Journals / Turkish Journal of Range and Forage Science / 2026 / Volume: 7 Issue: 1

Root System Architecture and Anatomical Adaptations under Drought Stress in Forage Species: Mechanisms, Phenotyping Advances, and Implications for Climate Resilience

Muhammad Khizar Hayat(SAKARYA UYGULAMALI BİLİMLER ÜNİVERSİTESİ, ZİRAAT FAKÜLTESİ)0009-0005-4998-7138
Mehmet Öten(SAKARYA UYGULAMALI BİLİMLER ÜNİVERSİTESİ, ZİRAAT FAKÜLTESİ)0000-0002-3737-2356
Rahime Cengiz(SAKARYA UYGULAMALI BİLİMLER ÜNİVERSİTESİ, ZİRAAT FAKÜLTESİ)0000-0001-6355-7496
Pages
—
Publication Date
2026-07-10
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AI Summary & Key Takeaways

AI Insight
⚡ TL;DR(Too Long; Didn't Read)

By integrating these multi-scale traits with high-throughput phenotyping and symbiotic interactions, this paper provides a framework for breeding climate-resilient forage ideotypes essential for sustaining livestock systems and global soil carbon stocks.

📌 3 Key Takeaways

🎯
Objective & Hypothesis:Drought stress represents the most significant threat to global forage productivity, necessitating a robust understanding of adaptive belowground mechanisms.
📊
Key Findings & Data:Under water-limited conditions, forages exhibit substantial morphological plasticity, characterized by a 20-40% increase in primary root elongation and a strategic rise in root-to-shoot ratios to prioritize deep-layer moisture acquisition.
💡
Conclusion & Impact:By integrating these multi-scale traits with high-throughput phenotyping and symbiotic interactions, this paper provides a framework for breeding climate-resilient forage ideotypes essential for sustaining livestock systems and global soil carbon stocks.
🔬Design:Review & Systematic Survey
📍Scope:Comprehensive Sample
⚙️Variables:Forage Crops · Drought Stress · Root System Architecture

Abstract

Drought stress represents the most significant threat to global forage productivity, necessitating a robust understanding of adaptive belowground mechanisms. This paper highlights the critical advancements in root system architecture (RSA) and root vascular architecture that underpin resilience in forages. Under water-limited conditions, forages exhibit substantial morphological plasticity, characterized by a 20-40% increase in primary root elongation and a strategic rise in root-to-shoot ratios to prioritize deep-layer moisture acquisition. Anatomically, they optimize hydraulic safety through a 10-40% reduction in xylem vessel diameter, which enhances embolism resistance by 20-35% despite a minor decline in hydraulic conductivity. Metabolic efficiency is further bolstered by the development of root cortical aerenchyma, which reduces cortical area by 30-50% and lowers maintenance respiration costs by up to 60%. Parallel to these metabolic shifts, the reinforcement of endodermal barriers specifically 1.5-2 times thickening of the Casparian strip and a 1.5-2.2 times elongation of root hairs maximizes water-use efficiency and soil-root contact. The review further examines the physiological drivers of these changes, indicating that moderate ABA accumulation can contribute to the maintenance or stimulation of primary root elongation under stress, with reported increases of 20–40% depending on genotype and environmental conditions. By integrating these multi-scale traits with high-throughput phenotyping and symbiotic interactions, this paper provides a framework for breeding climate-resilient forage ideotypes essential for sustaining livestock systems and global soil carbon stocks.

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Cite This Article

Hayat, M. K., Öten, M., Cengiz, R. (2026). Root System Architecture and Anatomical Adaptations under Drought Stress in Forage Species: Mechanisms, Phenotyping Advances, and Implications for Climate Resilience. Turkish Journal of Range and Forage Science, Volume: 7 Issue: 1. https://doi.org/10.51801/turkjrfs.1903705

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References (218)

  1. Abdalla, M., Bitterlich, M., Jansa, J., Püschel, D., & Ahmed, M. A. (2023). The role of arbuscular mycorrhizal symbiosis in improving plant water status under drought. Journal of Experimental Botany, 74(16), 4808-4824.
  2. Abdalla, G., Awad, H., & Rashed, U. M. (2022). Different cleaning techniques for removal of iron stain from archaeological bone artifacts: A review. Egyptian Journal of Chemistry, 65(5), 69-83.
  3. Abdullah, H. M., Atta, M., ul Haq, I., Shah, M. A. R., Mubashar, M., & Hayat, M. K. (2025). Soybean's Genetic Odyssey: Unveiling Diverse Transformation Strategies. In Policy Research Journal, 3(6), 297-326.
  4. Abou Seeda, M., Abou El-Nour, E. Z., Abdallah, M., & El-Bassiouny, H. (2022). Impacts of metal, metalloid and their effects in plant physiology: a review. Middle East J Agric Res, 11(3), 838-931.
  5. Acharya, A., Chatterjee, R., Das, M., & Latif, M. (2025). Impact of Climate Change-Induced Abiotic Stress on Plant Growth, Biodiversity, and Adaptations. In Impact of Climate Change on Medicinal and Herbal Plant microRNA (pp. 11-41). CRC Press.
  6. Adão, F., Pádua, L., & Sousa, J. J. (2025). Evaluating soil degradation in agricultural soil with ground-penetrating radar: A systematic review of applications and challenges. Agriculture, 15(8), 852.
  7. Adar, S., Sternberg, M., Argaman, E., Henkin, Z., Dovrat, G., Zaady, E., & Paz-Kagan, T. (2023). Testing a novel pasture quality index using remote sensing tools in semiarid and Mediterranean grasslands. Agriculture, Ecosystems & Environment, 357, 108674.
  8. Adepoju, S. A., & Faniyi, T. O. (2025). Synergizing plant breeding and animal nutrition: A review on the development of drought-tolerant and high-nutrient forage grasses and legumes. Innovations in Agriculture, 8, 1-10.
  9. Akhiyarova, G., Veselov, D., Ivanov, R., Sharipova, G., Ivanov, I., Dodd, I. C., & Kudoyarova, G. (2023). Root ABA accumulation delays lateral root emergence in osmotically stressed barley plants by decreasing root primordial IAA accumulation. International Journal of Plant Biology, 14(1), 77-90.
  10. Alagoz, S. M., Lajayer, B. A., & Ghorbanpour, M. (2023). Proline and soluble carbohydrates biosynthesis and their roles in plants under abiotic stresses. In Plant stress mitigators (pp. 169-185). Elsevier.
  11. Alahmad, S., Smith, D., Katsikis, C., Aldiss, Z., Brunner, S. M., Meer, S. V., Meijer, L., Heidariask, B., Chenu, K., & Chapman, S. (2025). Phenotyping the hidden half: combining UAV phenotyping and machine learning to predict barley root traits in the field. Journal of Experimental Botany, 76(17), 5161-5178.
  12. Alam, M. M., Rafi, A., Rahman, M. A., Mitu, M. A., Sakib, M. N., Rahman, M. A., Rahimi, M., Kordrostami, M., Biswas, A., & Rashad, M. M. I. (2026). Aquaporins as Natural Stress Integrator: Coordinating Transport, Signals, and Tolerance Mechanisms in Plants. Plant, Cell & Environment.
  13. Ali, M. F., Li, P., Ma, L., Feng, C., Gu, S., Lin, X., & Wang, D. (2025). Investigating the Impact of Drought During Returning Green and Jointing Stages on Different Tillers Position of Winter Wheat. Journal of Plant Growth Regulation, 44(9), 5542-5560.
