The history of ex situ conservation is relatively short, not more than a century old. During the middle of last century, triggered by the realization that genetic erosion was threatening the existing landraces and wild relatives of the major food crops, global efforts to collect and conserve the genetic diversity of these threatened resources were initiated, predominantly orchestrated by FAO. National and international genebanks were established to store and maintain germplasm materials, conservation methodologies were created, standards developed, and coordinating efforts were put in place to ensure effective and efficient approaches and collaboration. In the spontaneously developing global conservation system, plant breeders played an important role, aiming at the availability of genetic diversity in their breeding work. Furthermore, long-term conservation and the safety of the collected materials were the other two overriding criteria that led to the emerging international network of ex situ base collections.
Plant genetic resources are the foundation of our food production system, thanks to the genetic diversity they contain. It is this genetic diversity, both between and within crop species and their wild relatives, that allow crops to evolve and adapt to changing conditions, either natural or human-created conditions. Since the first steps of early farmers to start the process of domesticating species from wild plants in the Near East more than 10,000 years ago, plant genetic resources and their diversity allowed humankind to develop crops according to its needs and to spread them around the world; thus, securing our plant food basis.
Since these ancient times, the number of domesticated crops has steadily increased, and the cultivated forms or varieties of most of these crops have also increased and collectively become more diverse when moving from one region to another. Human and natural selection have been the driving force behind this diversification, but this process was only possible because of the genetic diversity available within and between these crop varieties and the related wild species that collectively form the diversity gene pool 
. Genetic mutations in the crop genome are a permanent source of genetic diversity that allowed and continue to allow human and natural selection to be successful. Human exploitation of genetic diversity drastically increased when plant breeding became established, some 150 years ago 
. This process of purposely generating new diversity through crossing different individuals followed by subsequent selection, resulted in high(er) yielding elite varieties. The success of this human managed evolution meant a steady replacement of older and usually well-adapted cultivars and even of entire crops. The loss of genetic diversity is called genetic erosion and was the trigger for targeted conservation efforts worldwide 
With the steady and increasing loss of genetic diversity since the middle of the last century for many of the crops cultivated worldwide and particularly for the main food crops, the need for systematic collecting and conservation of this diversity was recognized, and global conservation activities were initiated. Gradually, the Food and Agricultural Organization of the United Nations (FAO) in Rome assumed a coordinating role, supported by the International Board for Plant Genetic Resources (IBPGR), founded in 1974, one of the CGIAR centres, whose secretariat was initially based at FAO, thus serving as a technical and advisory institute for FAO and its political bodies such as the Commission on Plant Genetic Resources and later the International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA). Gradually, through IBPGR’s research, coordination and scientific advice and training were provided to countries worldwide, and a global network of plant genetic resources conservation centres, called genebanks, was established 
. Political debates at FAO, and IBPGR’s research efforts aimed at collecting and improving the conservation of plant genetic resources for food and agriculture (PGRFA), somehow led to a more or less spontaneous creation of a global long-term conservation system of PGRFA 
. This system underwent an evolutionary process itself, taking advantage of new scientific and technical developments and adjusting to evolving political conditions. It is this system that we will critically assess and review, with its strengths and weaknesses, with the aim to provide a perspective on how the system can be strengthened and can be made more rational in order to enable effective and efficient long-term conservation.
Whereas natural and human-made ecosystems harbour the biodiversity of plants, animals, and microbes embedded in a physical environment, the focus in this paper will be on the plant genetic resources that are used for food and agriculture, i.e., PGRFA, or plant agrobiodiversity. These PGRFA comprise landraces and primitive and obsolete cultivars, crop wild relatives and modern varieties. Sometimes, plant breeding and other research materials are also regarded as genetic resources that should be included in genebanks.
Regarding the conservation activities, the main focus of this paper will be on long-term ex situ conservation, i.e., genebanks that manage seed, field, in vitro, and cryopreserved collections as well as DNA samples. Thus, not only are seeds important organs for conservation, but entire plants, pollen, tissues, cell suspensions, and more recently, DNA are also used. As not all plant agrobiodiversity can be collected and stored in genebanks, e.g., many wild food plants, many crop wild relatives, etc., we also look at in situ or nature conservation as well as at the on-farm maintenance of landraces and other genetic resources that require keeping the population structures of the material to be protected intact and/or to ensure a continuous evolution or the maintenance through steady cultivation or management. This dynamic evolutionary conservation stands in contrast to the frozen and static conditions that genebanks practice. Whereas ex situ conservation tends to focus on genotypes, in situ and on-farm conservation aim at natural and/or human-made populations and mixtures. It might be obvious that a balanced integration of these different conservation approaches will be needed to optimize the conservation system, as these approaches are complementary.
As conservation is frequently undertaken with the aim of keeping genetic diversity available and easily accessible for use, i.e., by farmers, breeders, or researchers, availability aspects are also important to be considered when deciding on the conservation ‘approach’. Therefore, due attention will be given to how to increase the use of materials conserved under long-term conservation conditions.
Detailed knowledge of the conserved genetic resources is a key requirement for rational, effective, and efficient conservation as well as to facilitate the use of the resources. Thus, research on plant genetic resources in situ or in genebanks is an essential activity to support these requirements. This aspect will be given due attention.
Besides the importance of creating new knowledge of the materials under conservation and facilitating their use, the application of new technologies in conservation and use is critically important to achieve rational, efficient, and effective long-term conservation and to facilitate the use of plant agrobiodiversity.
Plant agrobiodiversity is distributed across the world; therefore, the sovereignty of national states is an important legal aspect that was recognized in the Convention on Biological Diversity (CBD), and the accessibility to these resources is thus determined by individual states. Moreover, genetic resources might be protected by intellectual property rights, hence the legal and policy framework for the conservation and the use of PGRFA is an important element to ensure rational, efficient, and effective conservation and use.
Other aspects that might directly or indirectly impact conservation decisions include training and capacity building, awareness creation, participatory approaches, economic considerations, and possibly others. These aspects are not the focus of this paper or of this Special Issue but can be of critical importance to achieving a rational and sustainable long-term conservation system.
