A significantly enhanced role for plant genetic resource centres in linking in situ and ex situ conservation to aid user germplasm access

REVIEW AND POSITION PAPER Genetic Resources (2025), (S2), 203–223 DOI: 10.46265/genresj.UNVV5571 https://www.genresj.org ISSN: 2708-3764 A significantly enhanced role for plant genetic resource centres in linking in situ and ex situ conservation to aid user germplasm access Nigel Maxteda, Anne-Franc¸oise Adam-Blondonb, Catherine Hazel Aguilarc, Ana Maria Baratad, B´ ela Barthae, Riccardo Boccif, Domenica De Paolag, Heli Susanna Fitzgeraldh, Louis John Frestai, Pietro Fusanij, Giovanni Giulianok, Filippo Guzzonl, Philipp Holzherre, Vojtech Holubecm, Jos´ e Mar´ ıa Iriondo Alegr´ ıan, Juozas Labokaso, Lorenzo Maggionil, Joana Magos Brehma,d, Anna Palm´ ep, Jade Phillipsa, Jaime Prohensq, Lorenzo Raggir, Parthenopi Rallis, Dainis Run¸g¸ist, Karuine Sarikyanu, Jelka ˇ Suˇ star Vozliˇ cv, Imke Thormannwand Goran Zduni´ cx aSchool of Biosciences, University of Birmingham, B15 2TT, Birmingham, United Kingdom bInstitut national de recherche pour l’agriculture, l’alimentation et l’environnement, 78026 Versailles, France cLeibniz Institute of Plant Genetics and Crop Plant Research (IPK) - Gatersleben, 06466 Seeland, Germany dBanco Portuguˆ es de Germoplasma Vegetal, Instituto Nacional de Investigac¸˜ ao Agr´ aria e Veterin´ aria I.P. (BPGV-INIAV), Braga, 4700-859, Portugal eProSpecieRara, Unter Br¨ uglingen 6, Basel, CH-4052, Switzerland fRete Semi Rurali, via di Casignano 25, Scandicci (FI), 50018, Italy gCNR-IBBR Institute of Biosciences and Bioresources, National Research Council of Italy, Bari, Italy hFinnish Museum of Natural History, University of Helsinki, Helsinki, FI-00014, Finland iPlant Protection Directorate, Rural Affairs Department, Ministry for Agriculture, Fisheries, Food and Animal Rights, LJA1910 Lija, Malta jCouncil for Research in Agriculture and Economics, Research Centre for Forestry and Wood (CREA-FL), Trento, 38100, Italy kENEA, Centro Ricerche Casaccia, Via Anguillarese 301, Rome, 00123, Italy lECPGR Secretariat, Alliance of Bioversity International and the International Center for Tropical Agriculture (CIAT), Via di San Domenico, 1, Rome, 00153, Italy mCzech Agrifood Research Centre, Drnovska 507, 16100, Praha 6 -Ruzynˇ e, Czech Republic nInstituto de Investigaci´ on en Cambio Global (IICG-URJC), Universidad Rey Juan Carlos, c/ Tulipan s/n, M´ ostoles, Madrid, 28933, Spain oState Scientific Research Institute Nature Research Centre, Akademijos g. 2, 08412, Vilnius, Lithuania pNordic Genetic Resource Center (NordGen), Box 162, 234 23, Lomma, Sweden qUniversitat Polit` ecnica de Val` encia, Camino de Vera 14, ES46022, Valencia, Spain rDipartimento di Scienze Agrarie, Alimentari e Ambientali (DSA3), Universita `degli Studi di Perugia, Borgo XX Giugno 74, 06121, Perugia, Italy sHellenic Agricultural Organization - DIMITRA (ELGO - DIMITRA), Institute of Plant Breeding & Genetic Resources, PO Box 60458, 570 01 Thermi – Thessaloniki, Greece tGenetic Resource Centre, Latvian State Forest Research Institute (LSFRI) Silava, LV-2169 Salaspils, Latvia uScientific Center of Vegetable and Industrial Crops, Ministry of Economy, Ararat Marz, Darakert Community, 808 Ararat region, Armenia vAgricultural Institute of Slovenia, Hacquetova ulica 17, 1000 Ljubljana, Slovenia wFederal Office for Agriculture and Food, Information and Coordination Centre for Biological Diversity (IBV), 53179 Bonn, Germany xInstitute for Adriatic Crops and Karst Reclamation, 21000 Split, Croatia Citation: Maxted, N., Adam-Blondon, A., Aguilar, C. H., Barata, A. M., Bartha, B., Bocci, R., De Paola, D., Fitzgerald, H. S., Fresta, L. J., Fusani, P., Giuliano, G., Guzzon, F., Holzherr, P., Holubec, V., Iriondo Alegr´ ıa, J. M., Labokas, J., Maggioni, L., Magos Brehm, J., Palm´ e, A., Phillips, J., Prohens, J., Raggi, L., Ralli, P., Run¸g¸is, D., Sarikyan, K., ˇ Suˇ star Vozliˇ c, J., Thormann, I., Zduni´ c, G. (2025). A significantly enhanced role for plant genetic resource centres in linking in situ and ex situ conservation to aid user germplasm access. Genetic Resources (S2), 204–223. doi: 10.46265/genresj.UNVV5571. © Copyright 2025 the Authors. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Received: 03.04.2025 Accepted: 27.04.2025 Published online: 09.05.2025 Abstract: Plant genetic resources (PGR) serve as the cornerstone for global varietal enhancement and food security. However, these resources face significant threats, including diversity erosion and extinction, are often inadequately conserved, and frequently remain inaccessible for practical use. Traditionally, PGR have been primarily conserved through population seed samples stored ex situ in genebanks. In contrast, complementary in situ techniques – whether involving crop wild relatives (CWR) in genetic reserves or crop landraces (LR) on-farm – have largely remained experimental. The demand from breeders for a broader diversity is driving a more integrated approach that combines ex situ and in situ methods. This paper posits that such an integrated strategy would be mutually advantageous for PGR, biodiversity, and farmer-based conservation communities. As a foundation for future PGR science, we propose the three ‘Principles of PGR Conservation and Use Congruence’ and outline the practical processes involved in in situ and on-farm conservation. We also review the challenges associated with integrating ex situ and in situ conservation, specifically addressing how collaborative resource management can be established, how potential resource users can access in situ and on-farm conserved PGR, how to promote user access to in situ conserved populations, and the progress made thus far in integrating in situ and ex situ efforts. While it is acknowledged that full integration may be unrealistic without adequate resources for Genetic Resource Centres and the rectification of skill gaps, the potential to significantly enhance the long-term, sustainable conservation of PGR diversity holds profound existential benefits for humanity in the 21st century. Keywords: crop wild relatives, ex situ, genebank, genetic reserves, in situ, landraces, on-farm conservation, integrated conservation Citation: Maxted, N., Adam-Blondon, A., Aguilar, C. H., Barata, A. M., Bartha, B., Bocci, R., De Paola, D., Fitzgerald, H. S., Fresta, L. J., Fusani, P., Giuliano, G., Guzzon, F., Holzherr, P., Holubec, V., Iriondo Alegría, J. M., Labokas, J., Maggioni, L., Magos Brehm, J., Palmé, A., Phillips, J., Prohens, J., Raggi, L., Ralli, P., Rungis, D., Sarikyan, K., Šuštar Vozlič, J., Thormann, I., Zdunić, G. (2025). A significantly enhanced role for plant genetic resource centres in linking in situ and ex situ conservation to aid user germplasm access. Genetic Resources (S2), 203–222. doi: 10.46265/genresj.UNVV5571. © Copyright 2025 the Authors. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 204Maxted et al Genetic Resources (2025), (S2), 203–223 Genetic Resources (2025), (S2), 203–223 Plant GRC roles in linking in situ and ex situ conservation 205 Introduction Plant genetic resources (PGR) conservation is unique among conservation methods as it aims to preserve biodiversity while also utilizing conserved resources (Maxted et al,1997a). This process involves several steps: identifying genetic diversity across plant species, prioritizing target taxa, planning and implementing conservation actions, and characterizing, evaluating and utilizing resources by farmers, breeders or researchers. Clarity and expediency in this model’s application are essential for global, regional, national and local initiatives focused on food security, poverty reduction, and enhancing human well-being, thereby supporting many UN Sustainable Development Goals (UN, 2015). PGR conservation employs two main strategies: in situ, where resources are conserved in their natural habitats, and ex situ, where resources are relocated to safer environments for conservation and accessibility (see definitions in Supplemental Table 1). It is widely accepted that in situ and ex situ actions should complement each other, enhancing overall conservation effectiveness (FAO,1996). Historically, formal PGR conservation and germplasm application for orthodox-seeded species have relied heavily on ex situ seed storage in genebanks and, latterly, cryogenic preservation, with field genebanks and tissue culture techniques primarily used for recalcitrant-seeded species and clonally propagated crops. Genebanks can secure long-term viability at low cost and have successfully made this diversity available to plant breeders and researchers (FAO,1998,2011). However, ex situ approaches alone do not fully address the growing demand for broader diversity in a rapidly changing environment. The science of in situ and on-farm PGR conservation has advanced significantly, with refined techniques and a solid evidence base (Maxted et al,1997c,2002,2020; Brush,2000;Eyzaguirre and Linares,2004;Heywood and Dulloo,2005;Jarvis et al,2007,2016;Iriondo et al,2008,2021;Vetel¨ ainen et al,2009;Hunter and Heywood,2011;Hunter et al,2017). Initially, in situ and ex situ techniques were viewed as independent, even competitive (Ford-Lloyd and Maxted,1993), but the case for their complementarity is now widely accepted, though their practical integration remains incomplete (Maxted et al,1997a,2020;van Hintum et al,2021). Lack of integration limits conservation effectiveness, resulting in unconserved resources being unavailable to users and preventing their potential utilization. The challenge of increasing food production to feed a growing human population while mitigating climate change impacts on agriculture is escalating for the PGR and breeding communities FAO (2010,2012). Lack of breadth and access to conserved genetic diversity is now a barrier to crop improvement (McCouch et al, 2013;IPCC,2014;Dempewolf et al,2017;Zhang et al, 2017). There is an opportunity to better serve farmers and breeders by integrating in situ conservation, genebanks, and germplasm use into a cohesive continuum that could significantly enhance breadth and access to diversity for users (Maxted and Brehm,2023). Failure to integrate these activities reduces the potential role of genebanks in leading PGR conservation and meeting user demands. Maxted et al (2016) suggested that expanding the role of genebanks to include both ex situ and in situ conservation was logical and required change to the PGR paradigm and would warrant their renaming as Genetic Resource Centres (GRC), as the term ‘genebank’ implies a more restrictive focus. To explore this enhanced GRC role, a questionnaire was prepared in 2024 on European genebank activities for the Horizon Europe project ‘Promoting a Plant Genetic Resource Community for Europe’ (PRO-GRACE -https://www.grace-ri.eu/pro-grace). The results indicated that 76% of genebanks (13 of 17 respondents) were interested in adopting complementary in situ/on-farm roles alongside traditional ex situ activities. Genebanks have historically succeeded in supporting breeders and farmers while maintaining the PGR foundation for diverse crop varieties, but human population increase and climate change’s impact on crop production and food security are forcing a change of practice. Although some GRC may face limitations in skills and resources, with appropriate support, their roles could evolve to become even more critical for humanity’s future. This discussion focuses on how to better integrate in situ,ex situ, and user access in PGR conservation to provide greater diversity. We highlight current opportunities to: (1) clarify PGR conservation aims through proposed Principles of PGR Conservation and Use Congruence; (2) summarize practical processes for in situ and on-farm conservation; (3) promote resource management collaboration; (4) enhance user access to in situ and on-farm conserved PGR populations; (5) facilitate access to in situ conserved populations via the European Search Catalogue for Plant Genetic Resources (EURISCO, http://eurisco.ecpgr.org); and (6) identify future ways to better integrate in situ and ex situ conservation. For PGR actors and germplasm users, the clear advantage lies in addressing current challenges and ensuring greater diversity availability, with an enhanced role for genebanks or GRC at the core, ultimately leading to increased sustainable food production and long-term food security. The Principles of PGR Conservation and Use Congruence The aim of PGR conservation may be summarized in three fundamental principles, to ensure: (1) longterm, sustainable maintenance of PGR1diversity, (2) 1The scope of PGR found outside of GRC, or breeding collections is commonly focused on crop wild relative (CWR) and landrace (LR) diversity both of which are highly threatened. 