response intensity, redox status, hormonal interactions, transcriptional control, biosynthetic-pathway regulation, and physiological and management implications. Evidence from different species and metabolite groups was then integrated to identify common response patterns as well as differences related to stress type, plant species, and response intensity. Finally, the relationships among redox signaling, hormonal regulation, gene expression, carbon allocation, and the biosynthesis of phenolic, terpenoid, and alkaloid compounds were synthesized into a conceptual framework linking environmental stress, plant physiological status, metabolic reprogramming, and active-compound quality.
Results: The reviewed evidence showed that secondary metabolism in medicinal plants is a dynamic system closely linked to plant physiological status. Drought, salinity, temperature, and radiation alter redox balance and, through controlled increases in reactive oxygen species, activate cellular signaling pathways and regulate the expression of key biosynthetic genes. These responses are strengthened by interactions among hormones such as jasmonic acid, salicylic acid, and abscisic acid and may redirect carbon flux toward the production of phenolic, terpenoid, and alkaloid compounds. Metabolic responses to stress were clearly nonlinear: mild stress often increased active-compound concentrations, whereas severe stress reduced metabolite-production capacity by limiting photosynthesis, energy supply, and carbon precursors. Morphological adjustments, including an increased glandular trichome density, may also enhance the capacity to produce certain medicinal compounds. Across studies, medicinal quality emerged as the result of interactions among photosynthetic status, resource allocation, redox and hormonal networks, and genetic capacity; therefore, similar stress intensity did not necessarily produce the same response across species or developmental stages. An increase in individual metabolite concentration did not necessarily translate into a higher total yield of the active compound because it could coincide with reduced biomass. Overall, controlled stress, appropriate timing of agronomic practices, and elicitor application were most effective when metabolic stimulation occurred without a severe reduction in growth and biomass.
Conclusion: The effects of environmental stress on medicinal-compound production depend on stress type and intensity, genotype, developmental stage, and photosynthetic capacity. Mild stress can enhance secondary-metabolite accumulation through redox signaling, hormonal pathways, and activation of biosynthetic genes, whereas severe stress can have the opposite effect by restricting photosynthesis and carbon supply. Therefore, active-compound concentration should be assessed alongside biomass and total metabolite yield. Determining the appropriate stress threshold and timing can serve as a scientific basis for the targeted use of controlled stress and elicitors to improve medicinal-plant quality.

Keywords

Main Subjects

- Abbasi, S., Houshmand, S. and Ahmadi, T., 2024. Volatile compositions and glandular trichomes of Zataria multiflora in different phenological stages under normal and drought stress conditions. BMC Plant Biology, 24: 483.
- Agati, G., Azzarello, E., Pollastri, S. and Tattini, M., 2020. Flavonoids as antioxidants in plants: Location and functional significance. Plant Science, 196: 67-76. https://doi.org/10.1016/j.plantsci.2012.07.014
- Alami, M.M., Guo, S., Mei, Z., Yang, G. and Wang, X., 2024. Environmental factors on secondary metabolism in medicinal plants: Exploring accelerating factors. Medicinal Plant Biology, 3: e016. https://doi.org/10.48130/mpb-0024-0016
- Alavi-Samani, S.M., Kachouei, M.A. and Pirbalouti, A.G., 2015. Growth, yield, chemical composition, and antioxidant activity of essential oils from two thyme species under foliar application of jasmonic acid and water deficit conditions. Horticulture Environment and Biotechnology, 56: 411-420. https://doi.org/10.1007/s13580-015-0117-y