2. History of the Development of the Long-Term Conservation Practices and the Evolving Global Conservation System
Crop and related genetic diversity underpin the productivity, sustainability, resilience, and adaptive capacity of agricultural systems and, thus, their evolutionary potential 
. This diversity, contained in the so-called plant genetic resources has played a key role in the developments of agriculture since the first steps towards the domestication of our crop plants, the subsequent diffusion of the domesticates as well as the associated weeds and wild relatives from the centres of domestication into the world and the ongoing improvement and adaptation of the crops to ever-changing environments, cultural practices, and human-made and natural threats. The first farmers started to migrate out of the Fertile Crescent to new geographic areas about 10,000 years ago, carrying genetic resources with them 
. Whereas this process will have caused bottlenecks and thus might have impacted the evolution of these crops, the introduction of new and possibly more genetic diversity, natural mutations as well as natural and human selection have resulted in an enormous diversity of crops and varieties. This traditional crop development process underwent significant changes through rediscovery, around the turn of the 20th century, of the laws of inheritance proposed by Gregor Mendel in 1865 and 1866, which formed the basis for the science of genetics and thus, the birth of scientific plant breeding 
One of the first persons to realize the importance and use the power of genetic diversity in crop improvement was Nicolai Vavilov, a Russian geneticist and a director of the Lenin All-Union Academy of Agricultural Sciences at Leningrad (now the Vavilov Federal Research Centre of Plant Genetic Resources—VIR) who was requested by Lenin, the head of the government of Soviet Russia and later the Soviet Union, to breed plants that could be cultivated in Siberia and thus would contribute to increased food production after the First World War 
. Collecting about 50,000 samples of crop plants systematically and throughout the world and evaluating them to assess their traits, he realized that the collected genetic diversity was largely confined to restricted areas, the so-called centres of diversity/origin of our crops 
Plant introduction centres that later grew out into genebanks were established in several countries to meet the increasing demand by plant breeders for more diversity. These included the All-Union Institute for Plant Industry in St Petersburg (in 1920), the Commonwealth Potato Collection in Cambridge, UK (before the Second World War), collections for the research programmes of the Rockefeller Foundation in the USA (1943), and The National Seed Storage Laboratory (NSSL) in Fort Collins, CO, USA (1958) 
. The latter became the long-term storage facility for valuable germplasm propagated by seeds from the four regional plant introduction stations and an inter-regional station for potatoes 
. During the 1950s and 1960s, several national plant introduction centres/genebanks were established on all continents, plant quarantine regulations were initiated (such as those in West Africa), and plant exploration and collecting started (such as the initiatives in Latin American countries). During the 1940s and 1950s, advanced and well-organized global germplasm collecting missions were coordinated by the Rockefeller Foundation in the USA 
With the increasing successes of plant breeding and the spread of modern and frequently high-yielding varieties, especially of the major food crops, a process of variety and later, even a process of crop replacement started and resulted in significant losses of genetic diversity, a development that was called ‘genetic erosion’ 
. As early as 1936, Harlan and Martini raised the issue of genetic erosion in a USDA report devoted to barley breeding 
, and Vavilov had noted the increased loss of landraces. Particularly, during the so-called ‘Green Revolution’, which started in the late 1950s until the early 1970s, the success of high-yielding (dwarf) varieties of wheat and rice, together with new agricultural technologies, led to drastic losses of the traditional landraces of these crops, and this triggered concern in organizations such as the European Society for Research and Plant Breeding (EUCARPIA) and FAO 
. In 1966, the EUCARPIA delegates advised European plant breeding institutes to foster continental collaboration through the establishment of four sub-regional genebanks in what was then West Germany (FAL in Braunschweig, for NW Europe); in East Germany (Gatersleben), Poland (Radzikow), Russia (St Petersburg) and/or others for Central and Eastern Europe; in Italy (Bari, for Southern Europe); and Sweden (Lund, for the Nordic countries) 
. Gradually, regional and global networking increased, and the contours of a global conservation system became visible.
2.1. The Role of FAO
During the 1950s and early 1960s, FAO became the major actor in the conservation of plant genetic resources. Besides the World Catalogues of Genetic Stocks for wheat, rice, maize, and barley, they started to publish the FAO Plant Introduction Newsletter and organized technical meetings/conferences (see below). Salient historical events with respect to the global conservation system are summarized in Table 1
and, where applicable, reference to the Table is made in the text. The first meeting was called the ‘Technical Meeting on Plant Exploration and Introduction’ and was held in 1961 (Table 1
. A Panel of Experts on Plant Exploration and Introduction was established in 1965. The panel included visionary scientists such as Sir Otto Frankel (CSIRO, Australia), Professor Jack Harlan (University of Illinois, Urbana, IL, USA), and Professor Jack Hawkes (University of Birmingham, Birmingham, UK); in addition, Ms. Erna Bennett, (FAO, Rome, Italy) served as one of the supporting secretarial staff members of the panel. Reports of the six-panel meetings were published between 1968 and 1974 
. This panel also played an important role in the planning and steering of the first two International Technical Conferences that the FAO organized in collaboration with their partners 
Table 1. Historical events of relevance to the establishment and evolution of the global PGRFA conservation, including the international network of base collections.
||Main Outputs and (References)
||Underpinning Principles (Reference)
||Establishment of first genebanks
||VIR, St. Petersburg (1920); Commonwealth Potato Collection, Cambridge (<2nd World War); research collections by Rockefeller Foundation, USA (1943); Fort Collins, CO, USA (1958) 
||Recognition of genetic erosion in landraces by 
||Publication Studies on the Origin of Cultivated Plants by N. Vavilov
||Monograph in Bulletin of Applied Botany and Plant-Breeding; 
||‘This monograph, dedicated to the memory of De Candolle, seems to be the most substantial contribution made since his day to the history of our main cultivated plants’ .
||Founding of IRRI
||Jointly established by Government of the Philippines’ and the Ford and Rockefeller Foundations 
||One of the first international genebanks; focus on rice genepool.
||Technical Meeting on Plant Exploration and Introduction, FAO Rome
||Report of the meeting 
||Mission-driven approach: conservation and use closely linked, tied to plant breeding, dominance of ex situ collections, mainly in developed countries.
||Establishment of the FAO Panel of Experts on Plant Exploration and Introduction.
||Six meetings and reports of same during period from 1967–1975 
||Formulation of criteria, standards, and procedures for the conservation and use of PGR.
||Formal establishment of CIMMYT
||Joint Mexican—Ford Foundation breeding project in progress since 1943 
||Norman Borlaug awarded Nobel Peace Prize (as wheat breeder) in 1970.
||Recommendation to foster continental collaboration through the establishment of four sub-regional genebanks in Europe 
||First indications of establishing a (global) conservation system or network.
||FAO/IBP (first) Technical Conference on Plant Exploration, Utilization and Conservation of Plant Genetic Resources, Rome
||Publication of Genetic Resources in Plants—Their Exploration and Conservation 
||Need for surveys; concern about genetic erosion of landraces and wild relatives; long-term ex situ collections; guidelines for establishment of global network for ex situ long-term conservation; international collaboration; in situ conservation as a complementary strategy.