206 Maxted et al active2conservation and characterization of crop, varietal and related wild taxon diversity using complementary3techniques and (3) conserved resource documentation and availability for utilization within the applicable legislative context. The use of complementary techniques provides additional security by employing multiple, diverse approaches to conserve these resources, ensuring greater security as each technique backs up and supplements the others. There could as well be other subordinate objectives, such as maintaining seed viability, phenotypic and genotypic characterization and evaluation of conserved resources, and ensuring standard material transfer agreement (SMTA) enforcement, but the three fundamental objectives should hold true for whatever form of conservation strategy is applied. Together, these objectives may be referred to as the Principles of PGR Conservation and Use Congruence; overall, conservation should, in the long-term, maintain the full breadth of genetic diversity, employ multiple conservation techniques, and make the conserved resources available to actual or potential users. These three objectives are met for most ex situ holdings (except for the requirement to link to complementary in situ conservation). Ex situ PGR conservation and use is well tested, and we know it already ‘works’, but there is now an urgent need to further develop in situ conservation approaches. Hawkes (1991) commented in the early 1990s that in situ techniques were in their “infancy”, and although advances in this area have been made (Maxted et al, 2020), in situ and on-farm conservation is still largely experimental and not based on more than 60 years of practice and the associated extensive evidence-base available for ex situ conservation. Additionally, effective standardization of in situ conservation techniques is itself challenging, as their application occurs in natural or semi-natural environments, or in on-farm locations, where diverse environmental, socioeconomic and cultural factors impact the target taxa, and effective PGR population managers (e.g. farmers, foresters, estate managers, etc.), may not be professional conservationists or have the necessary skills to maintain intrinsic genetic diversity. This is not to devalue the efforts of farmers or other landrace (LR) maintainers, or landscape managers, who have retained crop wild relative (CWR) populations on the estates they manage for extended periods of time. However, if in situ PGR conservation is to function as intended and be appropriately resourced, it must meet all three principles and objectives, as do ex situ approaches. Populations and diversity of in situ resources must be maintained in the long term via the application of complementary techniques, and the conserved resources must be 2Active conservation implies targeted management and monitoring of conserved CWR or LR populations, as opposed to passive maintenance of CWR or LR populations, where there may be a conservation ethos but no targeted management and monitoring. 3Complementary conservation implies the use of both ex situ and in situ techniques to conserve CWR or LR populations. available to users. If in situ PGR conservation does not ensure availability of the conserved resource, it will not meet the Principles of PGR Conservation and Use Congruence and it is unlikely ever to be seen as truly complementary to ex situ conservation. It should also be noted that the third principle, which conserved resources are available for use, may not always be achievable, for example, when the in situ conserved populations are rare or threatened, and few, or an ex situ conserved accession has limited seed numbers and low viability. In both cases, the sample may need to be multiplied or regenerated before it can be made available to users. The principle remains that resource availability is paramount, and any periods of unavailability should be temporary until germplasm can be offered. Practical processes of in situ and on-farm conservation To identify potential opportunities for integration, we need first to summarize and understand how in situ and on-farm conservation operate. The conservation–utilization continuum for in situ conservation is divided into four component steps and summarized in Figure 1 (adapted from Maxted et al (2020)): 1. Conservation planning. This involves: (i) selection of target conservation units, either CWR or wild food plant (WFP) taxa or crop LR (Maxted et al,1997c;Brehm et al,2017); (ii) prioritization, usually based on potential use value, relative crop value and threat, identifying an easily implementable inventory of highest priority CWR, WFP or LR (Brehm et al,2017;Nilsen et al,2017; FAO,2019b); (iii) ecogeographic and gap analyses to identify concentrations of the conservation units and predict which sites with target populations (Maxted and Kell,2008;Maxted et al,2012b; FAO,2019b); and (iv) field exploration to check the validity of the previous prediction and establish where the target diversity will be conserved in genetic reserves, other effective area-based conservation measures (OECM), on-farm, or in home garden. 