Task 1. Assess the applicability of the various conservation methods in your country Some conservation methods (cryoconservation in particular) will require special equipment and expertise. Lack of these resources will limit the options available. For example, liquid nitrogen can be a limiting resource for cryoconservation in many countries. Techniques for cryoconservation of germplasm also differ in their practicality across species and according to the type of germplasm to be conserved. Capacity to collect and freeze semen is available in many countries, whereas cryopreservation of pig embryos requires a high level of technical capacity. Ex situ in vivo conservation requires access to animal housing facilities and possibly land for grazing or crop production. An inventory of stakeholders and the available expertise, technology and facilities should be taken. All types of conservation require a long-term investment by stakeholders if they are to be successful. Commitment by stakeholders to cooperate with the government and with other stakeholders should be secured before embarking on a programme for which such cooperation will be fundamental.
Task 2. Match breeds to the most appropriate conservation methods 1. Identify the conservation objectives relevant for each breed.
For each breed, consider the question: why is this breed on the priority list for conservation? The answer may influence the choice of conservation method. For example, if the main reason is the breed’s contribution to the future genetic diversity of the species and to the genetic flexibility, then cryoconservation is the primary method of choice. If the main reason is to ensure the continuation of the breed’s present functions in rural areas, then in situ conservation is the preferable method.
Action 2. Rank potential conservation methods according to their efficacy Not all conservation methods will have the same efficacy with respect to the conservation objective.
To accommodate cross-breeding (e.g. to introgress some unique alleles), ex situ in vivo conservation is very efficient. A relatively small number of pure-bred animals are maintained at a central facility, while the breed’s genes are transmitted more widely in the commercial population. If a breed is to be cryoconserved for the purposes of regeneration at a later date, then collection and storage of the breed’s germplasm in the form of semen will, in most species, be less expensive than the collection and storage of embryos. However, in the long term, breed regeneration using semen will be more time-consuming than regeneration using embryos, because in the former case, several generations of backcrossing are needed in order to obtain the conserved breed in a nearly pure state. When a breed is conserved in situ, farmers’ enthusiasm for keeping the breed (and thus the success of the programme) will depend strongly on the breed’s productivity and market prices for its products. Improving these aspects will likely increase efficacy of in situ conservation (see Sections 7 and 8).
Action 3. Consider the possible pitfalls and risk of failure of each conservation method and discard approaches with unacceptably high risk When a breed is conserved in situ, risk factors include:
• disasters and infectious diseases, which may destroy the population, especially if the breed is concentrated within a small geographical area;
• disconnection between the management and the operation of a conservation programme;
farmers have the right to operate their farms according to their own prerogatives and may even decide to abandon the breed if maintaining it is not financially attractive;
• genetic bottlenecks and high relationships between animals;
small populations are at greater risk of inbreeding and loss of alleles by random drift if the population is not maintained correctly;
and CGRFA/WG-AnGR-7/12/Inf.6 • changes in government programmes;
when a breed is used for purposes such as landscape management (see Section 8), subsidies may contribute a substantial amount of the breeders’ income, and termination of the subsidy may induce breeders to forsake the conservation of the breed.
When a breed is conserved ex situ in vivo, risk factors include the following:
• high relationships between animals may cause inbreeding and loss of alleles by random drift, leading to decreased genetic variability, and perhaps poor fertility, fecundity and viability;
• opportunities for improving the population through a breeding programme are limited, so farmers participating in the conservation scheme often have to be subsidized;
if the subsidy stops, the risk for loss of the breed increases;
and • populations conserved in vivo on government farms are vulnerable to changes in the financial priorities of the central government or the respective ministry.
When a breed is cryoconserved risk factors include the following:
• material for cryoconservation (gametes, embryos) must usually meet high sanitary requirements and animal disease may disturb or inhibit the collection of this material;
• freezing, maintenance and thawing of frozen material require special skills and reliable equipment and infrastructure;
lack of plans to maintain these supporting resources will put the stored germplasm at risk;
and • electrical blackouts and ruptures of storage vessels can result in loss of viability in the stored materials.