||Third Session of the FAO Panel of Experts on Plant Exploration and Introduction, Rome
||Establishment of collecting priorities by crops (and later) by regions.
||Second FAO Technical Conference on crop genetic resources, Rome, Italy
||Book on Crop Genetic Resources for Today and Tomorrow 
||Plan of action agreed; panel of experts formulated basic criteria for conservation and use of genetic material (availability; maintaining genetic variability for the long-term; categorizing ex situ collections: base, active, and working collections.
||FAO/IBP Technical Conference on Genetic Resources, Rome, Italy
||Plan of Action 
||Recommendation to establish in situ collections.
||Establishment of IBPGR
||Established as secretariat for its board of trustees, administered by FAO and, technically, as one of the international centres of the CGIAR 
||Expected to coordinate global exploration and collecting efforts and to orchestrate a global network of genebanks.
||Third FAO, UNEP and IBPGR Technical Conference on PGR, Rome, Italy
||Clear focus on routine genebank operations; in vitro and in situ (CWRs) conservation; concerns about NUS.
||22nd Session of the FAO Conference, Rome, Italy
||Adoption of the International Undertaking on Plant Genetic Resources; establishment of the Commission on Plant Genetic Resources for Food and Agriculture (CGRFA) and of the Global System on Plant Genetic Resources 
||Shared principles; IU non-legally binding; PGRs are a common heritage of humankind; genetic stocks and breeding lines included; germplasm exchange through a network of genebanks; commission provides oversight to system.
||3rd Regular Session of Commission on GRFA, Rome, Italy
||Call for the development of the International Network of Ex Situ Collections under the Auspices of FAO 
||Lack of clarity regarding the legal situation of the ex situ collections.
||25th Session of the FAO Conference, Rome, Italy
||Resolution 4/89: Adoption of an agreed interpretation of the IU; Resolution 5/89: Farmers’ Rights 
||Plant breeders’ rights are not inconsistent with IU; recognition of Farmers’ Rights.
||26th Session of the FAO Conference, Rome, Italy
||Resolution 3/91 
||Recognition of the sovereign rights of nations over their PGRFA; agreement on development of 1st State of the World’s PGRFA and Global Plan of Action on PGR.
||UN Conference on Environment and Development (UNCED), Rio de Janeiro, Brazil
||Convention on Biological Diversity (CBD) (entered into force on 29 December 1993);
||Biodiversity vs. genetic resources; national sovereignty of states over their resources.
|Chapter 14 of Agenda 21
||Call for the strengthening of the FAO Global System on Plant Genetic Resources.
|Chapter 16 of Agenda 21
||Biotechnology can assist in the conservation of biological resources (e.g., ex situ techniques); risk assessment of LMOs, biosafety issues.
|Adoption of Resolution 3 of the Nairobi Final Act 
||Recognises matters not addressed by the convention: a. access to existing ex situ collections; b. questions on Farmers’ Rights; requests FAO forum to address these matters.
||1st Extraordinary Session of the CGRFA, Rome
||Start of negotiations for revision of IU; 12 centres of CGIAR sign agreement with FAO, placing their collections under the Auspices of FAO )
||CGIAR centres agree to hold the designated germplasm in trust for the benefit of the international community.
||4th International Technical Conference on PGR, Leipzig, Germany
||Global Plan of Action for the Conservation and Sustainable Use of PGRFA ; First Report on the State of the World’s PGRFA 
||Recognition of in situ and ex situ approaches; fair and equitable sharing of benefits arising from the use of PGRFA.
||31st Session of the FAO Conference, Rome, Italy
||Resolution 3/2001: adoption of the International Treaty (entered into force on 11 September 2004) 
||A legally binding agreement; recognition of Farmers’ Rights (a national responsibility); access and benefit-sharing
||Establishment of the Global Crop Diversity Trust
||Endowment fund, the income from which will be used to support the conservation of distinct and important crop diversity in perpetuity through existing institutions .
||Coordinates the Genebank Platform (of the CGIAR operated genebanks)
||First meeting of the Governing Body of the ITPGRFA, Madrid, Spain
||Standard Material Transfer Agreement (SMTA); relationship between the Treaty and the Crop Trust; agreement between GB and CGIAR centres (Art. 15) .
||SMTA is the legal instrument through which the MLS operates; recognition of the Crop Trust as an ‘essential element’ of the Treaty’s funding strategy; ex situ genebank collections of CGIAR are put under the Treaty (replacing agreement between CG centres and FAO).
||Establishment of the Svalbard Global Seed Vault
||Additional safety back-up for long-term ex situ collections.
||12th Regular Session of the CGRFA, Rome, Italy
||Second Report on the State of the World’s PGRFA 
||Report developed through a participatory approach with member countries
||143rd Session of the FAO Council, Rome, Italy
||Second Global Plan of Action for the Conservation and Sustainable Use of PGRFA 
||Need for a roadmap on climate change and genetic resources for food and agriculture
The first International Technical Conference was held in 1967 in Rome and was organized by FAO and the International Biological Programme (IBP) under the title ‘Technical Conference on the Exploration, Utilization and Conservation of Plant Genetic Resources’ (Table 1
. Some of the major recommendations of the 1967 conference included the need to survey genetic resources in nature and in genebanks and the need for a stronger emphasis on conservation, efficient documentation, and the improved international coordination of PGR activities. It also generated important guidelines for the establishment of a global network for ex situ long-term conservation. It should also be noted that in situ conservation, especially of landraces, was a big issue, but it was given little to no importance compared to ex situ conservation 
In 1971, the second international conference on crop genetic resources was held in Rome, and its proceedings were published in the book Crop Genetic Resources for Today and Tomorrow
, which included a plan of action (Table 1
. At this conference, the panel of experts made some major contributions with respect to global conservation plans, including the formulation of basic criteria for the conservation and the use of genetic material. These were: (i) that plant material was to be made available immediately and without restriction to all breeders requesting it and (ii) that genetic variability had to be maintained for future generations in long-term storage under conditions for maximum physical and genetic security. A third important result of the panel was a categorization of ex situ collections: base collections (for long-term conservation), active collections (for research and distribution), and working collections (usually maintained at plant breeding institutions) (for details, see 
. They also identified regions and crops for priority collecting 
. These collecting priorities were reformulated during the panel’s last meeting in 1975, with a clear shift from crops to regions 
The third international conference on crop genetic resources was held in Rome in 1981, jointly organized by FAO, UNEP, and IBPGR (Table 1
. The conference addressed most of the routine genebank operational topics, including sampling, seed storage and viability monitoring, recalcitrant seeds, in vitro conservation and the genetic stability of cultures, principles of germplasm regeneration, in situ conservation, the use of back-garden and genetic reserves for regeneration, the principles and practice of germplasm distribution and exchange, the safe and rapid transfer of plant genetic resources, including a proposal to distribute only germplasm materials completely free from plant pests and pathogens, principles of characterization and evaluation, data capturing and germplasm documentation, and under-exploited and minor crops 
The fourth technical conference was in the context of the FAO global system for the conservation and use of plant genetic resources and was held in Leipzig, Germany in 1996 (Table 1
. The major outcome of this conference was the Global Plan of Action (see below) and, in addition, ample information on the global conservation system 
The rising concern regarding the genetic erosion of landraces and wild relatives due to modern agriculture, and the more general, increasing need of the agro-industry for a steady flow of new germplasm convinced the members of the FAO conference to give more consideration to a generalist approach to conservation 
. During the second conference, the availability of new cold-storage techniques was noted, thus allowing long-term ex situ storage to be undertaken, whereas advocated in situ conservation, based on genecological premises, did not materialize until much later. The focus remained on ex situ conservation, despite the arguments for in situ approaches 
It should be noted that during the 1960s, the discussions on PGR in general as well as within FAO were dominated by plant breeders, and this resulted in a close conceptual link between conservation and use. Moreover, germplasm was predominantly stored in industrial countries and was closely tied to plant breeding institutes. During 1967, the FAO unit of Crop Ecology and Genetic Resources was established and thus provided FAO with more in-house specialized expertise.