  14. Allmendinger, C. (2025). Back to the roots: Ecological extremes shape barrier formation in plant roots across species
  15. Alomair, A. M. (2025). Enhancing Alfalfa Persistence in Rangelands: Evaluating Morphological Traits of Medicago sativa Subsp. Falcata Germplasm South Dakota State University].
  16. Andersen, T. G., & Drapek, C. (2024). The endodermis 1. Plant roots, 90-103.
  17. Angidi, S., Madankar, K., Tehseen, M. M., & Bhatla, A. (2025). Advanced high-throughput phenotyping techniques for managing abiotic stress in agricultural crops—A comprehensive review. Crops, 5(2), 8.
  18. Araújo, S. S., Beebe, S., Crespi, M., Delbreil, B., González, E. M., Gruber, V., Lejeune-Henaut, I., Link, W., Monteros, M. J., & Prats, E. (2015). Abiotic stress responses in legumes: strategies used to cope with environmental challenges. Critical Reviews in Plant Sciences, 34(1-3), 237-280.
  19. Asadullah, Kalhoro, S. A., Farhad, W., Iqbal, A., Sultan, Waheed, A., Rashid, M., & Shah, S. R. U. (2024). Exploring the variability of root system architecture under drought stress in heat-tolerant spring-wheat lines. Plant and Soil, 502(1), 103-119.
  20. Bai, Y., & Cotrufo, M. F. (2022). Grassland soil carbon sequestration: Current understanding, challenges, and solutions. science, 377(6606), 603-608.
  21. Becker, S. M. (2025). Soil Characterization with Proximal Geophysics The University of Nebraska-Lincoln].
  22. Benmrid, B., Ghoulam, C., Zeroual, Y., Kouisni, L., & Bargaz, A. (2023). Bioinoculants as a means of increasing crop tolerance to drought and phosphorus deficiency in legume-cereal intercropping systems. Communications Biology, 6(1), 1016.
  23. Berlingeri, J., Fuentes, A., Ranario, E., Yun, H., Rim, E. Y., Garrett, O., Howard, A., LaPorte, M.-F., Lo, S., & Pauli, D. (2025). Integration of crop modeling and sensing into molecular breeding for nutritional quality and stress tolerance. Theoretical and Applied Genetics, 138(9), 205.
  24. Bhattacharya, A. (2022). Low temperature stress and plant-water relationship: A review. Physiological processes in plants under low temperature stress, 107-197.
  25. Bian, L., Zhang, H., Ge, Y., Čepl, J., Stejskal, J., & El-Kassaby, Y. A. (2022). Closing the gap between phenotyping and genotyping: review of advanced, image-based phenotyping technologies in forestry. Annals of Forest Science, 79(1), 22.
  26. Bisht, D., Sharma, U., & Kumar, N. (2025). Anatomical Traits: Adaptations for Resource Acquisition and Allocation. Enhancing Forest Resilience through Plant Functional Traits, 32-45.
  27. Branca, F., Argento, S., Paoletti, A. M., & Melilli, M. G. (2022). The physiological role of inulin in wild cardoon (Cynara cardunculus L. var. sylvestris Lam.). Agronomy, 12(2), 290.
  28. Bucior, E. R., & Sorensen, R. B. (2025). A Shovel-omics Facelift: Exploring Inexpensive and Simple Root Phenotyping Techniques in Peanut. Peanut Science, 52(1).
  29. Byrt, C. S., Zhang, R. Y., Magrath, I., Chan, K. X., De Rosa, A., & McGaughey, S. (2023). Exploring aquaporin functions during changes in leaf water potential. Frontiers in Plant Science, 14, 1213454.
  30. Cai, G., & Ahmed, M. A. (2022). The role of root hairs in water uptake: recent advances and future perspectives. Journal of Experimental Botany, 73(11), 3330-3338.
  31. Cavus, Y., Stahl, K., & Aksoy, H. (2023). Drought intensity–duration–frequency curves based on deficit in precipitation and streamflow for water resources management. Hydrology and Earth System Sciences, 27(18), 3427-3445.
  32. Chandnani, R., & Soolanayakanahally, R. (2025). The Invisible Frontline: High‐Tech Root Imaging for Crop Stress Adaptation. Physiologia Plantarum, 177(5), e70572.
  33. Chen, A., Liu, T., Wang, Z., & Chen, X. (2022). Plant root suberin: a layer of defence against biotic and abiotic stresses. Frontiers in Plant Science, 13, 1056008.
  34. Chen, Z., Zhu, L.-T., & Luo, Z.-H. (2023). Characterizing flow and transport in biological vascular systems: A review from physiological and chemical engineering perspectives. Industrial & Engineering Chemistry Research, 63(1), 4-36.
  35. Chepkemoi, J., Onwonga, R. N., Gitau, A., & Bitange, N. (2025). Potential of perennial forages on soil carbon sequestration across agroecological zones with varying management practices in Meru County, Kenya. Discover Soil, 2(1), 3.
  36. Correa, J., Postma, J. A., Watt, M., & Wojciechowski, T. (2019). Soil compaction and the architectural plasticity of root systems. Journal of Experimental Botany, 70(21), 6019-6034.
  37. Cruz, N. T., Jardim, R. R., de Lana Sousa, B. M., Seixas, A. A., Fries, D. D., Pires, A. J. V., Dias, D. L. S., Bonomo, P., Ramos, B. L. P., & Alcantara, W. (2022). Energy flows and organic reserves of forage plants. Research, Society and Development, 11(12).
  38. Cseresnyés, I., Rajkai, K., Szitár, K., Radimszky, L., Ónodi, G., & Kröel-Dulay, G. (2020). Root capacitance measurements allow non-intrusive in-situ monitoring of the seasonal dynamics and drought response of root activity in two grassland species. Plant and Soil, 449(1), 423-437.
  39. Cui, Y.-N., Yan, S.-J., Zhang, Y.-N., Wang, R., Song, L.-L., Ma, Y., Guo, H., & Yang, P.-Z. (2024). Physiological, metabolome and gene expression analyses reveal the accumulation and biosynthesis pathways of soluble sugars and amino acids in sweet sorghum under osmotic stresses. International Journal of Molecular Sciences, 25(16), 8942.
  40. Danelia, G. V., Emelyanov, V. V., & Shishova, M. F. (2024). The role of water-transporting aquaporins of the PIP and TIP subfamilies in plant development and adaptation to stress factors. Ecological genetics, 22(4), 343-368.
  41. De Bruijn, F. J., & Hungria, M. (2022). Biological nitrogen fixation. Good microbes in medicine, food production, biotechnology, bioremediation, and agriculture, 466-475. de Moraes, D. H. M., Mesquita, M., Graciano-Ribeiro, D., de Araújo, D. S., Battisti, R., Flores, R. A., de Melo, H. C., &
  42. Casaroli, D. (2022). The effect of xylem vessel diameter on potential hydraulic conductivity in different rice stem longitudinal positions. Flora, 295, 152147.
  43. De Silva, N. (2019). The roles of suberin biopolymer and associated waxes in protecting plants against abiotic stresses Carleton University].
  44. Dimattia, B. G., Righi, A., Bettuzzi, M., Koestel, J., Morigi, M. P., Brancaccio, R., Salvi, S., Hernandez-Soriano, M. C., & Bittelli, M. (2025). Root traits of different wheat cultivars influence soil structure: an X-ray computed tomography and root morphology study. Geoderma, 459, 117349.
  45. Ding, R., Nóbrega, R. L., & Prentice, I. C. (2025). Global Assessment of Environmental and Plant‐Trait Influences on Root: Shoot Biomass Ratios. Global Change Biology, 31(10), e70543.