Action 4. Rank techniques for costs Once a decision has been taken as to which conservation methods have an acceptable level of risk, the costs of implementing these methods should be calculated. In the case of cryoconservation, the major cost consists of two parts: first, the collection and freezing of the material;
and second, the use of the material to meet the conservation objective (e.g. introgression of alleles or regeneration of the breed).
The maintenance costs of an animal gene bank are relatively low. The costs of in situ conservation may consist of subsidies provided to the keepers of the target breeds and the costs of realizing a breeding programme with special emphasis on the maintenance of the genetic variation. As noted above, many of these costs will recur for many years, and this must be accounted for.
Action 5. Choose the conservation method Finally, the rankings for efficacy, risk of failure and costs should be considered together. The weight given to each factor will depend on the country’s priorities and strategic preferences and the availability of resources, capacities and institutions. When artificial reproduction methods are well developed and widely applied, cryoconservation may be preferred. When only natural mating can be used to maintain the breed, in situ conservation is the first choice.
Task 3. Apply the chosen methods to reach the conservation objectives The remainder of these guidelines provides advice on establishing and operating in vivo conservation programmes. For cryoconservation, see Cryoconservation of animal genetic resources (FAO, 2012).
References FAO. 2012. Cryoconservation of animal genetic resources. FAO Animal Production and Health Guidelines. No. 12. Rome (available at http://www.fao.org/docrep/016/i3017e/i3017e00.htm).
CGRFA/WG-AnGR-7/12/Inf.6 V. ORGANIZING THE INSTITUTIONS FOR IN VIVO CONSERVATION The context in which an in vivo conservation programme is undertaken will vary greatly from country to country and from one species to another. Nevertheless, there are aspects that will be common among all programmes. Among the most important of these commonalities is the need for organization and for a plan for the sustainability of the programme. Organization is critical, because in most cases, many stakeholders will be involved in the programme. Although these stakeholders will have objectives of their own, they should all share the common goal of maintaining the breed in sufficient numbers to avoid its extinction or genetic erosion.
A wide array of stakeholders can contribute to breed conservation. For any individual breed, some types of stakeholders will be more important others, but the involvement of a range of stakeholders is usually critical to the long-term success of a conservation programme. Common stakeholders in the management of animal genetic resources include breeders (farmers and pastoralists), owners (farmers and pastoralists), users (e.g. of draught animals and breeding bulls), government institutions, breeders’ associations, breeding companies, research organizations, NGOs and animal genetic resource societies, consumers of livestock products and marketers (Oldenbroek, 2007;
Breeders usually own a significant proportion of the genetic resource, and are the most essential stakeholders. Most of the breeders will also be producers and they are of primary importance if the goal is to maintain the breed without economic subsidies (EURECA, 2010). “Buy in” by breeders is an absolutely essential component of any in vivo conservation programme. Success depends on the breeders having an understanding of and commitment to the conservation of pure-bred, viable populations. Successful conservation efforts generally involve multiple owners, working together for the survival of the breed. The pattern of ownership of animal genetic resources is distinct in important ways from that of plant genetic resources.
Breeders’ associations contribute in several ways to the conservation of animal genetic resources, including through participation in and communication with the National Advisory Committee (FAO, 2009 – see Section 1), serving as a source of information on the breed and its roles, product development and promotion, marketing and providing technical support for breeders. Associations manage herdbooks and performance recording and are centres of organization and support. They may, however, be a biased sample of owners, with a disproportionate share of larger herds and of herds with high levels of management and innovation.
Stakeholders that are not private owners can have important roles in conservation, but it is always important for them to work closely with private breeders. Generally, the non-private institutions (governmental bodies and NGOs) should support private efforts and the involvement of private breeders.
Especially in several Asian, South American and African countries, governmental breeding farms are important reservoirs of animal genetic resources, and make breeding animals and semen available to private farmers in situations where they would otherwise be unable to access selected breeding material. Such institutions also contribute to breed characterization and other research. They have a very real responsibility to ensure that their programmes lead both to short-term and long-term benefits for farmers. Where appropriate, such institutions should be strengthened or established.
Governmental organizations can be effective in promoting and rewarding cultural and social benefits provided by breeds. European countries increasingly recognize the value of locally adapted breeds of grazing animals in the management of natural areas and the maintenance of historically and culturally significant countryside. These values are difficult to recognize and reward via the private sector alone.
