Reconsidering how to dry orthodox seeds for improved ex situ conservation outcomes

• Baum, K. J., Norton, S. L., Keeble-Gagnère, G., Hayden, M. (2025). Safeguarding seed longevity under genebank storage:Evidence based viability monitoring intervals for six tropical species at the Australian Grains Genebank (AGG). Plant Genetic Resources: Characterization and Utilization, 1-11. https://doi.org/10.1017/S1479262125100488 • BGCI. (2025). https://www.bgci.org/our-work/saving-plants/seed-conservation/. Accessed 16-Nov-25. • Bray, E. A. (1997). Plant responses to water deficit. Trends in Plant Science, 2, 48-54. • Commander, L. E. (Ed.). (2021). Florabank Guidelines – best practice guidelines for native seed collection and use, 2nd edition, Florabank Consortium, Australia. • CPC. (2019). CPC Best plant conservation practices to support species survival in the wild. Center for Plant Conservation, Escondido, CA. • Cromarty, A. S. (1982). Techniques for drying seeds. In Seed management techniques for genebanks, Eds Dickie, J.B., Linington, S. and Williams, J.T. pp. 88-125. International Board for Plant Genetic Resources, Rome, Italy. • Cromarty, A. S., Ellis, R. H., & Roberts, E. H. (1982). The Design of Seed Storage Facilities for Genetic Conservation, International Board for Plant Genetic Resources, Rome. • Dasgupta, J., Bewley, J. D., & Yeung, E. C. (1982). Desiccation-tolerant and desiccation-intolerant stages during the development and germination of Phaseolus vulgaris seeds. Journal of Experimental Botany, 33, 1045-1057. • Demir, I., & Ellis, R. H. (1992). Changes in seed quality during seed development and maturation in tomato. Seed Science Research 2, no. 2: 81-87. doi:10.1017/S0960258500001173 • De Vitis, M., Hay, F. R., Dickie, J. B., Trivedi, C., Choi, J., & Fiegener, R. (2020). Seed storage: maintaining seed viability and vigor for restoration use. Restoration Ecology, 28, S249-S255. • Ellis, R. H. (2022). Seed ageing, survival and the improved seed viability equation; forty years on. Seed Science and Technology, 50, no. 1, Suppl.: 1-20. • Ellis, R. H., & Hong, T. D. (2006). Temperature sensitivity of the low-moisture-content limit to negative seed longevity-moisture content relations in hermetic storage. Annals of Botany, 97, 785-791. • Ellis, R. H., & Roberts, E. H. (1980). Improved equations for the prediction of seed longevity. Annals of Botany, 45, 13-30. • Ellis, R. H., Hong, T. D., & Roberts, E. H. (1989). A comparison of the low-moisture-content limit to the logarithmic relation between seed moisture and longevity in twelve species. Annals of Botany, 63, 601-611. • Ellis, R. H., Nasehzadeh, M., Hanson, J., & Woldemariam, Y. (2018). Medium-term seed storage of 50 genera of forage legumes and evidence-based genebank monitoring intervals. Genetic Resources and Crop Evolution, 65, 607-623. • Ellis, R. H., Nasehzadeh, M., Hanson, J., Ndiwa, N., & Woldemariam, Y. (2019). Medium-term seed storage of diverse genera of forage grasses, evidence-based genebank monitoring intervals, and regeneration standards. Genetic Resources and Crop Evolution, 66, 723-734. • FAO. (2014). Genebank Standards for Plant Genetic Resources for Food and Agriculture. Rev. ed. Rome. • FAO. (2022). Practical guide for the application of the Genebank Standards for Plant Genetic Resources for Food and Agriculture: Conservation of orthodox seeds in seed genebanks. Commission on Genetic Resources for Food and Agriculture. Rome. https://doi.org/10.4060/cc0021en • FAO/IPGRI. (1994). Genebank Standards. Food and Agriculture Organization of the United Nations, Rome, International Plant Genetic Resources Institute, Rome. • FAO. (2025). The Third Report on the State of the World's Plant Genetic Resources for Food and Agriculture. FAO Commission on Genetic Resources for Food and Agriculture