2.2. The Establishment of the International Board for Plant Genetic Resources (IBPGR)
During a meeting of the Technical Advisory Committee (TAC) of the CGIAR in Beltsville, USA, a group of invited external experts, including several members of the FAO panel of experts, presented an ambitious plan to establish a world network of genetic resources centres 
. This plan consisted of four elements. The first one was to establish a coordinating centre (to become IBPGR); the second one was to stimulate the establishment of genebanks in already existing international centres in developing countries (i.e., IRRI, established in 1960; CIMMYT (1966); CIAT (1967); and IITA (1968). The third element was to establish genebanks in new international centres (WARDA, 1971; CIP, 1971; and ICRISAT, 1972). Soon thereafter, the ILCA was established in 1974, and ICARDA was established in 1976. The fourth element was the establishment of new ‘regional’ centres in the Vavilovian centres for crop diversity. The establishment of the International Board for Plant Genetic Resources (IBPGR) took place in 1974, as a secretariat for its board of trustees, administered by FAO and technically as one of the international institutes of the CGIAR. It was expected to coordinate global exploration and collecting efforts and to orchestrate a global network of genebanks (see also the details of this international undertaking below). Its main task was formulated as ‘to promote and assist in the worldwide effort to collect and conserve the plant germplasm needed for future research and production
The main achievements of IBPGR and its successor institute IPGRI, particularly those related to long-term conservation and the global conservation system, are updated from a list in 
Organization of collecting missions, partly using consultants in addition to its own staff and through contracts with national (selected) genebanks (for details, see IBPGR Annual Reports, e.g., 
; for an overview: 
Support for national and regional PGR programmes, predominantly in developing countries with the establishment of conservation facilities, documentation systems, and capacity building/training 
Establishment of regional and global PGR networks with national programmes as principal stakeholders as well as regional and global crop networks, frequently with and through CGIAR centres and their leading roles in crop-specific conservation and breeding, thus trying to ensure a close link between conservation and use. The European Cooperative Program for Plant Genetic Resources (ECPGR), formerly the ‘European Cooperative Programme for Crop Genetic Resources Networks’—ECP/GR), was founded in 1980 on the basis of the recommendations of the United Nations Development Programme (UNDP), the Food and Agriculture Organization of the United Nations (FAO), and the Genebank Committee of the European Association for Research on Plant Breeding (EUCARPIA); its secretariat was hosted by IBPGR 
The establishment of an international network of base collections in 52 selected genebanks located in almost 40 countries across all continents for the long-term conservation of crops or crop groups, including 80 genera and approximately 250 species 
, and the so-called Registry of Base Collections containing a total of 144,000 accessions 
Support for an international MSc course in the conservation and use of PGR at the University of Birmingham and the organization of training courses 
Establishment of a digitalized information system for genebank documentation and germplasm management.
Initiating, coordinating, and/or conducting plant genetic resource conservation and use research and publishing the results and procedures.
More recently, the successor institutes of IBPGR (IPGRI and Bioversity International), especially after their administrative separation from FAO, played an active role in developing legal and policy proposals and acted as the CGIAR representative in international meetings and activities.
2.3. The International Undertaking (IU)
The International Undertaking (IU) was established by the FAO Commission on PGR in 1983 as a non-binding intergovernmental agreement to promote the conservation, exchange, and use of plant genetic resources 
. Its objective was to ensure that plant genetic resources of economic and/or social interest, particularly for agriculture, would be explored, preserved, evaluated, and made available for plant breeding and scientific purposes. The Undertaking was based on the universally accepted principle that plant genetic resources are a heritage of mankind and, consequently, should be available without restriction. It defined ‘plant genetic resources
’ as the reproductive or vegetative propagating material of the following categories of plants: (i) cultivated varieties (cultivars) in current use and newly developed varieties; (ii) obsolete cultivars; (iii) primitive cultivars (landraces); (iv) wild and weedy species, near relatives of cultivated varieties; (v) special genetic stocks (including elite and current breeder lines and mutants)
. It defined ‘base collection of plant genetic resources
’ as a collection of seed stock or vegetative propagating material (ranging from tissue cultures to whole plants) held for long-term security in order to preserve the genetic variation for scientific purposes and as a basis for plant breeding
; ‘active collection
’ was defined as ‘a collection which complements a base collection, and is a collection from which seed samples are drawn for distribution, exchange and other purposes such as multiplication and evaluation
’, and ‘centre
’ was defined as an institution holding a base or an active collection of plant genetic resources 
Furthermore, the IU foresaw the development of a global system as to ensure that (Article 7.1):
A well-coordinated international network of national, regional, and international genebanks, including the international network of base collections, would develop. The unrestricted availability of materials included in the active and base collections of such a network was assumed.
Through the progressive growth of the network, comprehensive coverage of species and regions was aspired, and an adequate safety duplication of the germplasm was involved.
The exploration, collection, conservation, maintenance, rejuvenation, evaluation, and exchange of plant genetic resources should be conducted by the genebanks in accordance with scientific standards.
Adequate funding should be provided.
A global information system should be developed.
Genebanks should give an early warning to the FAO in the case of hazards that threaten the efficient maintenance of the collection.
IBPGR is expected to liaise with FAO while conducting its programme of work aiming at building institutional and human capacity within developing countries for the development and distribution of improved crop varieties.