  46. Dini‐Andreote, F., Wells, D. M., Atkinson, J. A., Atkinson, B. S., Finkel, O. M., & Castrillo, G. (2025). Microbial drivers of root plasticity. New Phytologist, 248(1), 52-67.
  47. Dolezal, J., Jandova, V., Macek, M., & Liancourt, P. (2021). Contrasting biomass allocation responses across ontogeny and stress gradients reveal plant adaptations to drought and cold. Functional Ecology, 35(1), 32-42.
  48. Dollete, D., Lumactud, R. A., Carlyle, C. N., Szczyglowski, K., Hill, B., & Thilakarathna, M. S. (2024). Effect of drought stress on symbiotic nitrogen fixation, soil nitrogen availability and soil microbial diversity in forage legumes. Plant and Soil, 495(1), 445-467.
  49. Dowd, T., McInturf, S., Li, M., & Topp, C. N. (2021). Rated-M for mesocosm: allowing the multimodal analysis of mature root systems in 3D. Emerging Topics in Life Sciences, 5(2), 249-260.
  50. Duan, D., Feng, X., Wu, N., Tian, Z., Dong, X., Liu, H., Nan, Z., & Chen, T. (2023). Drought eliminates the difference in root trait plasticity and mycorrhizal responsiveness of two semiarid grassland species with contrasting root system. International Journal of Molecular Sciences, 24(12), 10262.
  51. Duan, H.-X., Luo, C.-L., Wang, X., Cheng, Y.-S., Abrar, M., & Batool, A. (2024). Responses of legumes to rhizobia and arbuscular mycorrhizal fungi under abiotic stresses: a global meta-analysis. Agronomy, 14(11), 2597.
  52. El Hamss, H., Radouane, N., Belabess, Z., & Lahlali, R. (2023). Microbial consortia: an approach to enhance the effectiveness of beneficial soil microbes. In Novel biological and biotechnological applications in plant nematode management (pp. 133-166). Springer.
  53. Farooq, M., Rafique, S., Zahra, N., Rehman, A., & Siddique, K. H. (2024). Root system architecture and salt stress responses in cereal crops. Journal of Agronomy and Crop Science, 210(6), e12776.
  54. Farooq, M. S., Uzair, M., Maqbool, Z., Fiaz, S., Yousuf, M., Yang, S. H., & Khan, M. R. (2022). Improving nitrogen use efficiency in aerobic rice based on insights into the ecophysiology of archaeal and bacterial ammonia oxidizers. Frontiers in Plant Science, 13, 913204.
  55. Franco‐Navarro, J. D., Padilla, Y. G., Álvarez, S., Calatayud, Á., Colmenero‐Flores, J. M., Gómez‐Bellot, M. J., Hernández, J. A., Martínez‐Alcalá, I., Penella, C., & Pérez‐Pérez, J. G. (2025). Advancements in Water‐Saving strategies and crop adaptation to drought: A comprehensive review. Physiologia Plantarum, 177(4), e70332.
  56. Franklin, O., Fransson, P., Hofhansl, F., Jansen, S., & Joshi, J. (2023). Optimal balancing of xylem efficiency and safety explains plant vulnerability to drought. Ecology letters, 26(9), 1485-1496.
  57. Freschet, G. T., Pagès, L., Iversen, C. M., Comas, L. H., Rewald, B., Roumet, C., Klimešová, J., Zadworny, M., Poorter, H., & Postma, J. A. (2021). A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements. New Phytologist, 232(3), 973-1122.
  58. Freschet, G. T., Violle, C., Bourget, M. Y., Scherer‐Lorenzen, M., & Fort, F. (2018). Allocation, morphology, physiology, architecture: The multiple facets of plant above‐and below‐ground responses to resource stress. New Phytologist, 219(4), 1338-1352.
  59. Galindo-Castañeda, T., Lynch, J. P., Six, J., & Hartmann, M. (2022). Improving soil resource uptake by plants through capitalizing on synergies between root architecture and anatomy and root-associated microorganisms. Frontiers in Plant Science, 13, 827369.
  60. Galindo-CastanedaLynch, T. (2018). Microbial tradeoffs of root cortical aerenchyma in maize. The Pennsylvania State University.
  61. Gano, B., Bhadra, S., Vilbig, J. M., Ahmed, N., Sagan, V., & Shakoor, N. (2024). Drone‐based imaging sensors, techniques, and applications in plant phenotyping for crop breeding: A comprehensive review. The Plant Phenome Journal, 7(1), e20100.
  62. Gao, Y.-Q., Su, Y., & Chao, D.-Y. (2024). Exploring the function of plant root diffusion barriers in sealing and shielding for environmental adaptation. Nature Plants, 10(12), 1865-1874.
  63. Gao, Y., Yang, Z., Wang, G., Sun, J., & Zhang, X. (2020). Discerning the difference between lumens and scalariform perforation plates in impeding water flow in single xylem vessels and vessel networks in cotton. Frontiers in Plant Science, 11, 246.
  64. Garbowski, M., Avera, B., Bertram, J. H., Courkamp, J. S., Gray, J., Hein, K. M., Lawrence, R., McIntosh, M., McClelland, S., & Post, A. K. (2020). Getting to the root of restoration: considering root traits for improved restoration outcomes under drought and competition. Restoration Ecology, 28(6), 1384-1395.
  65. Gardner, M. J., Condon, J. R., Dear, B. S., Conyers, M. K., & Li, G. D. (2023). Comparison of nitrogen mineralisation patterns of Cichorium intybus and Medicago sativa. European Journal of Soil Science, 74(6), e13443.
  66. Gavrilescu, M. (2021). Water, soil, and plants interactions in a threatened environment. Water, 13(19), 2746. Ghosh, T., Maity, P. P., Rabbi, S. M., Das, T., & Bhattacharyya, R. (2023). Application of X-ray computed tomography in soil and plant-a review. Frontiers in Environmental Science, 11, 1216630.
  67. Giehl, R. F., & von Wirén, N. (2014). Root nutrient foraging. Plant physiology, 166(2), 509-517.
  68. Gong, W., Ju, Z., Chai, J., Zhou, X., Lin, D., Su, W., & Zhao, G. (2022). Physiological and transcription analyses reveal the regulatory mechanism in oat (Avena sativa) seedlings with different drought resistance under PEG-induced drought stress. Agronomy, 12(5), 1005.
  69. Grossiord, C., Ulrich, D. E., & Vilagrosa, A. (2020). Controls of the hydraulic safety–efficiency trade-off. Tree Physiology, 40(5), 573-576.
  70. Guo, T., Niu, T., Kuka, K., & Tippkötter, N. (2025). Optimizing silage efficiency: The role of ryegrass varieties, harvest time, and additives in enhancing perennial ryegrass (Lolium perenne) fermentation. Fermentation, 11(4), 192.
  71. Gusain, S., Kumari, K., & Joshi, R. (2024). Physiological, hormonal and molecular dynamics of root system architectural response to drought stress signaling in crops. Rhizosphere, 31, 100922.
  72. Hanif, S., Hayat, M. K., Zaheer, M., Raza, H., Ain, Q. U., Razzaq, A., Sehar, A., & Raza, A. (2024a). Impact of climate change on agriculture production and strategies to overcome. Cornous Biology, 2(2), 1-7.
  73. Hanif, S., Sial, T. N., Hayat, M. K., Tariq, M., Mehwish, S., Usman, M., Shabbir, M. A., Benish, S. H., Bilal, M., & Abdullah, M. H. (2024b). Enhancing sesame yield in arid conditions through NPK fertilizer optimization. Asian Plant Research Journal, 12(3), 1-11.