- Al-Huqail, A., El-Dakak, R.M., Sanad, M.N., Badr, R.H., Ibrahim, M.M., Soliman, D. and Khan, F., 2020. Effects of climate temperature and water stress on plant growth and accumulation of antioxidant compounds in sweet basil (Ocimum basilicum L.) leafy vegetable. Scientifica, 2020: 3808909. https://doi.org/10.1155/2020/3808909
- Ali, S. and Baek, K.H., 2020. Jasmonic acid signaling pathway in response to abiotic stresses in plants. International Journal of Molecular Sciences, 21(2): 621. https://doi.org/ 10.3390/ijms21020621
- Amooaghaie, R. and Rajaie, N., 2025. Foliar application of yeast extract mitigates water deficit stress and elicits hypericin and phenolic production in Hypericum perforatum L. Scientific Reports, 15: 20896. https://doi.org/10.1038/s41598-025-06013-w
- Annunziata, M.G., Apelt, F., Carillo, P., Krause, U., Feil, R., Mengin, V., Lauxmann, M.A., Köhl, K., Nikoloski, Z., Stitt, M., Lunn, J.E. and Raines, C., 2017. Getting back to nature: A reality check for experiments in controlled environments. Journal of Experimental Botany, 68(16): 4463-4477. https://doi.org/10.1093/jxb/erx220
- Ataei Jaliseh, S. and Pourghafar, H., 2026. Investigating the effect of the allicin and cinnamon active ingredients on β-lactam-resistant Pseudomonas aeruginosa: In vitro and in silico study. Iranian Journal of Medicinal and Aromatic Plants Research, 42(1): 40-55.
- Ataeinejad, M., Moradi-Telavat, M.R., Siadat, S.A., Moshtatti, A. and Ghatei, A., 2025. Investigating the combined effect of nitrogen chemical fertilizer, organic fertilizer, sugarcane residue compost and biofarm biological fertilizer on the growth, yield and essential oil of fennel seed (Foeniculum vulgare L.). Iranian Journal of Medicinal and Aromatic Plants Research, 41(4): 580-597.
- Babaee, K., Amini Dehaghi, M., Modares Sanavi, S.A.M. and Jabbari, R., 2010. Water deficit effect on morphology, proline content and thymol percentage of thyme (Thymus vulgaris L.). Iranian Journal of Medicinal and Aromatic Plants Research, 26(2): 239-251. https://doi.org/10.22092/ijmapr.2010.6939
- Bakhshi, F., Fattahi, M. and Ashrafi-Saeidlou, S., 2026. Optimization of bioactive compound extraction from licorice (Glycyrrhiza glabra L.) leaves using response surface methodology (RSM). Iranian Journal of Medicinal and Aromatic Plants Research, 42(1): 75-91. https://doi.org/10.22092/ijmapr.2026.369432.3542
- Baxter, A., Mittler, R. and Suzuki, N., 2014. ROS as key players in plant stress signalling. Journal of Experimental Botany, 65(5): 1229-1240. https://doi.org/10.1093/jxb/ert375
- Berens, M.L., Berry, H.M., Mine, A., Argueso, C.T. and Tsuda, K., 2017. Evolution of hormone signaling networks in plant defense. Annual Review of Phytopathology, 55: 401-425.
- Boscari, A. and Frendo, P., 2025. Redox metabolism and signalling in plants. Journal of Experimental Botany, 76(13): 3629-3633.
- Briskin, D.P., 2000. Medicinal plants and phytomedicines: Linking plant biochemistry and physiology to human health. Plant Physiology, 124(2): 507-514.
- Caser, M., Chitarra, W., D’Angiolillo, F., Perrone, I., Demasi, S., Lovisolo, C., Pistelli, L., Pistelli, L. and Scariot, V., 2019. Drought stress adaptation modulates plant secondary metabolite production in Salvia dolomitica Codd. Industrial Crops and Products, 129: 85-96.
- Catani, M.V., Rinaldi, F., Tullio, V., Gasperi, V. and Savini, I., 2021. Comparative analysis of phenolic composition of six commercially available chamomile (Matricaria chamomilla L.) extracts: Potential biological implications. International Journal of Molecular Sciences, 22(19): 10601. https://doi.org/10.3390/ijms221910601
- Chezem, W.R. and Clay, N.K., 2016. Regulation of plant secondary metabolism and associated specialized cell development by MYBs and bHLHs. Phytochemistry, 131: 26-43.