Article 7 of the IU on International Arrangements addresses aspects of the global system and access to germplasm in the base collections. Countries are invited to notify the FAO in case their base collections are to be recognized as part of the international network of base collections. The participating genebanks are expected to make the materials in these base collections available to the participants in the IU for the purposes of scientific research, plant breeding, or conservation, free of charge and based on mutual exchange or mutually agreed on terms 
The IU was replaced by the International Treaty on Plant Genetic Resources in 2002 (see further below).
Another component of the global system is the International Code of Conduct for Plant Germplasm Collecting and Transfer 
. It was adopted by the FAO Conference at its 27th session in 1993. The voluntary code aims to promote the rational collecting and sustainable use of genetic resources to prevent genetic erosion and to protect the interests of both germplasm collectors and donors. It is based on the principle of national sovereignty over PGR and is in harmony with the CBD 
2.4. The Convention on Biological Diversity (CBD)
The negotiation of the Convention on Biological Diversity (CBD) in the eighties and early nineties, under the auspices of the United Nations Environment Programme 
, did result in drastic changes with respect to the conservation and use of PGRFA. Besides creating a general, globally, and legally binding framework for the conservation and sustainable use of biodiversity, the CBD, which entered into force in 1993, required that access to valuable biological resources must be conducted on ‘mutually agreed terms’ and is subject to ‘prior informed consent’ of the country of origin. The national sovereignty of states over biodiversity within their borders was recognized as a key principle in the CBD, and consequently, this became the ‘driving force’ in the thinking and approaches to the negotiations and future developments. Besides the fact that states were expected to ‘look after their own biological resources and conserve them, whenever possible in their own country’, this also caused a strong incentive for countries to favour bilateral rather than multilateral arrangements for the exchange of genetic resources.
From an agricultural perspective, it should be noted that the negotiations of the CBD were strongly influenced by environmentalists and nature conservationists and, consequently, a bias towards wild (i.e., non-domesticated and non-agricultural) plant and animal species could be observed 
. In fact, agriculturalists were hardly present in the negotiations, and it was only through a separate resolution (Resolution 3 of the Nairobi Final Act) 
that the FAO was asked to address two important but unresolved agricultural genetic resources issues, i.e., the question of Farmers’ Rights and the need to address the legal status of existing genetic resource collections established prior to 1993 
The negotiation process of the CBD caused a dramatic shift concerning the overall conservation approach, i.e., from a rather technologically driven ex situ conservation approach (‘putting the germplasm safely away for the future’), towards a much more people-centred conservation, with a strong emphasis on in situ and on-farm conservation and sustainable use efforts. Alongside this, due attention was being paid to participatory research (and conservation) activities to recognize the important role of local communities in the management of and their dependency on biodiversity. This also led to the recognition of traditional and indigenous knowledge to be an important component of biodiversity that needs to be collected and/or conserved. The importance of technology for the conservation and use of genetic resources should be recognized as well as the provision of access to such ‘enabling’ technologies. These aspects facilitated (and required) a much closer link between conservation and development and led to greater participation of local communities and subsistence farmers in conservation and use related activities. It is against this background that the access and benefit-sharing guidelines were developed and agreed upon in 2002 within the framework of the CBD by an Ad Hoc Open-Ended Working Group on Access and Benefit-Sharing 
that eventually, in 2010, resulted in the adoption of the Nagoya Protocol on Access and Benefit Sharing (ABS), which entered into force in 2014 
. It is a supplementary agreement to the CBD convention of 1992 and aims at the implementation of one of the three objectives of the CBD: the fair and equitable sharing of benefits arising out of the use of genetic resources, thereby contributing to the conservation and sustainable use of biodiversity 
. Its rather strong focus on wild species and the bureaucracy involved to apply the protocol have resulted in concerns that the added bureaucracy and legislation could be damaging to the monitoring and collecting of biodiversity, conservation, and research because the protocol severely limits access to genetic resources.
The CBD recognizes the application of intellectual property rights (IPRs) on biological materials as a means of protecting inventions and stimulating innovation. This led to a further expansion of the scope and/or application of IPRs, especially patents and plant breeder rights (PBRs), in agricultural research and plant breeding. Due to concerns that the development and use of genetically modified varieties could cause a threat to the environment and its biological resources, a legal framework on biosafety aspects was demanded, and thus, the so-called Cartagena Protocol on Biosafety was developed and came into force in 2003 as a legal framework for biosafety legislation and is yet another supplementary agreement of the CBD 
At present, the negotiation process on the development of the post-2020 global biodiversity framework is ongoing for its adoption during the forthcoming meeting later in 2021 in Kunming, China 
2.5. Global Plan of Action (GPA)
The first Global Plan of Action (GPAI) for conserving and using crop diversity was adopted in 1996 by 150 countries 
. The GPAI called for a rational global conservation system based on the principles of effectiveness, efficiency, and transparency. The Second Global Plan of Action (GPAII) reiterated that call and provided a strategic framework for the conservation and sustainable use of plant genetic diversity. It was adopted by the FAO Council in November 2011 and reaffirmed the commitment of governments to the promotion of plant genetic resources as essential components of food security through sustainable agriculture in the face of climate change (Table 1
. It is a rolling action plan and is based on the findings of the Second Report on the State of the World’s PGRFA 
and inputs from a series of regional consultations and from experts. The GPAs are a supporting component of the International Treaty on Plant Genetic Resources for Food and Agriculture 
The GPAII consists of four main groups of priority activities, i.e., in situ conservation and management, ex situ conservation, sustainable use, and building sustainable institutional and human capacities 
. The in situ conservation group of four priority activities comprises: 1. surveying and inventorying PGRFA; 2. supporting on-farm management and improvement of PGRFA; 3. assisting farmers in disaster situations to restore crop systems; and 4. promoting in situ conservation and management of crop wild relatives and wild food plants. The ex situ group of priority activities includes: 5. the targeted collecting of PGRFA; 6. sustaining and expanding ex situ conservation; and 7. regenerating and multiplying ex situ accessions. The sustainable use priority activities consist of: 8. the characterization and evaluation and development of subsets of collections to facilitate use; 9. plant breeding, genetic enhancement, and base broadening; 10. promoting the diversification of crop production and broadening crop diversity; 11. the development and commercialization of varieties, primarily of farmer varieties/landraces and underutilized species; and 12. supporting seed production and distribution. The set of capacity building activities comprises: 13. building and strengthening national programmes; 14. promoting and strengthening networks for PGRFA; 15. constructing and strengthening comprehensive information systems; 16. developing and strengthening systems for monitoring and safeguarding genetic diversity and minimizing genetic erosion of PGRFA; 17. building and strengthening human resource capacity; and 18. promoting and strengthening public awareness of the importance of PGRFA 