  74. Hasan, M. M., Liu, X.-D., Rahman, M. A., Hazzazi, Y., Wassem, M., Ghimire, S., Alabdallah, N. M., Alharbi, B. M., Kabir, A. H., & Yao, G. (2025). Plants breathing under pressure: mechanistic insights into soil compaction-induced physiological, molecular and biochemical responses in plants. Planta, 261(3), 52.
  75. Hayat, M. K., Khan, T. Y., Azam, M. A., Imran, F., Ifra, I., Shan, M., Belizaire, P. J., Rehman, M. N. U., Majeed, U., & Oumar, B. (2025). Introduction to Organic Agriculture-From Haber-Bosch to Howard: A Century of Agricultural Transformation. World News of Natural Sciences, 60, 40-64.
  76. Hayat, M. K., Mumtaz, M. M., Ahsan Ali, M., Huaira, A., Ur Rehman, M., Shabbir, M. A., Usman, M., & Afzal, A. (2024). Optimizing Livestock Feed Systems: A Multi-Faceted Approach for Sustainable and Resilient Animal Agriculture; A Comprehensive Review. J. Plant Biota, 3, 27-33.
  77. Hochholdinger, F., & Yu, P. (2026). The Genetic Basis and Domestication of Root System Architecture in Cereals. Annual review of plant biology, 77.
  78. Hölttä, T., & Sperry, J. (2013). Plant water transport and cavitation. In Transport and reactivity of solutions in confined hydrosystems (pp. 173-181). Springer.
  79. Hose, E., Clarkson, D. T., Steudle, E., Schreiber, L., & Hartung, W. (2001). The exodermis: a variable apoplastic barrier. Journal of Experimental Botany, 52(365), 2245-2264.
  80. Hu, Y., Pandey, A. K., Wu, X., Fang, P., & Xu, P. (2022). The role of arbuscular mycorrhiza fungi in drought tolerance in legume crops: a review. Legume Research: An International Journal, 45(1).
  81. Huang, W. (2024). Boosting soil health: the role of rhizobium in legume nitrogen fixation. Molecular Soil Biology, 15.
  82. Islam, M. S., Fahad, S., Hossain, A., Chowdhury, M. K., Iqbal, M. A., Dubey, A., Kumar, A., Rajendran, K., Danish, S., & Rahman, M. H. U. (2021). Legumes under drought stress: plant responses, adaptive mechanisms, and management strategies in relation to nitrogen fixation. In Engineering tolerance in crop plants against abiotic stress (pp. 179-207). CRC Press.
  83. Islam, W., Zhihao, Z., Khan, K. A., & Zeng, F. (2026). Resilience and Adaptation in Desert Ecosystems: Unveiling Microbial Legacies and Plant Functional Trait Coordination Under Climate Change. Global Change Biology, 32(1), e70678.
  84. Jayasinghe, C., Jacobs, J., Thomson, A., & Smith, K. (2023). Evaluation of the relationship between cultivar, endophyte and environment on the expression of persistence in perennial ryegrass populations using high-throughput phenotyping. Agronomy, 13(9), 2292.
  85. Jayasinghe, C., Thomson, A., Smith, K., & Jacobs, J. (2024). Early indicators of declining pasture persistence: sensor-based tools for paddock-scale identification. Crop & Pasture Science, 75(12), CP24124.
  86. Jeandet, P., Formela-Luboińska, M., Labudda, M., & Morkunas, I. (2022). The role of sugars in plant responses to stress and their regulatory function during development. In (Vol. 23, pp. 5161): MDPI.
  87. Jiang, H., Jiang, L., He, L., Murengami, B. G., Jing, X., Misiewicz, P. A., Cheein, F. A., & Fu, L. (2024). Yield prediction of root crops in field using remote sensing: A comprehensive review. Computers and Electronics in Agriculture, 227, 109600.
  88. Jiang, H., Zeng, Y.-Y., Liu, X.-D., Corpas, F. J., Hasan, M. M., Li, F.-P., Tian, X.-Q., Zhang, Y.-H., He, R.-Y., & Fang, X.-W. (2025). Rooting for survival: ABA-mediated hormonal integration driving primary root elongation under drought stress. Journal of Plant Ecology, rtaf172.
  89. Jinagool, W. (2015). Genetic variability of drought tolerance of trees of agronomic interest: the role of vulnerability to xylem cavitation Université Blaise Pascal-Clermont-Ferrand II].
  90. Judd, L. A., Jackson, B. E., & Fonteno, W. C. (2015). Advancements in root growth measurement technologies and observation capabilities for container-grown plants. Plants, 4(3), 369-392.
  91. Kagan, I. A. (2022). Water-and ethanol-soluble carbohydrates of temperate grass pastures: a review of factors affecting concentration and composition. Journal of equine veterinary science, 110, 103866.
  92. Kakusu, E. M., Kinuthia, R., Ott, J. P., & Koech, O. (2025). Belowground resprouting potential and nutrient acquisition traits of three perennial grasses during the short dry season in Kenya. African Journal of Range & Forage Science, 42(2), 146-157.
  93. Kalra, A., Goel, S., & Elias, A. A. (2024). Understanding role of roots in plant response to drought: Way forward to climate‐resilient crops. The Plant Genome, 17(1), e20395.
  94. Karthika, K., Rashmi, I., & Parvathi, M. (2018). Biological functions, uptake and transport of essential nutrients in relation to plant growth. In Plant nutrients and abiotic stress tolerance (pp. 1-49). Springer.
  95. Kaur, V., Yadav, S. K., Yadav, B., Madaan, S., Kheralia, M., & Chinnusamy, V. (2026). Root system architecture and drought adaptation: emerging tools and genetic insights. Frontiers in Plant Science, 17, 1753086.
  96. Kearney, S. P., Porensky, L. M., Augustine, D. J., Derner, J. D., & Gao, F. (2022). Predicting spatial‐temporal patterns of diet quality and large herbivore performance using satellite time series. Ecological Applications, 32(2), e2503.
  97. Keerthi, G. M., Mallikarjuna, M. G., Jha, S. K., Pandey, R., Veeraya, P., Lohithaswa, H. C., & Chinnusamy, V. (2025). Unravelling root system architecture plasticity in response to abiotic stresses in maize. Scientific Reports, 15(1), 20433.
  98. Keret, R. (2024). The physiological and genetic underpinnings that influence the xylem properties of Eucalyptus grandis, when subjected to different levels of water availability. Stellenbosch University.
  99. Khangura, R., Ferris, D., Wagg, C., & Bowyer, J. (2023). Regenerative agriculture—A literature review on the practices and mechanisms used to improve soil health. Sustainability, 15(3), 2338.
  100. Kirbas, G. (2025). The Impact of Climate Change and Soil Management on Root System Architecture and rhizosphere processes in Winter Wheat-Insight from an Ecotron experiment.
  101. Klein, S. (2021). Functional implications and association mapping of root hydraulic traits for improved drought tolerance in maize (Zea mays L.).
  102. Kou, X., Han, W., & Kang, J. (2022). Responses of root system architecture to water stress at multiple levels: A meta-analysis of trials under controlled conditions. Frontiers in Plant Science, 13, 1085409.
  103. Kramer-Walter, K. R., & Laughlin, D. C. (2017). Root nutrient concentration and biomass allocation are more plastic than morphological traits in response to nutrient limitation. Plant and Soil, 416(1), 539-550.
  104. Kumar, M., Kumar Patel, M., Kumar, N., Bajpai, A. B., & Siddique, K. H. (2021). Metabolomics and molecular approaches reveal drought stress tolerance in plants. International Journal of Molecular Sciences, 22(17), 9108.