2. Conservation technique implementation. Conservation targets are actively managed either in nature for CWR or WFP or cultivated on-farm or in-garden for LR diversity. This involves: (i) selection of sites with targeted resource diversity (Hawkes et al,2000;Maxted et al,2002;Dulloo et al,2008;Vetel¨ ainen et al,2009;Iriondo et al,2021); (ii) formulation of the management plan, a detailed plan for how the population(s) of the target taxa/crop are to be maintained and enhanced (Maxted et al,2002,2008;Dulloo et al, 2008;Vetel¨ ainen et al,2009;Iriondo et al,2021); (iii) implementation of the management plan, including the site interventions, implementation of which is likely to be experimental initially until tarGenetic Resources (2025), (S2), 203–223 207 get population retention is sustainable (Vetel¨ ainen et al,2009;Iriondo et al,2021); (iv) resource monitoring at set time intervals to check the success the management regime (Vetel¨ ainen et al,2009; Iriondo et al,2021); and (v) formation and upkeep of partnerships essential for in situ and on-farm conservation of the genetic resources to occur. 3. Conserved resource description. The preutilization stage will involve characterization and evaluation (Maxted et al,2020). These data may be uploaded alongside passport data in EURISCO to facilitate germplasm selection. 4. Conserved resource utilization. The in situ conserved resource should be available for use by breeders, farmers, researchers and other potential bona fide users. Forms of traditional utilization should be encouraged, provided it is not detrimental to the target taxon or taxa, thus fostering local support for conservation actions. Proposed resource management collaboration It is important to clarify not only how the target populations are managed, but also (1) who should provide oversight of the networks of in situ or on-farm sites and populations, and (2) who should practically implement the management interventions of individual in situ or on-farm sites and populations. There are several potential communities that might fulfil these roles: existing population managers, national GRC staff and other diverse PGR stakeholders (including allied nongovernmental organizations (NGOs), research centres and universities). As noted above, given that often the conserved in situ or on-farm genetic resources have been managed by the reserve/protected area (PA) manager, landowner, farmer or gardener for extended periods, one might assume they are the most appropriate to play both roles. While existing in situ and on-farm site managers should continue their successful management of individual in situ or on-farm sites and populations, the question is: do they have the necessary skills, tools and resources to provide oversight of the network(s) of in situ or onfarm sites and populations established? It can be argued that it would be impractical for individual in situ and on-farm site managers to provide oversight of the network(s) of in situ or on-farm sites and populations given they: (1) are unlikely themselves to use trait diversity from the conserved CWR or WFP populations; (2) lack skills and expertise in international and national policy and legislation; (3) lack skills and expertise in field trials or genomic analysis; (4) lack access to a PGR information system to aid in situ population management and transfer of germplasm to the end user; and (5) already have an existing heavy core load of activities in managing biodiversity populations or producing food and their scope to adding a significant additional activity is limited. Therefore, it would seem appropriate that national GRC staff (or other appropriate national PGR agency or PGR-focused NGOs) would be better placed with the necessary skills, tools, resources and long-term experience from ex situ PGR applications, to provide multi-site PGR governance and overall oversight of the networks of in situ or on-farm sites and populations, including overall monitoring of natural reserves, other effective areabased conservation measures (OECM) sites or on-farm systems to prevent population losses (Maxted et al, 2016). However, it is important to stress that national GRC staff cannot work in isolation. The PGR conservation goal of maximum PGR diversity conservation and availability can only be achieved by the three communities working in integrated collaboration, with national GRC staff providing national PGR leadership and oversight, individual PGR field population maintainers (i.e. reserve/PA manager, landowner, farmer or gardener) managing the genetic resources under their responsibility, and other PGR stakeholders (allied NGOs, research centres and universities) providing the necessary additional support. Furthermore, as the in situ or on-farm resource is maintained outside of a controlled unit, like a GRC, the local community within the vicinity of the in situ/on-farm resource site should also be involved in the conservation project management and associated decisions. Individual roles will vary depending on multiple factors (e.g. taxa included, whether wild or cultivated, resources available, value of resource conserved, etc.), therefore stakeholder discussions and negotiations will form part and parcel of the process of defining the roles of each actor, however it can be safely stated that the key expertise and areas of responsibility are likely to include those presented in Table 1 . To aid clarity, Figure 2 highlights those components managed by GRC staff, and in situ site maintainers alone, and which may be managed jointly. Collaboration between the three communities would be critical and involve periodic meetings of a PGR In Situ Population Management Committee. Such an integrated approach to in situ and ex situ collaboration would extend each communities roles and responsibilities. However, for those maintaining PGR populations (PA, OECM or on-farm field maintainers) and given the target populations were selected because of their ‘health’, the additional workload is not foreseen as being significant, at least initially, as it would primarily involve monitoring target populations, while the provision of additional ecosystem and food services from the site would underpin the public good value of maintaining PGR populations. Furthermore, in some countries, additional targeted PGR conservation could generate additional subsidies or added income for the site maintainers/owners through government funding (such as payments for ecosystem