The GPAII does not contain specific activities related to long-term conservation and the global system, but several comments and supporting actions are referred to throughout the text, e.g., that the network of international ex situ collections of major crops played an important role in the negotiations of the International Treaty. These collections continue to form the backbone of the global system. The Svalbard Global Seed Vault now provides an additional level of security to existing ex situ collections 
. Furthermore, the development of a global portal of accession-level data and the imminent release of an advanced genebank information management system (recently released and called GLIS) are additional important steps towards the strengthening and more effective operation of a global system for ex situ conservation 
. Enhancing capacity at all levels is a key strategy to implement the priority activities of the GPA, including those related to long-term conservation, sustainable use (i.e., plant breeding, genetic enhancement, and base-broadening efforts) and the global system. Whereas countries have national sovereignty over and responsibility for the PGRFA they conserve, there is nevertheless a need for the greater rationalization of the global system for ex situ collections. The fostering of partnerships and synergies among countries is a requirement to develop a more rational and cost-effective global system. Furthermore, the GPAII plays an important role in the international policy framework for world food security and as a supporting component of the International Treaty. It contributes to achieving the Millennium Development Goals and aids in the implementation of the Strategic Plan for Biodiversity 
2.6. International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA)
The International Treaty for Plant Genetic Resources for Food and Agriculture (ITPGRFA or Treaty) aims to recognize the enormous contribution of farmers to the diversity of crops that feed the world; it aims to establish a global system to provide farmers, plant breeders, and scientists with access to plant genetic materials; and it aims to ensure that recipients share the benefits they derive from the use of these genetic materials with the countries where they originated 
. The preparations and negotiations of the revision of the IU were initiated in 1994 and were concluded in 2001 by the adoption of the International Treaty. It encompasses all PGRFA and came into force in 2004 
Through the Treaty, countries agree to promote the development of national integrated approaches to the exploration, collecting, characterization, evaluation, conservation, and documentation of their PGRFA, including the development of national surveys and inventories 
. They also agree to develop and maintain appropriate policies and legal measures to promote the sustainable use of these resources, including on-farm management, strengthening research, promoting plant-breeding efforts, broadening the genetic bases of crops, and expanding the use of locally adapted crops and varieties and underutilized species. These activities would be supported, as appropriate, by international cooperation provided in the Treaty.
The most important part of the ITPGRFA is the establishment of the so-called Multilateral System (MLS) of Access and Benefit-Sharing 
. The MLS applies to 64 genera, including the major food crops and forages, which were agreed upon on the basis of two criteria: their importance for food security and the level of interdependence among countries. At the global level, these crops provide approximately 80% of the food that is produced by plants. Through the MLS, sovereign nations have agreed to share resources and benefits. The genetic resources included in the MLS will be made available for research, breeding, and training, and their recipients should not claim any intellectual property or other rights that limit access to these resources or their genetic parts or components in the form received from the MLS 
. The peculiarities of PGRFA compared to biodiversity in general, e.g., the difficulty of applying the country-of-origin concept, the strong interdependency of nations on genetic diversity for crop improvement, and the critical role of these resources in traditional agriculture and in food security, formed the basis for the establishment of a multilateral rather than a bilateral system for their exchange 
. This thinking eventually led to the establishment of the MLS, which keeps the genetic resources of Annex 1 listed species that are formally in the public domain and under governmental control and facilitates easy access to and the use of these resources 
. It should be noted that the diversity of the crop species or the groups of species listed in Annex I is rather limited and, for instance, the majority of the vegetable genetic resources conserved by the World Vegetable Center in Taiwan, which consist of a large proportion of indigenous vegetables that are critically important for the diversification of cropping systems, nutritional security, and livelihoods 
, are not included in Annex 1. Discussions on the extension of the Annex 1 list have been ongoing for several years, but no final decision has been reached.
The benefits arising from the use of materials from the MLS shall be shared fairly and equitably through the exchange of information, access to and transfer of technology, and capacity-building, considering the priority activity areas indicated in the above mentioned GPAII and under the guidance of the Governing Body of the Treaty. It further establishes the payment, which is in certain cases mandatory, of an equitable part of the monetary benefits that are derived from the use of PGRFA into the funding strategy of the Treaty 
. The funding strategy aims at mobilizing funds for activities, plans, and programmes to support the implementation of the Treaty and, in particular, its implementation in developing countries while keeping in line with the priorities that have been identified in the GPA. The funding strategy includes the monetary benefits that are paid in accordance with the MLS as well as the Global Crop Diversity Trust, which is described below. The Treaty recognizes the enormous contributions that local and indigenous communities and farmers of all regions of the world have made and will continue to make for the conservation and development of PGRFA. The Treaty makes governments responsible for the realization of Farmers’ Rights, including the protection of relevant traditional knowledge; provisions for farmers to participate equitably in sharing benefits; and farmer participation in national policy decision-making 
. Through Article 15, the Treaty establishes its relationship with the CGIAR and other international centres: ‘Ex Situ Collections of Plant Genetic Resources for Food and Agriculture held by the International Agricultural Research Centres of the Consultative Group on International Agricultural Research and other International Institutions
’ and arranges that the materials listed in Annex 1 of the Treaty and that are held by the centres as well as other species than those listed in Annex 1 of this Treaty and collected before its entry into force that are held by IARCs shall be made available in accordance with the provisions of the standard material transfer agreement (SMTA) 
2.7. International Network of Ex Situ Collections
The international network of ex situ base collections in genebanks that are managed by national, regional, or international centres was a component of the section on the international arrangements of the International Undertaking. It was foreseen that through a steady increase of the number of genebanks participating in the network, adequate coverage in terms of species and geographical distribution would eventually be achieved. It was further foreseen in the IU to conclude agreements (four ‘model agreements’ were available to choose from) with countries to place their base collections within this network and/or to provide storage space for the long-term storage of base collections from elsewhere. A few countries and institutions made concrete offers to place (part of) their collections in the network. The latter would operate under the auspices and/or the jurisdiction of the FAO and a number of contracts were concluded (see below).
In 1994, the CGIAR centres expressed the wish that their designated germplasm be recognized as part of the international network of ex situ collections and signed individual agreements with FAO 
; Chapter 3.1. in 
. The salient features of these agreements based on one of the above-mentioned model agreements include that:
The centre shall hold the designated germplasm in trust for the benefit of the international community.
The centre shall not claim legal ownership over the designated germplasm, nor shall it seek any intellectual property rights over that germplasm or its related information.
The designated germplasm shall remain in the charge of the centre.