  105. Kurepa, J., & Smalle, J. (2025). The Evolution of Plant Hormones: From Metabolic Byproducts to Regulatory Hubs. International Journal of Molecular Sciences, 26(15), 7190.
  106. Lal, M. A., & Bhatla, S. C. (2023). Metabolism of storage carbohydrates. In Plant physiology, development and metabolism (pp. 231-259). Springer.
  107. Li, M., & Gao, F. (2025). Discussion on the Symbiotic Relationship between Legume Crops and Soil Microorganisms and Their Ecological Benefits. Legume Genomics and Genetics, 16.
  108. Li, S., Wang, J., Lu, S., Salmon, Y., Liu, P., & Guo, J. (2023). Trade-off between hydraulic safety and efficiency in plant xylem and its influencing factors. Forests, 14(9), 1817.
  109. Li, W., Zhang, Z., Chen, D., He, X., Huang, Y., Zhao, X., Sun, W., Liu, Z.-J., & Zou, S.-Q. (2026). Physiological and transcriptomic insights into the molecular mechanisms of salt stress adaptation in Gardenia jasminoides. BMC Plant Biology, 26(1), 203.
  110. Li, X., Guo, Y., Ling, Q., Guo, Z., Lei, Y., Feng, X., Wu, J., & Zhang, N. (2024). Advances in the structure, function, and regulatory mechanism of plant plasma membrane intrinsic proteins. Genes, 16(1), 10.
  111. Liang, X., Yu, S., Ju, Y., Wang, Y., & Yin, D. (2025). Multi-scale remote-sensing phenomics integrated with multi-omics: Advances in crop drought–heat stress tolerance mechanisms and perspectives for climate-smart agriculture. Plants, 14(18), 2829.
  112. Liang, X., Zhou, Y., Xu, W., & Liang, J. (2025). An intracellular CPK-ECA1 phosphoregulatory circuit couples calcium signatures to ABA homeostasis for plant osmosensivity. Science Advances, 11(40), eadz2428.
  113. Liu, H., Song, S., Zhang, H., Li, Y., Niu, L., Zhang, J., & Wang, W. (2022). Signaling transduction of ABA, ROS, and Ca2+ in plant stomatal closure in response to drought. International Journal of Molecular Sciences, 23(23), 14824.
  114. Liu, R., & Chen, Y. (2024). Mycorrhizal symbiosis: evolution, opportunities, challenges, and prospects. Arbuscular mycorrhizal fungi in sustainable agriculture: inoculum production and application, 1-35.
  115. Liu, X., Dong, X., & Leskovar, D. I. (2016). Ground penetrating radar for underground sensing in agriculture: a review. International Agrophysics, 30(4), 533-543.
  116. Loconsole, D., Elia, M., Conversa, G., De Lucia, B., Cristiano, G., & Elia, A. (2025). Soil Moisture Sensing Technologies: Principles, Applications, and Challenges in Agriculture. Agronomy, 15(12), 2788.
  117. Lucas, M., & Vetterlein, D. (2022). X-ray imaging of root–soil interactions. In X-Ray imaging of the soil porous architecture (pp. 129-157). Springer.
  118. Lynch, J. P. (2022). Harnessing root architecture to address global challenges. The Plant Journal, 109(2), 415-431.
  119. Ma, J., Wang, W., Yang, J., Qin, S., Yang, Y., Sun, C., Pei, G., Zeeshan, M., Liao, H., & Liu, L. (2022). Mycorrhizal symbiosis promotes the nutrient content accumulation and affects the root exudates in maize. BMC Plant Biology, 22(1), 64.
  120. Ma, L., Li, J., Li, J., Huo, Y., Yang, Y., Jiang, C., & Guo, Y. (2025). Plant salt-tolerance mechanisms: Classic signaling pathways, emerging frontiers, and future perspectives. Molecular Plant.
  121. MacAdam, J. W., & Nelson, C. J. (2020). Plant‐Water Relations in Forage Crops. Forages: The Science of Grassland Agriculture, 2, 113-126.
  122. Mao, H., Kumi, F., Li, Q., & Han, L. (2016). Combining X-ray computed tomography with relevant techniques for analyzing soil–root dynamics–an overview. Acta Agriculturae Scandinavica, Section B—Soil & Plant Science, 66(1), 1-19.
  123. Maqbool, S., Hassan, M. A., Xia, X., York, L. M., Rasheed, A., & He, Z. (2022). Root system architecture in cereals: progress, challenges and perspective. The Plant Journal, 110(1), 23-42.
  124. Marik, D., & Sadhukhan, A. (2025). Unearthing the hidden organ: vital role of the root in drought tolerance of plants. Plant Molecular Biology, 115(6), 1-35.
  125. Martins, S., Brito, C., Baltazar, M., Dinis, L.-T., & Pereira, S. (2026). Exploring the Role of Root Exudates in Shaping Plant–Soil–Microbe Interactions to Support Agroecosystem Resilience. Horticulturae, 12(1), 90.
  126. Mehra, P., Banda, J., Ogorek, L. L. P., Fusi, R., Castrillo, G., Colombi, T., Pandey, B. K., Sturrock, C. J., Wells, D. M., & Bennett, M. J. (2025). Root growth and development in “real life”: Advances and challenges in studying root–environment interactions. Annual review of plant biology, 76.
  127. Merino, V. M., Aguilar, R. I., Rivero, M. J., Ordóñez, I. P., Piña, L. F., López-Belchí, M. D., Schoebitz, M. I., Noriega, F. A., Pérez, C. I., & Cooke, A. S. (2024). Distribution of non-structural carbohydrates and root structure of Plantago lanceolata L. under different defoliation frequencies and intensities. Plants, 13(19), 2773.
  128. Michalak, A., Małas, K., Dąbrowska, K., Półrolniczak, K., Bronowska, L., Misiewicz, A., Maj, A., Stabrowska, M., Wnuk, I., & Kabała, K. (2026). Molecular Mechanisms and Experimental Strategies for Understanding Plant Drought Response. Plants, 15(1), 149.
  129. Milosavljević, A., Kostić Kravljanac, L., Kostić, I., Trailović, M., Blagojević, J., Petrović, S., Trkulja, N., & Nikolić, M. (2025). Silicon mitigates Macrophomina phaseolina root infection in sugar beet. 9th International Conference on Silicon in Agriculture (ICSA 2025). Belgrade, Serbia, 15-19/09/2025.,
  130. Misra, V., & Mall, A. K. (2025). Application of phytohormones exogenously to ameliorate sugarcane's response to water stress. Agricultural Ecology and Environment, 1(1).
  131. Mohammadi Alagoz, S., Zahra, N., Hajiaghaei Kamrani, M., Asgari Lajayer, B., Nobaharan, K., Astatkie, T., Siddique, K. H., & Farooq, M. (2023). Role of root hydraulics in plant drought tolerance. Journal of Plant Growth Regulation, 42(10), 6228-6243.
  132. Morisse, M., Wells, D. M., Millet, E. J., Lillemo, M., Fahrner, S., Cellini, F., Lootens, P., Muller, O., Herrera, J. M., & Bentley, A. R. (2022). A European perspective on opportunities and demands for field-based crop phenotyping. Field Crops Research, 276, 108371.
  133. Morota, G., Jarquin, D., Campbell, M. T., & Iwata, H. (2022). Statistical methods for the quantitative genetic analysis of high-throughput phenotyping data. In High-throughput plant phenotyping: Methods and protocols (pp. 269-296). Springer.