Assessments, Rome. • Fernández Marín, B., Buchner, O., Kastberger, G., Piombino, F., García Plazaola, J. I., & Kranner, I. (2019). Non-invasive diagnosis of viability in seeds and lichens by infrared thermography under controlled environmental conditions. Plant Methods, 15, 147. • Goodale, U. M., Antonelli, A., Nelson, C. R., & Chau, M. M. (2023). Seed banks needed to restore ecosystems. Science, 379(6628), 147. • Guzzon, F., Costich, D. E., Afzal, I., Barboza Barquero, L., Monge Vargas, A. A., Vargas Ramírez, E., Bello, P., Dahal, P., Sánchez Cano, C., Zavala Espinosa, C., Imran, S., Patolo, S., Tukia, T. N., Van Asbrouck, J., Nabubuniyaka-Young, E., Gianella, M., & Bradford, K. J. (2024). Applications of dry chain technology to maintain high seed viability in tropical climates. PeerJ, 12, e18146 https://doi.org/10.7717/peerj.18146 • Hay, F. R. (1997). Seed maturity and the effects of different drying conditions on seed longevity in four wild plant species collected in the Sahel region of West Africa. In The Development of Seed Longevity in Wild Plant Species, pp. 179-225, PhD thesis, King’s College, University of London. • Hay, F. R., & Probert, R. J. (2011). Collecting and handling seeds in the field. In Collecting Plant Genetic Diversity: Technical Guidelines – 2011 Update. • Hay, F. R., de Guzman, F., Ellis, D., Makahiya, H., Borromeo, T., & Sackville Hamilton, N. R. 2013. Viability of Oryza sativa (L.) seeds stored under genebank conditions for up to 30 years. Genetic Resources and Crop Evolution, 60, 275–296. • Hay, F. R., Whitehouse, K. J., Ellis, R. H., Sackville Hamilton, N. R., Lusty, C., Ndjiondjop, M. N., Tia, D., Wenzl, P., Santos, L. G., Yazbek, M., Azevedo, V. C. R., Peerzada, O. H., Abberton, M., Oyatomi, O., de Guzman, F., Capilit, G., Muchugi, A., & Kinyanjui, Z. 2021. CGIAR genebank viability data reveal inconsistencies in seed collection management. Global Food Security, 30, 100557. • HSBP. (Hawaiʻi Seed Bank Partnership) (2025). Best Management Practices—Seed Drying. Available online at https://laukahi.org/wp-content/uploads/2025/06/Seed-Drying-Protocol-Final_18Jun2025.pdf and https://seedsofhawaii.org/wp-content/uploads/2025/06/Seed-Drying-Protocol-Final_18Jun2025_NK.pdf • IBPGR. (1976). Report of IBPGR working group on engineering, design and cost aspects of long-term seed storage facilities. International Board for Plant Genetic Resources, Rome. • IBPGR. (1982). IBPGR ad hoc advisory committee on seed storage: report of the first meeting. International Board for Plant Genetic Resources, Rome. • IBPGR. (1985). IBPGR advisory committee on seed storage: report of the third meeting. International Board for Plant Genetic Resources, Rome. • Jawad, R., Hay, F. R., Yazbek, M., & Ellis, R. H. (2025). Seed collection and processing practices affect subsequent seed storage longevity in durum wheat and wild relatives. Seed Science and Technology, 53, 277-310. • Kameswara Roa, N., Dulloo, M. E., & Engels, J. M. M. (2017). A review of factors that influence the production of quality seed for long-term conservation in genebanks. Genetic Resources and Crop Evolution 64, no. 5: 1061-1074. • Kermode, A. R., Bewley, J. D., Dasgupta, J., & Misra, S. (1986). The transition from seed development to germination: a key role for desiccation? HortScience, 21, 1113-1118. • Leprince, O., Pellizzaro, A., Berriri, S., & Buitink, J. (2017). Late seed maturation: drying without dying. Journal of Experimental Botany 68, no. 4: 827–841. • National Academies of Sciences, Engineering, and Medicine. (2023). An Assessment of Native Seed Needs and the Capacity for Their Supply: Final Report. Washington, DC: The National Academies Press. • Pieta Filho, C., & Ellis, R. H. (1991). The development of seed quality in spring barley in four environments. I. Germination