The FAO shall have a right of access to the premises at any time and has the right to inspect all activities performed therein.
The centre shall undertake the management and the administration of the designated germplasm in accordance with internationally accepted standards with respect to the storage, the exchange and distribution of seeds, the international genebank standards endorsed by the Commission and that all designated germplasm is duplicated.
The centre recognizes the intergovernmental authority of the Commission in setting policies for the International Network and shall undergo consultation with the FAO and its Commission on proposed policy changes related to the conservation of the germplasm.
The centre shall undertake the creation of samples of the designated germplasm and will make related information available directly to users or through the FAO for the purposes of scientific research, plant breeding, or genetic resource conservation without restriction.
The centre shall ensure that such other people or institutions and any further entity receiving samples of the designated germplasm from such a person or institution are bound by the conditions to not claim ownership over the materials or to seek any intellectual property rights over that material and, in the case of samples duplicated for safety purposes, to manage these in accordance with internationally accepted standards.
A related network, as mentioned above, is the so-called Register of Base Collections that was established by the IBPGR in the 1970s and includes genebanks that were prepared to accept a long-term commitment to conserve germplasm materials and to make these available to users. This register formed the backbone of the international network of base collections. For details, see the paper by Engels and Thormann 
It should be noted that further agreements have been concluded with several other international research centres (e.g., the World Vegetable Center, CATIE and CRU, and some regional organizations (e.g., South Pacific Community)). Agreements with individual countries have not been vigorously pursued. In October 2006, 11 CGIAR centres signed agreements with the Governing Body of the International Treaty to bring their in trust collections under the framework of the Treaty and to recognize the authority of the Governing Body providing policy guidance related to those collections 
With the establishment of the International Treaty and its Multilateral System, the network of ex situ collections, and the conclusion of the agreements with the centres of the CGIAR, these collections were brought under the International Treaty (Chapter 3.2 in 
). The commitments of countries to conserve germplasm for the long-term and to make the materials available (under an SMTA) have been made by countries and genebanks through the inclusion of germplasm in the MLS.
3. Description of Ex Situ Germplasm Conservation Methods and Their Strengths and Weaknesses
The vast majority (approx. 92%) of angiosperms comprising roughly 330,000 species of flowering plants has desiccation-tolerant and so-called orthodox seeds 
that survive drying to a low moisture content, 5% or less, and subsequent rehydration without a significant loss of viability 
. Orthodox seeds acquire desiccation tolerance during their late phase of development when they undergo pre-maturation drying and are later shed metabolically inactive 
. Desiccation tolerance is lost during germination 
. Moreover, most desiccation-tolerant species tolerate low temperature (sub-zero) storage and seed longevity increases, within certain limits, with a decrease in seed moisture content (SMC) and storage temperature 
. Harrington 
postulated two rules of thumb regarding seed longevity in storage that apply independently. Over the range of 14 to 4% SMC (fresh weight basis), a 1% reduction in SMC doubles the life span of the seed. Similarly, within the range of 50 to zero degrees Celsius, for each 5 °C drop in storage temperature, the life span of seed in storage would double. Therefore, the cold storage of dried seeds is a practical, efficient, and cost-effective method for the long-term storage of germplasm in genebanks. The FAO Genebank Standards recommend storage at −18 ± 3 °C and relative humidity of 15 ± 3 per cent for most original seed samples and safety duplicate samples intended for long-term storage 
. In case seed samples are stored in hermetically sealed pouches or containers, the control of the storage room RH is not required.
In contrast to orthodox seeds, so-called ‘recalcitrant’ seeds are desiccation-sensitive and rapidly lose viability upon drying and do not tolerate low-temperature storage 
. Recalcitrant seeds undergo extremely limited drying during maturation and consequently, have high SMC and are metabolically active during shedding 
. Desiccation sensitivity also seems to be linked to the non-dormant state of seeds upon shedding 
. The SMC below which viability is lost varies between species but is generally above 20% 
. Specifically, tree species of tropical provenance, such as avocado (Persea americana
), cacao (Theobroma cacao
), jackfruit (Artocarpus heterophyllus
), breadfruit (Artocarpus altilis
), lychee (Litchi chinensis
), mango (Mangifera indica
), mangosteens (Garcinia mangostana
), etc., produce recalcitrant seeds.
There is a third category of seed storage behaviour comprising so-called ‘intermediate’ seeds without sharp boundaries between orthodox and recalcitrant seeds 
. Species with intermediate seed storage behaviour can be dried to certain SMC levels but cannot be dried to a level as low as truly orthodox seeds 
and often do not survive sub-zero storage temperatures. Moreover, seeds with intermediate storage behaviour tend to lose viability much quicker than orthodox seeds 
. Coffee (Coffea arabica
) seeds fall into this category of intermediate seeds 
. Depending on the cultivar, coffee seeds tolerate drying to 5–10% SMC but viability at low or sub-zero temperatures is rapidly lost. Seeds of alpine species are also significantly shorter-lived than their lowland counterparts, possibly due to abnormal seed development under the cool and wet conditions of the alpine climate 
As species producing seeds with intermediate or particularly recalcitrant storage behaviour have extremely limited longevity in a seed genebank, they are commonly stored in field genebanks and/or as in vitro collections for medium-term conservation and/or in liquid nitrogen for long-term conservation.
3.1. Short-, Medium- and Long-Term Ex Situ Storage of Orthodox Seeds
In general, orthodox seeds are relatively small and require little storage space for the conservation of a representative sample of the source population and further sub-samples for distribution, viability checking, and safety backup. Crops commonly conserved in seed genebanks include cereals such as rice, wheat, barley, oats, sorghum, millet, maize, grain and forage legumes, most vegetables, and some fruit crops. True seeds of crops such as those from potato, which are commonly propagated vegetatively, can also be dried and stored at low temperatures 
. This is common practice with wild potato germplasm, and accessions are maintained as botanical seeds or true-potato seeds (TPS). A representative number of 20–50 individuals are typically collected from a wild population, and seeds are regenerated and combined to form a unique genebank accession of heterogeneous seed, which is expected to represent most alleles found in that population 
. Seed samples of such wild potato germplasm accessions thus represent a heterogeneous mix of genotypes, whereby each genotype represents a portion of the genetic make-up of the sampled population.
The core operations of a genebank conserving the seeds of orthodox species comprise cleaning, seed drying, viability and health testing, packing, storage, and distribution to users and for a safety backup 
. When seed stocks are running low or when seed viability drops below a minimum threshold, seed lots need to be regenerated for seed replenishment. All these genebank operational steps are documented and in many genebanks are supported by a genebank information system 
Most genebanks conserving PGRFA have the mandate to distribute germplasm to a range of different users and, for practical reasons, store the seeds of most collected or acquired accessions in a base and an active collection when justified. The most original seed samples are kept in the base collection for long-term conservation, aiming at the highest level of genetic integrity of the stored sample with the original sample 
. The active collection is oriented towards seed regeneration (triggered by low viability), characterization, evaluation, multiplication (triggered by low seed stock), and distribution and is generally kept under medium-term storage (MTS) conditions.