  134. Mpongwana, S., Manyevere, A., Mupangwa, J., & Mashamaite, C. V. (2024). Improving soil fertility through dual inoculation with arbuscular mycorrhizal fungi and Rhizobium on a eutric cambisol cultivated with forage legumes in a semi-arid region. Heliyon, 10(2).
  135. Mpongwana, S., Manyevere, A., Mupangwa, J., Mpendulo, C. T., & Mashamaite, C. V. (2023). Optimizing biomass yield of three herbaceous forage legumes through dual inoculation of Arbuscular Mycorrhizal Fungi and Rhizobia. South African Journal of Botany, 159, 61-71.
  136. Muhammad, A., Khan, Z. U., Khan, J., Mashori, A. S., Ali, A., Jabeen, N., Han, Z., & Li, F. (2025). A comprehensive review of crop stress detection: Destructive, non-destructive, and ML-based approaches. Frontiers in Plant Science, 16, 1638675.
  137. Muhammad, M., Wahab, A., Waheed, A., Hakeem, K. R., Mohamed, H. I., Basit, A., Toor, M. D., Liu, Y. H., Li, L., & Li, W. J. (2025). Navigating climate change: Exploring the dynamics between plant–soil microbiomes and their impact on plant growth and productivity. Global Change Biology, 31(2), e70057.
  138. Mukherjee, A., Dwivedi, S., Bhagavatula, L., & Datta, S. (2023). Integration of light and ABA signaling pathways to combat drought stress in plants. Plant Cell Reports, 42(5), 829-841.
  139. Naikwade, P. V. (2023). Plant responses to drought stress: Morphological, physiological, molecular approaches, and drought resistance. In Plant metabolites under environmental stress (pp. 149-183). Apple Academic Press.
  140. Nawaz, A. F., Gargiulo, S., Pichierri, A., & Casolo, V. (2025). Exploring the role of non-structural carbohydrates (NSCs) under abiotic stresses on woody plants: a comprehensive review. Plants, 14(3), 328.
  141. Nikolic, M., & Kirkby, E. (2025). Book of Abstracts/9 th International Conference on Silicon in Agriculture. 9 th International Conference on Silicon in Agriculture,
  142. Nuixe, M., Traoré, A. S., Blystone, S., Bonny, J.-M., Pagés, G., & Picon-Cochard, C. (2025). Use of a portable MRI to evaluate root structure-function relationships in water-use for several herbaceous species. Plant and Soil, 511(1), 483-499.
  143. Odone, A., Popovic, O., & Thorup-Kristensen, K. (2024). Deep roots: implications for nitrogen uptake and drought tolerance among winter wheat cultivars. Plant and Soil, 500(1), 13-32.
  144. Odone, A., & Thorup-Kristensen, K. (2025). Can rhizotron tube studies predict deep rooting in the field? A comparison of root phenotyping methods. Field Crops Research, 326, 109867.
  145. Paluch-Lubawa, E., & Polcyn, W. (2024). Tissue-specific accumulation of PIP aquaporins of a particular heteromeric composition is part of the maize response to mycorrhiza and drought. Scientific Reports, 14(1), 21712.
  146. Panchal, P., Preece, C., Peñuelas, J., & Giri, J. (2022). Soil carbon sequestration by root exudates. Trends in plant science, 27(8), 749-757.
  147. Pandey, J. K., & Biswal, B. (2024). Sugar homeostasis in leaves during senescence and abiotic stress: role of cell wall polysaccharides. Biologia, 79(8), 2347-2357.
  148. Pandey, R., Chinnusamy, V., Rathod, G., Paul, V., & Jain, N. (2017). Evaluation of root growth and architecture. Manual of ICAR Sponsored Training Programme on” Physiological Techniques to Analyze the Impact of Climate Change on Crop Plants”(16-25 Jan., 2017). Division of Plant Physiology, IARI, New Delhi. Pp, 95-102.
  149. Panigrahi, C. K., Jena, N., & Charitha, K. (2024). The role of phosphorus in root development and crop yield. Soil Fertility and Plant Nutrition, 21.
  150. Paul, M., Negrao, S., & Schulman, A. (2025). Barley under drought and vapour pressure deficit: Long-and short-term physiological and transcriptomic responses and recovery.
  151. Pereira, L. E. T., Herling, V. R., & Tech, A. R. B. (2022). Current scenario and perspectives for nitrogen fertilization strategies on tropical perennial grass pastures: a review. Agronomy, 12(9), 2079.
  152. Pflugfelder, D., Metzner, R., Van Dusschoten, D., Reichel, R., Jahnke, S., & Koller, R. (2017). Non-invasive imaging of plant roots in different soils using magnetic resonance imaging (MRI). Plant Methods, 13(1), 102.
  153. Picasso, V. D., Berti, M., Cassida, K., Collier, S., Fang, D., Finan, A., Krome, M., Hannaway, D., Lamp, W., & Stevens, A. W. (2022). Diverse perennial circular forage systems are needed to foster resilience, ecosystem services, and socioeconomic benefits in agricultural landscapes. Grassland Research, 1(2), 123-130.
  154. Piccini, C., Conti, V., Cantini, C., Parri, S., & Cai, G. (2025). Functional roles of osmotin, dehydrin and aquaporin in drought stress response: comparative analysis in tomato and olive. Plant Biosystems-An International Journal Dealing with all Aspects of Plant Biology, 159(6), 1377-1389.
  155. Portell, X., Pot, V., Ebrahimi, A., Monga, O., & Roose, T. (2022). Integrating X-ray CT Data into Models. In X-ray Imaging of the Soil Porous Architecture (pp. 183-222). Springer.
  156. Qin, Y., Yan, Y., Cheng, L., Lu, Y., Chen, J., Liu, F., & Tan, J. (2023). Arbuscular mycorrhizal fungi and rhizobium facilitate nitrogen and phosphate availability in soybean/maize intercropping systems. Journal of Soil Science and Plant Nutrition, 23(2), 2723-2731.
  157. Ragland, D. I. (2024). The Evaluation of Root System Architecture (RSA) As A New Breeding Target for Climate-Resilient Winter Wheat (Tritium aestivum L.) Virginia Tech].
  158. Rahman, M. M., Keya, S. S., Bulle, M., Ahsan, S., Rahman, M. A., Roni, M. S., Al Noor, M. M., & Hasan, M. (2025). Past trauma, better future: how stress memory shapes plant adaptation to drought. Functional Plant Biology, 52(6), FP24355.
  159. Ramachandran, P., Ramirez, A., & Dinneny, J. R. (2025). Rooting for survival: how plants tackle a challenging environment through a diversity of root forms and functions. Plant physiology, 197(1), kiae586.
  160. Rao, I. M. (2021). Digging deeper to define the physiological responses to environmental stress: The case of common bean and Brachiaria grasses. In Handbook of plant and crop physiology (pp. 1099-1140). CRC Press.
  161. Reeger, J. E. (2020). Rice Roots for Drought Tolerance: Functional Roles and Genetic Components of Root Anatomical and Architectural Traits in Oryza sativa The Pennsylvania State University].
  162. Renzetti, M., Funck, D., & Trovato, M. (2024). Proline and ROS: a unified mechanism in plant development and stress response? Plants, 14(1), 2.
  163. Rockwell, F., & Sage, R. F. (2022). Plants and water: the search for a comprehensive understanding. Annals of Botany, 130(3), i-viii.
  164. Ruckle, M. E., Meier, M. A., Frey, L., Eicke, S., Kölliker, R., Zeeman, S. C., & Studer, B. (2017). Diurnal leaf starch content: an orphan trait in forage legumes. Agronomy, 7(1), 16. Şahin, M., Bayram, G., Can, M., Ayan, İ., & Acar, Z. (2022). Perennıal Ryegrass (Lolium perenne L.) Oregon State University Corvallis, OR, USA].