services, subsidies for farmers who cultivate and conserve landraces that suffer from genetic erosion), so the benefit to PGR field population maintainer could be substantial. The proposed changes outlined for the national GRC would also be significant, possibly requiring additional staff with in situ expertise and Genetic Resources (2025), (S2), 203–223 Plant GRC roles in linking in situ and ex situ conservation 208 Maxted et al Figure 1. Schematic description of key elements of in situ conservation to utilization pathway. Green, in situ; brown, ex situ; red, threatened populations; gold, utilized PGR; blue, conservation steps; CWR, crop wild relatives; LR, landraces; OECM, other effective area-based conservation measures; WFP, wild food plants. additional resources, but the additional role would fall within the existing genebank’s remit – Genebank Managers Network (https://www.ecpgr.org/about/gen ebank-managers-network) and AEGIS initiative (https: //www.ecpgr.org/aegis) of the European Cooperative Programme for Plant Genetic Resources (ECPGR) – and would substantially boost the genebank’s role in national biodiversity conservation. For all three collaborating communities, increased collaboration will involve additional time and resource commitments, incurring additional costs. Therefore, it is crucial to identify sustainable funding mechanisms to cover these costs, even if they are anticipated to be minor. However, any additional costs incurred due to collaboration and changes in roles would be far outweighed by the potential benefits of increased diversity available for breeders and other stakeholder’s use (Maxted and Brehm,2023). Access to and conservation of additional germplasm significantly enhances the diversity of collections, a core GRC and genebank objective, thereby better fulfilling their professional mandate. As a final point, the collaboration as outlined in this document, involves the transfer of in situ or onfarm samples from their original locality to a nominated ex situ GRC for backup and to facilitate access for germplasm users. This means that the provisions emanating from the International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA) (FAO, 2001) and the Convention of Biological Diversity (CBD) Nagoya Protocol (CBD,1992,2011) are triggered and there is the need for an SMTA or Internationally Recognized Certificate of Compliance (IRCC) respectively, between the in situ maintainer and the recipient nominated GRC. This would need enacting even if the GRC had no intention to utilize the germplasm itself, but simply to conserve the in situ or on-farm sample and make it in turn available to more active users. By virtue of the relationship between the in situ/on-farm source, the GRC and the end user, the involved actors would be required to address the requirements to ensure fair and equitable sharing of benefits arising from the sample‘s potential final utilization, depending on the terms established under national regulations. The actual scope of the three-way (source, GRC and end user) relationship would require expert deconstruction and is therefore beyond the scope of this document but must be resolved before any germplasm transfer occurs. Genetic Resources (2025), (S2), 203–223 209 Table 1. Collaborative activities of national Genetic Resource Centre (GRC) staff, plant genetic resource (PGR) population maintainers and other stakeholders. CWR, crop wild relatives; LR, landraces; WFP, wild food plants CWR, WFP or LR in situ population conservation National GRC staff’s role PGR population maintainer’s role Other stakeholder’s role Lead preparation and periodic revision of National PGR Strategy and Action Plan, including in situ site selection and management plan production. Contribute to preparation and periodic revision of National PGR Strategy and Action Plan, lead in situ site selection and management plan production. Contribute to preparation and periodic revision of National PGR Strategy and Action Plan, and in situ site selection and management plan production. Lead national PGR in situ conservation site network management. Contribute to PGR in situ conservation site network management. Contribute to PGR in situ conservation site network management. Assist with implementing the site’s individual management plan. Lead implementation of individual site management plan. Assist with implementing of individual site management plan. Assist with periodic monitoring of target populations and analysis of demographic and genetic trends. Lead periodic monitoring of target populations and analysis of demographic and genetic trends. Assist with periodic monitoring of target populations and analysis of demographic and genetic trends. Assist with periodic revision of individual site management plan and building evidence base. Lead periodic revision of individual site management plan and building evidence base. Assist with periodic revision of individual site management plan and building evidence base. Target population characterization and evaluation. Ensuring user access to in situ conserved resources (via ex situ backup samples). Periodic collection of target populations for ex situ representative backup samples. Diverse research projects focused on aiding effective PGR diversity conservation and use. Ensure integration of in situ and ex situ conservation activities and support tools and applications to aid in situ conservation site network management, e.g. national inventories, management and monitoring, germplasm access, characterization and evaluation databases and information management. Collation of site and PGR population data, and integration with national PGR databases, activities assisted by network tools and applications. Diverse research projects supporting national PGR population management and use, and associated tool and application development. Promotion of national integration into international PGR community. Promotion of PGR integration into the broader biodiversity community. Participation in national and international research actions. Lead and participate in the National PGR In Situ Population