Many hobbyists keep and breed locally adapted breeds as a leisure activity. These non-production activities offer a great opportunity for breed conservation, but need institutional support to ensure proper conservation and management of genetic diversity.
Educational (university) and private research institutes also play roles in conserving breeds. These organizations can be especially important in providing the technical support needed to ensure that the genetic viability of small populations is maintained through proper attention to population structure and mating strategies. Private breeding companies likewise manage important breed populations in various species, although these resources may not be widely available for distribution. Many breeds or CGRFA/WG-AnGR-7/12/Inf.6 lines no longer developed for immediate commercial goals are set aside without long-term conservation plans.
This section gives the rationale, the tasks and the actions required by the different stakeholders with special emphasis for the breeding organizations.
Involving livestock keepers in community-based in situ conservation Rationale The conservation of animal genetic resources under sustainable management by farmers is one of the most effective and practical ways of conserving these resources with a minimum of inputs. This approach does not involve large financial expense and is feasible under field conditions. However, major attention should be paid to the economics of the targeted breeds. Breed conservation with the participation of farmers will be successful only if it is economically viable and if sufficient technical support is provided. Hence conservation projects aimed at the improvement and utilization of breeds should define their objectives clearly, especially with respect to the characteristics for which the breed has traditionally been valued. Conservation efforts should begin with characterization and evaluation of the breeds and identification of characteristics of economic, social and cultural value specific to each breed. Further advice on organizing such studies is provided in Phenotypic characterization of animal genetic resources (FAO, 2012b). A participatory approach, involving the farmers, breeders and other livestock keepers as much as possible is important, both to increase the accuracy of the information upon which the conservation programme will be based and to ensure interest and ownership of the programme on the part of the livestock keepers and thereby increase its sustainability. The livestock keepers and breeders will rarely accept a programme that deviates from their preferences (see Box 21). The role of an outside entity will primarily be to provide the inputs to improve the long-term survival of the target community and to provide technical support. FAO has produced a publication on community-based programmes for management of animal genetic resources that includes a number of practical examples (FAO, 2003a).
Box Conservation of Hallikar cattle in India The Timbaktu Collective is an NGO working in the Anantpur area in the State of Andhra Pradesh, India. Initially, their activities primarily dealt with improving the land and plant biodiversity in the drought-prone rainfed area. However, in 2005 they decided to expand their activities to cattle production because water availability had recently improved in the area and local farmers had begun cultivation. The Timbaktu team took the farmers to dairy centres in nearby towns and showed them the high milk producing Holstein-Friesian crosses. The farmers were not impressed, however. They opined that such animals would not suit the village production environment, and expressed their preference for the local breed – Hallikar – which used to be common in the area. The farmers knew that their main need was for draught power, with milk production as a secondary bonus. This opinion surprised the Timbaktu management, which had expected the farmers to want the higher producing cross-breeds, but they accepted the farmers’ opinion. In 2007 and 2008, about Hallikar cattle were purchased from the adjoining state and re-introduced to the Anantpur area.
Now, five years later, the whole experiment can be deemed a success. All animals are used for draught purposes in crop cultivation and the cows produce 1 to 2 kg of milk per day, which meets the needs and expectations of the farmers. The number of Hallikar cattle has increased significantly in the village since their reintroduction.
Provided by Devinder K. Sadana.
CGRFA/WG-AnGR-7/12/Inf.6 Objective: To design, with livestock keepers, an in situ conservation programme that they will implement with the assistance of outside agencies;
this ensures maintenance of the respective animal genetic resource because it promotes the autonomy of the community and the sustainability of the livelihoods of community members.
1. A breed at risk of extinction, but deemed of high value for conservation;
2. Basic knowledge of the location where the breed is raised and the lifestyle of the respective community of livestock keepers, their production system, their animals and their facilities;
3. An indication from the livestock keepers that they are interested in breeding and conservation;
and 4. Earmarked technical and financial resources.
• A sustainable in situ conservation programme based on the active participation of livestock keepers.
Task 1. Identify the specific location and collaborators for conservation activities Action 1. Pinpoint and study the geographic area where the breed is kept Using the background information on the targeted breed and results from breed surveys (FAO, 2011b), the core of the breed’s home region should be identified.