  165. Sargar, P., Jwala, P., Thakur, N., Mehtre, S., Kalpande, H., Pawar, G., Chavan, S., & Habyarimana, E. (2025). Decoding Sorghum Root System Architecture for Resource Use Efficiency and Climate Resilience Under Multifactorial Stress Conditions. Physiologia Plantarum, 177(6), e70672.
  166. Sha, S., Cai, G., Liu, S., & Ahmed, M. A. (2024). Roots to the rescue: how plants harness hydraulic redistribution to survive drought across contrasting soil textures. Advanced Biotechnology, 2(4), 43. Sharipova, G., Ivanov, R., Veselov, D., Akhiyarova, G., Seldimirova, O., Galin, I., Fricke, W., Vysotskaya, L., &
  167. Kudoyarova, G. (2022). Effect of salinity on stomatal conductance, leaf hydraulic conductance, HvPIP2 aquaporin, and abscisic acid abundance in barley leaf cells. International Journal of Molecular Sciences, 23(22), 14282.
  168. Sharma, A., Hazarika, M., Heisnam, P., Pandey, H., Devadas, V., & Wangsu, M. (2024). Controlled environment ecosystem: A plant growth system to combat climate change through soilless culture. Crop Design, 3(1), 100044.
  169. Sharma, N., Farooq, F., Srivastava, H., Saini, H., Nanda, D., Mehta, S., MeCarty, J. S., Attri, M., & Kumar, N. (2025). Perennial Grass‐Legume Integration: A Sustainable Approach to Fodder Production. Grass and forage science, 80(3), e12727.
  170. Shoaib, M., Banerjee, B. P., Hayden, M., & Kant, S. (2022). Roots’ drought adaptive traits in crop improvement. Plants, 11(17), 2256.
  171. Singh, L. P., Gill, S. S., & Tuteja, N. (2011). Unraveling the role of fungal symbionts in plant abiotic stress tolerance. Plant signaling & behavior, 6(2), 175-191.
  172. Siyiannis, V. F., Protonotarios, V. E., Zechmann, B., Chorianopoulou, S. N., Müller, M., Hawkesford, M. J., & Bouranis, D. L. (2012). Comparative spatiotemporal analysis of root aerenchyma formation processes in maize due to sulphate, nitrate or phosphate deprivation. Protoplasma, 249(3), 671-686.
  173. Slooten, E., Raath-Krüger, M., Archibald, S., & Siebert, F. (2026). Subtropical grassland root-to-shoot ratios and plant organic carbon estimates: why global averages fall short. Annals of Botany, mcag027. Smethurst, C. F. (2005). Waterlogging and salinity tolerance in lucerne (Medicago sativa) University of Tasmania (Hobert).
  174. Stanciu, A.-Ș. (2025). Sustainable forage yield improvement through perennial grass–legume intercropping in degraded grasslands. Turkish Journal of Agriculture and Forestry, 49(2), 215-241.
  175. Striker, G. G. (2023). An overview of oxygen transport in plants: diffusion and convection. Plant Biology, 25(6), 842-847.
  176. Striker, G. G., Yamauchi, T., & Mollard, F. P. (2025). Maize physiology under waterlogging: shoot and root adaptive responses to mitigate low oxygen stress. Journal of Experimental Botany, eraf424.
  177. Sun, X., Miao, Q., Gu, Y., Yang, L., & Wang, P. (2025). Research on the physiological mechanisms of nitrogen in alleviating plant drought tolerance. Plants, 14(18), 2928.
  178. Sun, Y., Robert, C. A., & Thakur, M. P. (2024). Drought intensity and duration effects on morphological root traits vary across trait type and plant functional groups: a meta-analysis. BMC ecology and evolution, 24(1), 92.
  179. Suura, S. R. (2025). Integrating artificial intelligence, machine learning, and big data with genetic testing and genomic medicine to enable earlier, personalized health interventions. Deep Science Publishing. https://doi.org/10.70593/978-93-49307-76-6
  180. Svane, S. F., Jensen, C. S., & Thorup-Kristensen, K. (2019). Construction of a large-scale semi-field facility to study genotypic differences in deep root growth and resources acquisition. Plant Methods, 15(1), 26. Takahashi, H., & Nakazono, M. (2024). Cavity tissue for the internal aeration in plants. In Responses of Plants to Soil Flooding (pp. 105-117). Springer.
  181. Tang, Y., Liu, X., Lian, J., Cheng, X., Wang, G. G., & Zhang, J. (2023). Soil depth can modify the contribution of root system architecture to the root decomposition rate. Forests, 14(6), 1092.
  182. Tao, H., Xu, S., Tian, Y., Li, Z., Ge, Y., Zhang, J., Wang, Y., Zhou, G., Deng, X., & Zhang, Z. (2022). Proximal and remote sensing in plant phenomics: 20 years of progress, challenges, and perspectives. Plant Communications, 3(6).
  183. Tariq, A., Graciano, C., Sardans, J., Zeng, F., Hughes, A. C., Ahmed, Z., Ullah, A., Ali, S., Gao, Y., & Peñuelas, J. (2024). Plant root mechanisms and their effects on carbon and nutrient accumulation in desert ecosystems under changes in land use and climate. New Phytologist, 242(3), 916-934.
  184. Tomasella, M., Häberle, K.-H., Nardini, A., Hesse, B., Machlet, A., & Matyssek, R. (2017). Post-drought hydraulic recovery is accompanied by non-structural carbohydrate depletion in the stem wood of Norway spruce saplings. Scientific Reports, 7(1), 14308.
  185. Tylová, E., Pecková, E., Blascheová, Z., & Soukup, A. (2017). Casparian bands and suberin lamellae in exodermis of lateral roots: an important trait of roots system response to abiotic stress factors. Annals of Botany, 120(1), 71-85.
  186. Undersander, D. J., & Sharpe, P. (2025). Forage plant structure, function, nutrition and growth. In Horse pasture management (pp. 1-10). Elsevier.
  187. Urfan, M., Sharma, S., Hakla, H. R., Rajput, P., Andotra, S., Lehana, P. K., Bhardwaj, R., Khan, M. S., Das, R., & Kumar, S. (2022). Recent trends in root phenomics of plant systems with available methods-discrepancies and consonances. Physiology and Molecular biology of Plants, 28(6), 1311-1321.
  188. Villarraga-Gómez, H., Brackman, P., Ziabari, A., Rahman, O., Snow, Z., Shahani, R., Bugelnig, K., Andreyev, A., Trenikhina, Y., & Johnson, N. (2025). Multiscale characterization of additive manufacturing components with computed tomography, 3D X-ray microscopy, and deep learning. Journal of Nondestructive Evaluation, 44(3), 110.
  189. Voothuluru, P., Wu, Y., & Sharp, R. E. (2024). Not so hidden anymore: advances and challenges in understanding root growth under water deficits. The Plant Cell, 36(5), 1377-1409.
  190. Wahab, A., Muhammad, M., Munir, A., Abdi, G., Zaman, W., Ayaz, A., Khizar, C., & Reddy, S. P. P. (2023). Role of arbuscular mycorrhizal fungi in regulating growth, enhancing productivity, and potentially influencing ecosystems under abiotic and biotic stresses. Plants, 12(17), 3102.
  191. Wang, G., Guo, H., Zhu, Q., An, D., Song, Z., & Ma, L. (2025). Optimizing Subsurface Drainage Pipe Layout Parameters in Southern Xinjiang’s Saline–Alkali Soils: Impacts on Soil Salinity Dynamics and Oil Sunflower Growth Performance. Sustainability, 17(11), 4797.