Management Committee. Participate in the National PGR In Situ Population Management Committee. Participate in the National PGR In Situ Population Management Committee. Genetic Resources (2025), (S2), 203–223 Plant GRC roles in linking in situ and ex situ conservation 210 Maxted et al Figure 2. Schematic description of key elements of in situ conservation, highlighting Genetic Resource Centre (GRC) staff (dark red), in situ populations manager (green) and joint (orange) responsibilities. CWR, crop wild relatives; LR, landraces; OECM, other effective area-based conservation measures; WFP, wild food plants. User access to in situ and on-farm conserved PGR populations The endpoint of PGR conservation is not conservation itself but ensuring that conserved germplasm is available for present or potential future utilization (Maxted et al, 1997a). The pathway of use for ex situ conserved PGR is tried and tested, but, apart from the positive activities of farmers and farming NGOs focusing on PGR diversity and farming systems, the in situ pathway to utilization has yet to be established. Without effective in situ conservation-to-use linkage, it is doubtful whether in situ conservation sites and site networks will ever be established (Maxted,2019). Therefore, establishing links between in situ resources and use is fundamental to ensure additional germplasm access and the promotion of in situ conservation itself (Maxted and Brehm,2023). Maxted and Kell (2008); Maxted and Palm´ e(2016) and Maxted (2019) each reviewed potential models for how in situ conserved resources might be linked to user access, either accessed for use directly from the in situ population or indirectly via an ex situ conservation facility (Figure 3). Five potential options have thus far been proposed for promoting user access to in situ and on-farm conserved PGR and are elaborated in Table 2. Except for Option 3, users request an in situ PGR population sample and ≈(20-) 40–50 viable seeds are dispatched to the end user, fulfilling the in situ to-use prerequisite outlined in the Principles of PGR Conservation and Use Congruence. The chosen option may vary based on GRC facilities, available resources, conservation practices, and constraints from PGR maintainers or national authorities. However, assuming resources are adequate and constraints do not limit distribution, Option 5 achieves the Principles of PGR Conservation and Use Congruence, making the in situ resource-to-user link via the GRC, while placing the minimum additional burden on the GRC staff and their resources. However, such an approach has not been practically implemented in any country. The reason is not thought to be that Option 5 or the other options are not conceptually sound, but due to funding limitations, risk aversion, lack of formal incentives, or the necessary skills and tools to promote in situ utilization. It could also simply be that active CWR, WFP in situ or LR on-farm conservation itself is only now being tentatively initiated, in situ conserved resources are uncharacterized and evaluated, the potential of in situ or on-farm germplasm access is unflagged so potential users are unaware such resources are accessible or how to access them. Genetic Resources (2025), (S2), 203–223 211 Figure 3. Five options to linkplant genetic resource (PGR) in situ conserved resources to user access: 1. Direct in situ supply, 2. Standard ex situ conservation, 3. Blackbox in situ safety backup, 4. In situ demand and supply, and 5. In situ backup & supply. OECM, other effective area-based conservation measures. Genetic Resources (2025), (S2), 203–223 Plant GRC roles in linking in situ and ex situ conservation 218 Maxted et al In situ conservation should be complemented by ex situ strategies to: (1) provide long-term backup for security and potential population reinforcement or reinstatement; (2) assist in characterization and evaluation; and (3) ensure ease of access for end users. Likewise, ex situ conservation should be complemented by in situ approaches to: (1) maximize the preservation of taxonomic and genetic diversity; (2) allow for the evolution of adaptive traits in changing environments; and (3) address the evolving demands of end users. Thus, both ex situ and in situ conservation methods are interdependent and should function in a mutually supportive manner. However, unlike ex situ conservation, which can be largely managed within controlled environments, in situ conservation necessitates the active participation of diverse actors with various skill sets (ecology, wild plant biology, field genoand phenotyping, remote monitoring, climate change management, invasive species and pest management) to enact conservation actions, adding layers of complexity and associated challenges. Integration of these diverse actors in a distributed Research Infrastructure on Plant Genetic Resources is likely to unite these additional actors. Moreover, a critical question arises: who will take primary responsibility for coordinating in situ conservation efforts? The experiences of the ECPGR CWR and On-farm Conservation and Management Working Groups have demonstrated that neither protected areas nor farming communities can effectively coordinate in situ PGR diversity conservation activities, and many are reluctant to engage in formal in situ PGR conservation. Protected area managers focus on biodiversity rather than crop diversity conservation, while farming communities are primarily engaged in commercial agricultural production rather than systematic diversity conservation. Therefore, there is a pressing need for additional training for GRC staff and/or extending collaboration with actors possessing the necessary skills and experience in ecology, pest management and field conservation to complement the existing GRC staff’s expertise in genotypic, phenotypic and agronomic evaluation, sampling, viability and phytosanitary testing, documentation, data upload to EURISCO, and distribution to users, including knowledge of national and international legislative implementation. This collaboration is fundamental to enhancing the conservation