- Chiappero, J., Cappellari, L.D.R., Palermo, T.B., Giordano, W. and Banchio, E., 2022. Influence of drought stress and PGPR inoculation on essential oil yield and volatile organic compound emissions in Mentha piperita. Horticulturae, 8(12): 1120. https://doi.org/10.3390/horticulturae8121120
- Culurciello, R., Power, K., Esposito, S., Di Nardo, I., Landi, S., De Vico, G., Palatucci, D., Pizzo, E., Naviglio, D. and Zarrelli, A., 2026. Cytotoxic potential evaluation of innovative pressurised cyclic solid-liquid extracts from Withania somnifera. Plants, 15(7): 1027. https://doi.org/10.3390/plants15071027
- Dong, Q., Wallrad, L., Almutairi, B.O. and Kudla, J., 2022. Ca²⁺ signaling in plant responses to abiotic stresses. Journal of Integrative Plant Biology, 64(2): 287-300. https://doi.org/10.1111/jipb.13228
- Erb, M. and Kliebenstein, D.J., 2020. Plant secondary metabolites as defenses, regulators, and primary metabolites: The blurred functional trichotomy. Plant Physiology, 184(1): 39-52. https://doi.org/ 10.1104/pp.20.00433
- Fernie, A.R. and Tohge, T., 2017. The genetics of plant metabolism. Annual Review of Genetics, 51: 287-310. https://doi.org/ 10.1146/annurev-genet-120116-024640
-Figueiredo, A.C., Barroso, J.G., Pedro, L.G. and Scheffer, J.J.C., 2008. Factors affecting secondary metabolite production in plants: Volatile components and essential oils. Flavour and Fragrance Journal, 23(4): 213-226. https://doi.org/ 10.1002/ffj.1875
- Gershenzon, J. and Dudareva, N., 2007. The function of terpene natural products in the natural world. Nature Chemical Biology, 3(7): 408-414. https://doi.org/ 10.1038/nchembio.2007.5
- Ghasemi, M., Babaeian Jelodar, N., Modarresi, M., Bagheri, N. and Jamali, A., 2016. Increase of chamazulene and α-bisabolol contents of the essential oil of German chamomile (Matricaria chamomilla L.) using salicylic acid treatments under normal and heat stress conditions. Foods, 5(3): 56. https://doi.org/10.3390/foods5030056
- Hasanuzzaman, M., Bhuyan, M.H.M.B., Zulfiqar, F., Raza, A., Mohsin, S.M., Mahmud, J.A., Fujita, M. and Fotopoulos, V., 2020. Reactive oxygen species and antioxidant defense in plants under abiotic stress. Antioxidants, 9(8): 681. https://doi.org/ 10.3390/antiox9080681
- Huang, H., Liu, B., Liu, L. and Song, S., 2017. Jasmonate action in plant growth and development. Journal of Experimental Botany, 68(6): 1349-1359. https://doi.org/10.1093/jxb/erw495
- Isah, T., 2019. Stress and defense responses in plant secondary metabolites production. Biological Research, 52: 39. https://doi.org/ 10.1186/s40659-019-0246-3
- Jadczak, D., Bojko, K., Kaymakanova, M. and Berova, M., 2022. Salinity-induced changes in the antioxidant status of common basil plants (Ocimum basilicum L.) grown under controlled conditions. Horticulturae, 8(9): 775. https://doi.org/10.3390/horticulturae8090775
- Jangpangi, D., Patni, B., Chandola, V. and Chandra, S., 2025. Medicinal plants in a changing climate: Understanding the links between environmental stress and secondary metabolite synthesis. Frontiers in Plant Science, 16: 1587337. https://doi.org/10.3389/fpls.2025.1587337
- Kabiri, R., Farahbakhsh, H. and Nasibi, F., 2014. Effect of drought stress on physiological and biochemical characteristics of Nigella sativa L. Iranian Journal of Medicinal and Aromatic Plants Research, 30(4): 600-610. https://doi.org/10.22092/ijmapr.2014.9841
- Kajrolkar, A., 2025. Integrating multi-omics data for plant stress response: Current advances and future directions. Premier Journal of Plant Biology, 3: 100012. https://doi.org/10.70389/ PJPB.100012
- Karayel, H.B., 2025. Examination of changes in essential oil and components in the medicinal sage (Salvia officinalis L.) species grown in different locations. Iranian Journal of Chemistry and Chemical Engineering, 44(8): 2108-2116.