  192. Wang, L., Dai, Y., Zhang, J., Meng, P., & Wan, X. (2022). Xylem structure and hydraulic characteristics of deep roots, shallow roots and branches of walnut under seasonal drought. BMC Plant Biology, 22(1), 440.
  193. Wang, W., Wang, X., Lv, Z., Khanzada, A., Huang, M., Cai, J., Zhou, Q., Huo, Z., & Jiang, D. (2022). Effects of cold and salicylic acid priming on free proline and sucrose accumulation in winter wheat under freezing stress. Journal of Plant Growth Regulation, 41(6), 2171-2184.
  194. Wang, X., An, Y., Wang, L., Zhang, M., Xing, X., Tong, S., & Wu, H. (2026). Leaf and root traits codetermine biomass production and allocation patterns in agriculturally managed wetlands in a semi-arid region. Agriculture, Ecosystems & Environment, 400, 110229.
  195. Wang, X., Hu, G., Zhang, Q., Quan, X., Zhang, H., Aubrey, D. P., & Wang, C. (2025). Annual Tree Biomass Increment Is Positively Related to Nonstructural Carbohydrate Pool Size and Depletion: Evidence for Carbon Limitation? Forests, 16(4), 619.
  196. Wang, Y., Jia, X., An, S., Yin, W., Huang, J., & Jiang, X. (2024). Nanozyme‐based regulation of cellular metabolism and their applications. Advanced Materials, 36(10), 2301810.
  197. Wang, Z., Li, Z., Zhang, Y., Liao, J., Guan, K., Zhai, J., Meng, P., Tang, X., Dong, T., & Song, Y. (2024). Root hair developmental regulators orchestrate drought triggered microbiome changes and the interaction with beneficial Rhizobiaceae. Nature Communications, 15(1), 10068.
  198. Wang, Z., & Wang, C. (2025). Dynamics of nonstructural carbohydrates during drought and subsequent recovery: a global meta-analysis. Agricultural and Forest Meteorology, 363, 110429.
  199. Wiley, E., & Estes, A. G. (2023). The impacts of late season defoliation and winter flooding on spring leaf flush and carbohydrate remobilization in pin oak. Trees, 37(4), 1069-1080.
  200. Wróbel, B., Zielewicz, W., & Paszkiewicz-Jasińska, A. (2025). Improving forage quality from permanent grasslands to enhance ruminant productivity. Agriculture, 15(13), 1438.
  201. Wu, Q., Wu, J., Hu, P., Zhang, W., Ma, Y., Yu, K., Guo, Y., Cao, J., Li, H., & Li, B. (2023). Quantification of the three-dimensional root system architecture using an automated rotating imaging system. Plant Methods, 19(1), 11.
  202. Xiao, T., Li, P., Vanapalli, S. K., Zheng, Z., Ren, G., Zhai, H., Han, K., & Wang, J. (2025). Effects of root exudates analogues of Robinia pseudoacacia L. on water retention capacity and capillary rise in loess: An experimental study. Journal of Hydrology, 134417.
  203. Xie, Q., Hou, H., Yan, P., Zhang, H., Lv, Y., Li, X., Chen, L., Pang, D., Hu, Y., & Ni, X. (2022). Programmed cell death associated with the formation of schizo-lysigenous aerenchyma in Nelumbo nucifera root. Frontiers in Plant Science, 13, 968841.
  204. Yang, M., Gao, X., Wang, S., & Zhao, X. (2022). Quantifying the importance of deep root water uptake for apple trees’ hydrological and physiological performance in drylands. Journal of Hydrology, 606, 127471.
  205. Yang, Q., Deng, X., Liu, T., Qian, J., Zhang, P., Zhu, E., Wang, J., Zhu, X., Kudoyarova, G., & Zhao, J. (2024). Abscisic acid root-to-shoot translocation by transporter AtABCG25 mediates stomatal movements in Arabidopsis. Plant physiology, 195(1), 671-684.
  206. Yetgin, A. (2024). Exploring the dynamic nature of root plasticity and morphology in the face of changing environments. Ecological Frontiers, 44(1), 112-119.
  207. Yi, K., Li, X., Chen, D., Yang, S., Liu, Y., Tang, X., Ling, G., & Zhao, Z. (2022). Shallower root spatial distribution induced by phosphorus deficiency contributes to topsoil foraging and low phosphorus adaption in sugarcane (Saccharum officinarum L.). Frontiers in Plant Science, 12, 797635.
  208. York, L. M., Slack, S., Bennett, M. J., & Foulkes, M. J. (2018). Wheat shovelomics I: a field phenotyping approach for characterising the structure and function of root systems in tillering species. bioRxiv, 280875.
  209. Zahedi, S. M., Karimi, M., Venditti, A., Zahra, N., Siddique, K. H., & Farooq, M. (2025). Plant adaptation to drought stress: the role of anatomical and morphological characteristics in maintaining the water status. Journal of Soil Science and Plant Nutrition, 25(1), 409-427.
  210. Zaman, F., Ali, A., Khattak, W., Khan, H., Anwar, Z., Rahimi, M., & Chen, F.-S. (2025). Diversity of arbuscular mycorrhizal fungi (AMF) and its and role in nutrient recycling and availability in subtropical forest ecosystem of China: A review. Eurasian Soil Science, 58(11), 145.
  211. Zhang, M., Pan, M., Li, H., Liu, B., Qiao, S., Ma, C., Mu, H., Zhao, W., & Guo, J. (2025). Plant survival strategies under heterogeneous salt stress: Remodeling of root architecture, ion dynamic balance, and coordination of metabolic homeostasis. Plant and Soil, 1-40.
  212. Zhang, W.-P., Surigaoge, S., Yang, H., Yu, R.-P., Wu, J.-P., Xing, Y., Chen, Y., & Li, L. (2024). Diversified cropping systems with complementary root growth strategies improve crop adaptation to and remediation of hostile soils. Plant and Soil, 502(1), 7-30.
  213. Zhang, Y., Chen, X., Geng, S., & Zhang, X. (2025). A review of soil waterlogging impacts, mechanisms, and adaptive strategies. Frontiers in Plant Science, 16, 1545912.
  214. Zhang, Z., Liu, X., Wang, Y., Zhou, J., Wan, Z., Zhang, X., Wang, J., Si, B., Luo, L., & Xu, W. (2025). Genome-Wide Identification of the ABF/AREB/ABI5 Gene Family in Ziziphus jujuba cv. Dongzao and Analysis of Its Response to Drought Stress. Genes, 16(7), 785.
  215. Zhiyong, Z., Baomin, F., Chao, S., Xiaoxian, Z., Qingwen, Z., & Bing, Y. (2022). Advances in root system architecture: functionality, plasticity, and research methods. Journal of Resources and Ecology, 14(1), 15-24.
  216. Zhou, Y., Case, M. F., & Staver, A. C. (2025). Root trait (multi) functionality in savanna trees: progress and challenges. Journal of Ecology, 113(5), 1022-1041.
  217. Zhou, Y., Wang, C., Tang, Q., Wang, J., & Li, M.-H. (2024). Morphological responses of root hairs to changes in soil and climate depend on plant life form. Frontiers in Forests and Global Change, 7, 1324405.
  218. Ziabari, A., Bedhief, M. H., Rahman, O., Venkatakrishnan, S., Brackman, P., & Katuch, P. (2025). Combining deep learning and scatterControl for high-throughput X-ray CT based non-destructive characterization of large-scale casted metallic components. Journal of Nondestructive Evaluation, 44(4), 148.