of in situ diversity, its description and its availability to end users. Additionally, the existential problem of user supply is often underestimated by the in situ PGR community. For PGR conservation to be effective, meaningful, and serve a utilitarian purpose beyond its intrinsic value in nature preservation, a link must exist between conservation and utilization. However, neither protected areas nor farming communities possess experience in germplasm supply within the context of access and benefit-sharing legislation. Consequently, it can be argued that without the involvement of the ex situ community in these roles, in situ implementation risks becoming limited to ’academia,’ ’hobbyists’ or shortterm project support without long-term sustainability. Therefore, it is evident that the application of ex situ and in situ strategies is mutually dependent, and their complementary integration should be led by GRC. Leadership from GRC would entail adjusting their perspective to encompass both ex situ and in situ aspects, along with appropriately increased resources to fund the necessary structural and skill provisions for practical implementation. Conversely, if the in situ or on-farm community was able to take such a leadership role, would the genebanks welcome the competition? There is also an economic argument for GRC to adopt a more proactive role in in situ conservation. As outlined, one justification for PGR conservation is to enhance user access and benefits, which may encompass various industries, with the most prominent being those related to economic and food security, medicinal products and material uses. The most recent estimate of the use value for CWR closely related to 29 globally important crops is US$42 billion, with a potential future value of $120 billion. The annual gross added value was $581 billion in 2010, indicating that CWR are already valued at about 7% of the annual production value of these 29 crops (PWC,2013). This valuation is conservative, as it does not account for the potential expansion of CWR use in breeding these or other crops, nor the value of utilizing LR diversity. Therefore, the overall annual gross added value of using PGR diversity in crop improvement could approach a trillion US dollars. This significant valuation raises the question: does not the potential revenue stream justify the modest investment required now in PGR conservation to secure future substantial benefits? The rationale for integrating ex situ and in situ conservation lies in the fact that ex situ collections typically capture only a snapshot of the genetic diversity present in natural populations at the time of collection. It also should be acknowledged that over time, genetic drift or selection during storage and regeneration can lead to the loss of some of this genetic variation. In contrast, in situ conservation allows the remaining spectrum of genetic diversity to persist and evolve naturally in response to environmental changes. Without leveraging both approaches, a significant portion of the genetic diversity available in natural populations remains untapped, limiting its potential contribution to crop improvement and other industries. While the practical establishment of CWR genetic reserves or LR on-farm diversity maintenance sites has progressed more slowly than anticipated, this may be partly attributed to the PGR community’s longstanding focus on the established ’in-nature and on-farm sampling to genebank to user’ paradigm (Guarino et al, 1995,2012;Hawkes et al,2000;Smith et al,2003; FAO,2014). This paradigm has proven resilient and successful over the last century, consistently meeting the needs of breeders and consumers. However, the very success of this established paradigm poses a significant challenge to the adoption of in situ conservation approaches. To gain wider acceptance, these approaches Genetic Resources (2025), (S2), 203–223 219 must articulate an equally robust and straightforward model that demonstrates long-term effectiveness – the PGR germplasm user is indifferent to the conservation source if it meets their trait needs. Promoting in situ application includes clearly communicating the value of the proposed in situ to ex situ to use paradigm and its mutual advantage in diversity breadth. Although the clarification of the Principles of PGR Conservation and Use Congruence and the derived proposals presented provide an initial foundation for a proposed in situ to ex situ to use paradigm led by the national GRC, further development will be necessary based on a growing evidence base. Another related topic that has progressed more slowly than anticipated is the systematic ex situ and in situ conservation of WFP. These include fruits, leafy vegetables, woody foliage, bulbs and tubers, cereals and grains, nuts and kernels, saps and gums, mushroomsand seaweeds (Wunder,2014). WFP have historically served as a coping strategy for many rural households, particularly during the ’hungry season’ before the next season’s crops ripen and as part of subsistence farming systems (Hunter et al,2015;Kennedy et al, 2017). FAO (2019a) estimates that around one billion people globally incorporate wild foods into their diets regularly, and forests alone provide livelihoods and food for approximately 300 million people through non-timber forest products. However, WFP are rarely included in PGR conservation initiatives and are unlikely to be targeted for biodiversity conservation only if they are threatened or rare. FAO (2019a) calls for (1) active ex situ and in situ conservation and sustainable use, (2) breeding of improved varieties, and (3) raising awareness of the importance of WFP, particularly local and traditional foods that are vital for nutritionally balanced, healthy diets and food security. WFP, like CWR, are simply wild species with specific food value, although the former is associated with direct consumption rather than trait provision. Therefore, WFP planning and conservation implementation are unlikely to differ significantly from CWR-based actions, making it timely to test this assumption. Implementing