The base collection for any given species or a crop genepool may be distributed over several institutions, as is the case in Europe, with the implementation of a European Genebank Integrated System, abbreviated as AEGIS 
. In contrast, the United States Department of Agriculture (USDA-ARS) has a network of genebanks holding the active collections for different crops in 19 different locations across the country, with one main base collection held at the National Laboratory for Genetic Resources Preservation (NLGRP) in Fort Collins, Colorado, serving all of the regional genebanks. The NLGRP maintains the US system backup of more than 445,000 accessions, representing 86% of the seed collections and 15% of the clonal collections 
. Seeds are not distributed from the base collection to the users, but rather, they are distributed from the active collections.
The active collections comprising the bulk of orthodox seeds stored in most genebanks are to be kept under medium-term storage (MTS) conditions at temperatures ranging from 5 °C to 10 °C and at a relative humidity (RH) of 15 ± 3 per cent for seeds that are stored in open containers 
. Frequently, MTS conditions have a narrower range from +2 to +5 °C 
, and RH adjustment is not required if seeds are stored in hermetically sealed pouches or containers. Refrigerated seed storage under MTS conditions is adequate for up to 30 years 
. It should be noted that seeds stored in hermetically closed containers are to be dried in a controlled environment with a temperature range between 5 and 20 °C and a RH between 15 and 25%, depending on the species.
The base collections are stored under long-term storage (LTS) conditions at sub-zero temperatures of typically −18 to −20 °C 
, and the seeds are dried as mentioned above for MTS, maintaining high seed quality over long, species-specific periods of up to 100 years or more.
Other genebanks whose major focus is not the use of plant agrobiodiversity facilitation but rather whose focus is on the long-term conservation of globally threatened species (with relatively few sample requests), store all of their seeds exclusively under LTS conditions. This applies, for example, to the Millennium Seed Bank (MSB) of the Royal Botanic Gardens Kew, where dried seeds are transferred to air-tight glass containers or aluminum foil bags and are stored in the seed vault at −20 °C 
Assessing 42,000 seed accessions representing 276 species in the USDA National Plant Germplasm System provided evidence that some species produce orthodox seeds of short longevity in dry storage 
. Some plant families had typically short-lived seeds (e.g., Apiaceae and Brassicaceae) or long-lived ones (e.g., Malvaceae and Chenopodiaceae). Moreover, environmental factors seem also to determine seed longevity, as seeds from species originating from certain localities in Europe had short shelf lives, while seeds of the same species originating from localities in South Asia and Australia had much longer shelf lives. For these reasons, some genebanks additionally cryopreserve samples of those orthodox seeds that are expected to be very short-lived, even under LTS conditions 
Under short-term storage (STS) conditions, the seed quality and the viability of orthodox seeds with long shelf lives can be maintained for a minimum of eight years under ambient conditions if 25 °C is not exceeded, and the relative humidity in the storage room is kept at 10–25% 
. At the World Vegetable Center in Taiwan, working collections of breeders and other researchers are kept in STS conditions at 15 °C and 40–45% RH 
3.2. Field Genebanks
Although seed desiccation sensitivity affects only about 8% of flowering plants 
, there are many field and horticultural crops as well as (agro)forestry species that cannot be conserved long-term in conventional seed storage and that require different forms of conservation, such as in field genebanks, in in vitro collections, and/or in liquid nitrogen 
. Among those are species that only produce recalcitrant or intermediate seeds with a short storage life span. Moreover, some species take several years to produce seeds, such as yucca (Yucca
sp.) and bamboo (a species of the Poaceae subfamily Bambusoideae), while other crop species hardly produce seeds and are only vegetatively propagated, such as edible banana and plantain (Musa
Major food crops that are commonly clonally propagated and therefore conserved in field genebanks include herbs, shrubs, vines, and trees, and these food crops belong to about 34 families 
. Among those are sub-tropical and tropical shrub and tree species, such as coffee (Coffea
sp.), cacao (Theobroma cacao
), rubber (Hevea brasiliensis
), coconut (Cocus nucifera
), peach palm (Bactris gasipaes
), breadfruit (Artocarpus altilis
), mango (Mangifera indica
), citrus (Citrus
sp.), avocado (Persea americana
) many temperate fruit trees, root and tuber crops such as potato (Solanum tuberosum
), cassava (Manihot esculenta
), yams (Dioscorea
sp.), sweet potato (Ipomoea batatas
), taro (Colocasia esculenta
), other aroids, bananas, garlic (Allium sativum
), shallot (Allium cepa
), grasses such as sugarcane (Saccharum officinarum
), and forages. Additionally, temperate and sub-tropical fruit trees like peach (Prunus persica
) and apricot (P. armeniaca
) are typically clonally propagated to maintain the genetic constitution of the variety. As their seeds are non-orthodox, i.e., they cannot be dried to low seed moisture content and thus cannot be stored for longer periods at low temperatures, they are maintained in field genebanks and increasingly as in vitro materials (see Section 3.3
) or cryopreserved (see Section 3.4
). Although some of those crops are sexually fertile, they do not breed true to type, hence, the preferred method is vegetative propagation which enables the maintenance of genotypes as clones.
In field genebanks, the plant genetic resources are kept as live plants that undergo continuous growth and require regular care and maintenance. Accessions maintained in field genebanks need considerable space, especially tree species, and require much more attention in their day-to-day management than seed or in vitro collections, as the plants are continuously exposed to biotic and abiotic stresses. Integrated pest and disease measures are essential to ensure that plants are free of pathogens 
Given the exposure of plants in field genebanks to biotic and abiotic stresses and physical security threats (invading animals, theft), these do not present the most secure methods of germplasm conservation; however, they are often the only practical and cost-effective choice to conserve the germplasm of clonal crops, especially when resources and skills for alternative conservation approaches, such as in vitro conservation or cryopreservation, are out of reach.
When field genebank conservation is the only viable alternative, careful planning of site selection and appropriate field management can help to mitigate those risks. The revised and updated Genebank Standards of the FAO 
indicate the best practices for the safe establishment and management of field genebanks, including the choice of location, the acquisition of germplasm, the establishment of field collections, appropriate field management, the regeneration and propagation of plant material, characterization, evaluation, documentation, distribution, and security and safety duplication.