and longevity. Seed Science Research, 1, 163-177. • Probert, R., Adams, J., Coneybeer, J., Crawford, A., & Hay, F. (2007). Seed quality for conservation is critically affected by pre-storage factors. Australian Journal of Botany, 55, 326-335. • Rao, N. K., Hanson, J., Dulloo, M. E., Ghosh, K., Nowell, D., & Larinde, M. (2006). Manual of seed handling in genebanks. Handbooks for Genebanks No. 8. Bioversity International, Rome, Italy. • Rezaei, S. (2023). Seed quality and moisture relations. PhD Thesis, Department of Agroecology, Aarhus University, Denmark. • Roberts, E. H., & Ellis, R. H. (1989). Water and seed survival. Annals of Botany, 63, 39-52. • Royal Botanic Gardens Kew. (2022a). The Millennium Seed Bank Partnership (MSBP) Seed Conservation Standards for ‘MSBP Collections’. https://brahmsonline.kew.org/Content/Projects/msbp/resources/Training/MSBP-Seed-Conservation-Standards.pdf • Royal Botanic Gardens Kew. (2022b). Post-harvest handling of seed collections. Technical Information Sheet 04. Millennium Seed Bank Partnership, Royal Botanic Gardens Kew. https://brahmsonline.kew.org/Content/Projects/msbp/resources/Training/04-Post-harvest-handling.pdf • Sackville-Hamilton, N. R., and Chorlton, K. H. 1997. Regeneration of accessions in seed collections: a decision guide. Handbooks for Genebanks No. 5. International Plant Genetic Resources Institute, Rome, Italy. • Sallon, S., Solowey, E., Gostel, M. R., Egli, M., Flematti, G. R., Bohman, B., Schaeffer, P., Adam, P., & Weeks, A. 2024. Characterization and analysis of a Commiphora species germinated from an ancient seed suggests a possible connection to a species mentioned in the Bible. Communications Biology 7: 1109. • Salvador, O. F., Ellis, R. H., Oyatomi, O. O., Abberton, M. T., & Hay, F. R. (2025). Two-step drying of soya bean seed germplasm often improves subsequent storage longevity. Genetic Resources and Crop Evolution, 72, 9577-9590. • SER/INSR/RBG Kew. (2023). Seed Information Database (SID). Society for Ecological Restoration, International Network for Seed Based Restoration and Royal Botanic Gardens Kew. Available from: https://ser-sid.org/ (February 2023) • Sinniah, U. R., Ellis, R. H., & John, P. (1998). Irrigation and seed quality development in rapid-cycling Brassica: soluble carbohydrates and heat-stable proteins. Annals of Botany, 82, 647-655. • Vertucci, C. W., & Roos, E. E. (1993). Theoretical basis of protocols for seed storage II. The influence of temperature on optimal moisture levels. Seed Science Research, 3, 201-213. • Walters, C. T., Wheeler, L. M., & Grotenhuis, J. M. (2005). Longevity of seeds stored in a genebank: species characteristics. Seed Science Research, 15, 1–20. https://doi.org/10.1079/SSR2004195 • Whitehouse, K. J., Hay, F. R., & Ellis, R. H. 2015. Increases in the longevity of desiccation-phase developing rice seeds: response to high-temperature drying depends on harvest moisture content. Annals of Botany 116, no. 2: 247–259. • Whitehouse, K. J., Hay, F. R., & Ellis, R. H. 2017. High-temperature stress during drying improves subsequent rice (Oryza sativa L.) seed longevity. Seed Science Research 27, no. 4: 281-291. • Whitehouse, K. J., Hay, F.R., & Ellis, R. H. 2018. Improvement in rice seed storage longevity from high-temperature drying is a consistent positive function of harvest moisture content above a critical value. Seed Science Research, 28, 332-339. • Whitehouse, K. J., Owoborode, O. F., Adebayo, O. O., Oyatomi, O. A., Olaniyan, A. B., Abberton, M. T., & Hay, F. R. (2018). Further evidence that the genebank standards for drying orthodox seeds may not be optimal for subsequent seed longevity. Biopreservation and Biobanking, 16, 327-336. • Whitehouse, K. J., Hay, F. R., & Lusty, C. (2020). Why seed physiology is important for genebanking. Plants, 9, 584.