WFP conservation falls within the remit of national GRC activities and should be integrated with other PGR activities. Most importantly, WFP can provide material for future domestication efforts, thereby expanding the foundation of our food production systems. Here much has been made of expanding in situ/ex situ integration, but there is also significant leverage in in situ/on-farm working more closely with biodiversity communities. CWR and LR could be used as ‘cultural ambassadors’ to help promote PA-based conservation or traditional cultivation practices. The collaboration offers opportunities to marry biodiversity conservation management with food security or traditional foods associated with healthier lifestyles. While such collaboration would also help conserve the critical PGR resource more extensively and effectively – demonstrating the mutual relevance of each community contribution – PA don’t only maintain birds, mammals and reptiles, they conserve the founding resource for our food. Traditional farming is not just picturesque, it sustains cultural benefits such as recreation, education, spiritual and creative enrichment, and improved mental health and wellbeing. Whilst PA management may recognize the importance of these ecosystem services, their consideration and usefulness in site management decision-making is worth closer understanding. There exists an opportunity and a central role for the proposed GRACE research infrastructure (see https:// www.grace-ri.eu/pro-grace), which builds on 55 years of ECPGR collaborative networking aimed at ensuring long-term conservation and facilitating utilization of PGR to implement the necessary transition from genebanks to GRC and enact more effective in situ PGR conservation. This role may prove existentially important for humanity in the future. Without appropriate financing, skills and capacity provision, and cooperation with the broader biodiversity community, establishing and maintaining in situ and on-farm networks would be unsustainable in the medium to long term, even under GRC direction. The core mission of the PGR community remains unchanged, as summarized in the Principles of PGR Conservation and Use Congruence, and it is essential to reassess and reconfigure this mission to ensure it is fit for purpose today and in the future. Conclusions The dual challenges of human population growth and climate change’s negative impact on crop production have resulted in increased demand from germplasm users and consumers for greater breadth of diversity. Ex situ genebanking alone is unable to secure such breadth of diversity, as are in situ or on-farm conservation activities; the urgency of the situation is such that the muchdiscussed but rarely applied implementation of complementary PGR conservation offers the only practical and expedient solution. The Principles of PGR Conservation and Use Congruence describe the fundamental principles of PGR conservation (long-term, sustainable conservation, application of complementary conservation techniques, and documentation and availability of the conserved resource for utilization) and provide a framework for indicating success. Evidence and experience have shown that neither ex situ,in situ nor on-farm conservation functions adequately in isolation, but further that systematic in situ and on-farm genetic conservation is not a priority for practitioners of either biodiversityfocused conservationists or production-based farmers. The comprehensive integration of ex situ,in situ and on-farm conservation communities and their activities, with the local communities where the bulk of the genetic resources exist, led by national GRC and CGIAR institutes, is now critical for global, regional, national and local food security; failure to address this issue could have devastating consequences for humankind in the 21st century. Specific recommendations are outlined for collaborative resource management, user access to in Genetic Resources (2025), (S2), 203–223 Plant GRC roles in linking in situ and ex situ conservation 220 Maxted et al situ and on-farm conserved PGR, improving user selection of in situ conserved populations and what the future challenges and opportunities there might be for future in situ–ex situ integration. Other recommendations will undoubtedly come from further steps toward PGR community integration. Although realistically this initiative is doomed to failure unless national GRC step up to take the lead, skill gaps are filled, and they are adequately resourced. Supplemental data Supplemental Table 1. Genetic conservation strategies and techniques (Maxted et al,2020). Supplemental Table 2. Horizon scanning issues associated with CWR in situ–ex situ conservation in 2025 that require resolution by 2035. Supplemental Table 3. Horizon scanning issues associated with LR in situ–ex situ conservation in 2025 that require resolution by 2035. Acronyms used • ABS − Access and Benefit Sharing • AEGIS − A European Genebank Integrated System • C & E − Characterization and evaluation • CBD − Convention on Biological Diversity • CWR − Crop wild relatives • CWR-NI − CWR-National Inventory • ECPGR − European Cooperative Programme for Plant Genetic Resources • EURISCO − European Search Catalogue for Plant Genetic Resources • FAO − Food and Agriculture Organization of the United Nations • GPA − Global Plan of Action • GR – Genetic reserve • GRC − Genetic resource centre • IRCC − Internationally Recognized Certificate of Compliance • ITPGRFA − International Treaty on Plant Genetic Resources for Food and Agriculture • LR − Crop landrace • NGO − Non-governmental organization • OECM − Other effective area-based conservation measures • PA − Protected area • PGR − Plant genetic resources • SMTA − Standard material transfer agreement • TAGS − Tools to Aid Germplasm Selection • WFP − Wild food plant Acknowledgements We gratefully acknowledge the support of the Horizon Europe programme, project ‘Promoting a Plant Genetic Resource Community for Europe (PRO-GRACE)’, N. 101094738 in preparation for this review. 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