- Karban, R., 2008. Plant behaviour and communication. Ecology Letters, 11(7): 727-739. https://doi.org/10.1111/j.1461-0248.2008.01183.x
- Kheyri, H., Rajabi Mazhar, A., Salimi, F., Tabaei-Aqdai, S.R., Souri, M., Hamzei, J. and Shabani, N., 2026. Selection of compatible genotypes of Rosa damascena Mill. for cold semi-arid conditions in Hamadan based on flower yield and related traits. Iranian Journal of Medicinal and Aromatic Plants Research, 42(1): 56-74.
- Kiani, M.T., Tabatabaeian Nimavard, J. and Nasri, M., 2025. Drought stress affecting the morphological and physiological properties of nettle (Urtica dioica) and tribulus (Tribulus terrestris) medicinal plants. Next Research, 2(4): 100772.
- Kuskun, U., 2026. Climate-driven modulation of plant secondary metabolites: Implications for medicinal quality, standardization, and therapeutic efficacy. Health Academy Kastamonu, 11(1): 50-61. https://doi.org/10.25279/sak.1895917
- Li, Y., Kong, D., Fu, Y., Sussman, M.R. and Wu, H., 2020. The effect of developmental and environmental factors on secondary metabolites in medicinal plants. Plant Physiology and Biochemistry, 148: 80-89. https://doi.org/10.1016/j.plaphy.2020.01.006
- Mahajan, R., Sagar, T., Hussain, S. and Kapoor, N.N., 2023. Secondary metabolite production in medicinal plants under abiotic stress: 127-152. In: Husen, A. and Iqbal, M., (Eds.). Medicinal Plants: Their Response to Abiotic Stress. Springer Nature Singapore, Singapore, 469p.
- Mistry, V., Darji, S., Tiwari, P. and Sharma, A., 2022. Engineering Catharanthus roseus monoterpenoid indole alkaloid pathway in yeast. Applied Microbiology and Biotechnology, 106(7): 2337–2347. https://doi.org/10.1007/s00253-022-11883-5
- Moazzami Farida, S.H., Rahmani, N., Taghizadeh, M. and Albrectsen, B.R., 2026. Dopamine modulates antioxidant and phenolic responses to alleviate nickel stress in Salvia officinalis. BMC Plant Biology, 26: 491. https://doi.org/10.1186/s12870-026-08365-5
- Naghdi Badi, H.A., Abdollahi, M., Mehrafarin, A., Ghorbanpour, M., Tolyat, S.M., Qaderi, A. and Ghiaci Yekta, M., 2017. An overview on two valuable natural and bioactive compounds, thymol and carvacrol, in medicinal plants. Journal of Medicinal Plants, 16(63): 1-32. http://jmp.ir/article-1-1370-en.html
- Neilson, E.H., Goodger, J.Q.D., Woodrow, I.E. and Møller, B.L., 2013. Plant chemical defense: At what cost? Trends in Plant Science, 18(5): 250-258. https://doi.org/10.1016/j.tplants.2013.01.001
- Ninkuu, V., Aluko, O.O., Yan, J., Zeng, H., Liu, G., Zhao, J., Li, H., Chen, S. and Dakora, F.D., 2025. Phenylpropanoid metabolism: Recent insights into stress tolerance and plant development cues. Frontiers in Plant Science, 16: 1571825. https://doi.org/10.3389/fpls.2025.1571825
- Noctor, G., Reichheld, J.P. and Foyer, C.H., 2018. ROS-related redox regulation and signaling in plants. Seminars in Cell & Developmental Biology, 80: 3-12. https://doi.org/10.1016/j.semcdb.2017.07.013
- Omrani, M., Ghasemi, M., Modarresi, M. and Salamon, I., 2023. Alternations in physiological and phytochemical parameters of German chamomile (Matricaria chamomilla L.) varieties in response to amino acid fertilizer and plasma-activated water treatments. Horticulturae, 9(8): 857.
- Pichersky, E. and Lewinsohn, E., 2011. Convergent evolution in plant specialized metabolism. Annual Review of Plant Biology, 62: 549-566. https://doi.org/10.1146/annurev-arplant-042110-103814
- Rahdar, S., Esmaeilzadeh Bahabadi, S. and Mohkami, Z., 2026. Effect of chitosan and carrageenan spraying on physiological parameters of indigo (Indigofera tinctoria L.). Iranian Journal of Medicinal and Aromatic Plants Research, 42(1): 1-14. https://doi.org/10.22092/ijmapr.2025.366494.3461
- Rai, A., Saito, K. and Yamazaki, M., 2017. Integrated omics analysis of specialized metabolism in medicinal plants. The Plant Journal, 90(4): 764-787. https://doi.org/10.1111/tpj.13485
- Safaii, L., Sharifi Ashoorabadi, E. and Amin Azarm, D., 2025. Comparison of quantitative and qualitative performance of Ziziphora clinopodioides, Z. capitata, and Z. tenuior grown under cultivated conditions in Isfahan province. Iranian Journal of Medicinal and Aromatic Plants Research, 41(4): 561-579. https://doi.org/10.22092/ijmapr.2025.367476.3488
- Santino, A., Taurino, M., De Domenico, S., Bonsegna, S., Poltronieri, P., Pastor, V. and Flors, V., 2013. Jasmonate signaling in plant development and defense response to multiple (a) biotic stresses. Plant Cell Reports, 32(7): 1085-1098. https://doi.org/10.1007/s00299-013-1441-2
- Selmar, D. and Kleinwachter, M., 2013. Stress enhances the synthesis of secondary plant products: The impact of stress-related over-reduction on the accumulation of natural products. Plant and Cell Physiology, 54(6): 817-826. https://doi.org/10.1093/pcp/pct054
- Schuurink, R.C. and Tissier, A., 2020. Glandular trichomes: Micro-organs with model status? New Phytologist, 225(6): 2251-2266.
- Scossa, F., Alseekh, S. and Fernie, A.R., 2021. Integrating multi-omics data for crop improvement. Journal of Plant Physiology, 257: 153352. https://doi.org/10.1016/j.jplph.2020.153352
- Shahhat, I.M.A., Guetat, A., Elsheik, S.Y.S., Abu-Tahon, M.A., Abdelwahab, A.T. and Awad, M.A., 2025. Sustainable medicinal plant production-responses of Lamiaceae plants to organic acid elicitor spraying during environmental stress: A review. Plant Protection Science, 61(2): 110-151. https://doi.org/10.17221/83/2024-PPS
- Shen, Q., Zhang, L., Liao, Z., Wang, S., Yan, T., Shi, P., Liu, M., Fu, X., Pan, Q., Wang, Y., Lv, Z., Lu, X., Zhang, F., Jiang, W., Ma, Y., Chen, M., Hao, X., Li, L., Tang, Y., Lv, G., Zhou, Y., Sun, X., Brodelius, P.E., Rose, J.K.C. and Tang, K., 2018. The genome of Artemisia annua provides insight into the evolution of the Asteraceae family and artemisinin biosynthesis. Molecular Plant, 11(6): 776-788. https://doi.org/10.1016/j.molp.2018.03.015
- Takubessi, M.I., Jalil, B. and Heinrich, M., 2025. The impact of climate change on medicinal plants and natural products: A scoping review. Frontiers in Pharmacology, 16: 1697581.
- Tan, U. and Gören, H.K., 2024. Comprehensive evaluation of drought stress on medicinal plants: A meta-analysis. PeerJ, 12: e17801.
- Tissier, A., 2012. Glandular trichomes: What comes after expressed sequence tags? The Plant Journal, 70(1): 51-68. https://doi.org/10.1111/j.1365-313X.2012.04913.x
- Wang, G., Sun, X., Li, Y., Wang, Y. and Jin, C., 2025. The role of UV-B radiation in modulating secondary metabolite biosynthesis and regulatory mechanisms in medicinal plants. BioResources, 20(2): 4776-4797. https://ojs.bioresources.com/index.php/BRJ/article/view/24205
- Wasternack, C. and Strnad, M., 2018. Jasmonates: News on occurrence, biosynthesis, metabolism and action of an ancient group of signaling compounds. International Journal of Molecular Sciences, 19(9): 2539. https://doi.org/10.3390/ijms19092539
- Waszczak, C., Carmody, M. and Kangasjärvi, J., 2018. Reactive oxygen species in plant signaling. Annual Review of Plant Biology, 69: 209-236. https://doi.org/10.1146/annurev-arplant-042817-040322
- Yang, C.Q., Fang, X., Wu, X.M., Mao, Y.B., Wang, L.J. and Chen, X.Y., 2012. Transcriptional regulation of plant secondary metabolism. Journal of Integrative Plant Biology, 54(10): 703-712. https://doi.org/10.1111/j.1744-7909.2012.01161.x
- Yang, L., Wen, K.S., Ruan, X., Zhao, Y.X., Wei, F. and Wang, Q., 2018. Response of plant secondary metabolites to environmental factors. Molecules, 23(4): 762.
- Yu, T., Ma, X., Zhang, J., Cao, S., Li, W., Yang, G. and He, C., 2025. Progress in transcriptomics and metabolomics in plant responses to abiotic stresses. Current Issues in Molecular Biology, 47(6): 421. https://doi.org/10.3390/cimb47060421
- Zandalinas, S.I., Mittler, R., Balfagón, D., Arbona, V. and Gómez-Cadenas, A., 2018. Plant adaptations to the combination of drought and high temperatures. Physiologia Plantarum, 162(1): 2-12. https://doi.org/10.1111/ppl.12540
- Zarei, Z., Khaliliaqdam, N., Saeidian, S., Rashidzadeh, B. and Ghadermarzi, M., 2026. Kinetic evaluation of ascorbate peroxidase in black mulberry leaf and fruit with the help of linear and non-linear regression analysis. Iranian Journal of Medicinal and Aromatic Plants Research, 42(1): 15-39. https://doi.org/10.22092/ijmapr.2026.368405.3515
- Zhang, H., Hedhili, S., Montiel, G., Zhang, Y., Chatel, G., Pré, M., Gantet, P. and Memelink, J., 2011. The basic helix-loop-helix transcription factor CrMYC2 controls the jasmonate-responsive expression of the ORCA genes that regulate alkaloid biosynthesis in Catharanthus roseus. The Plant Journal, 67(1): 61-71.
- Zheng, Z., Gao, H. and Yang, Q., 2026. WRKY-mediated regulation of secondary metabolite production in medicinal plants. Journal of Holistic Integrative Pharmacy, 7(2): 249-259. https://doi.org/10.1016/j.jhip.2026.05.002
- Zust, T. and Agrawal, A.A., 2017. Trade-offs between plant growth and defense against insect herbivory: An emerging mechanistic synthesis. Annual Review of Plant Biology, 68: 513-534. https://doi.org/10.1146/annurev-arplant-042916-040856