Action 2. Choose the communities with which conservation work will be undertaken Several villages distributed in different parts of the breed’s home region should be identified as candidates for participation in the conservation programme. The precise number of villages will depend on the population size of the breed to be conserved, its distribution across the area and across farms, and the resources available. If knowledge about the extent of livestock keepers’ interest in participating in a conservation programme is already available, this may facilitate the selection process. Clearly, a programme is more likely to be successful if livestock keepers and other stakeholders are keen to participate The livestock keepers who own the animals that are truest to the ideal type targeted for conservation (e.g. free from cross-breeding or having superior phenotypic traits) should be encouraged to participate.
Task 2. Undertake a detailed participatory study of the targeted communities Once the candidate communities have been chosen, the next step is to engage with them. In most cases, completing the activities described in Sections 1 to 3 will only have yielded the information needed to identify which breeds should be conserved and why. Much more information will be needed to develop a sustainable community-based programme for the conservation of these breeds. Community-based programmes will likely have to identify how breed conservation measures can complement the broader objective of improving the likelihoods of the community members. This will require multidisciplinary studies based on participatory approaches. Properly implemented, such approaches will not only offer an effective means of gathering information from the community, but also help to establish lines of communication that will facilitate collaboration during the subsequent implementation of the programme (Franzel and Crawford, 1987). General advice on participatory approaches in agriculture and rural development can be found in FAO (2003b). Their role in surveying and monitoring animal genetic resources is introduced in the respective guidelines in this series (FAO, 2011b). Specific examples of their use in animal genetic resource management include the work of Duguma et al. (2010).
Action 1. Undertake preparatory work including making initial contact with community leaders and agencies that may act as a liaison with the community A top-down approach in which a study team simply visits the community unannounced and undertakes the study will often not work well and is not recommended. A step-by-step process of engagement that informs the members of the community about the intentions and goals of conservation work and any complementary activities will usually be necessary. The first contact should be with the leaders of the village or with prominent farmers (also referred to in this context as “role model breeders” – see Section 8). It may be helpful to work with the assistance of an organization that already has experience working with the community, such as an NGO or government extension service.
CGRFA/WG-AnGR-7/12/Inf.6 Action 2. Enlist the research team and perform the studies At the community level, the problem of conserving animal genetic resources will likely be multi faceted, due to the large number of possible threats to breed sustainability (see Section 1). Thus the research team will need to address not only genetic factors, but also economic and social factors. It should therefore be multidisciplinary.
The study should collect information on a wide number of topics, including standard production and breeding practices, perceptions of the breeders on the strengths and weaknesses of the targeted animal genetic resource, outputs obtained from the animals and their use or marketing, sources of agricultural inputs and marketing opportunities. Constraints affecting the production system should be identified.
Much of this information should already be available if the activities described in Sections 1 to 3 have been undertaken. However, details particular to the communities targeted by the conservation programme will need to be obtained. In particular, the specific threats to the sustainability of the targeted breed should be identified in detail. Finally, an assessment of the willingness and capacity of community members to participate in conservation activities should be included.
Action 3. Evaluate the results The results of the participatory surveys should be evaluated in a comprehensive manner in order to determine the most practical and efficient means of conserving the targeted animal genetic resource. A follow-up meeting with the community members is recommended.
Task 3. Facilitate the implementation of an in situ conservation programme based on the participation of the community Figure 2 shows the possible interactions among stakeholders in a community-based programme for management and in vivo conservation of animal genetic resources. The ellipses indicate the major stakeholders (groups of livestock-keeping communities, the government, and breeders’ associations and other NGOs). Rectangular text boxes indicate the “goods” exchanged by each pair of stakeholders, with solid arrows showing the flow of these goods. Other stakeholders that may be involved in a community based conservation programme include the private sector (marketing of products and provision of inputs) and the general public (consumption of products). Box 22 explains how farmers, the government and a breeders’ association have worked together to conserve of an important breed in Argentina.
Box Public-private partnerships for conservation of Criollo Argentino cattle The Criollo Argentino is a local breed of cattle that evolved from a mix of the cattle breeds brought by the Spaniards to Argentina in the fifteenth century. Over time, the Criollo Argentino has co evolved with the local environment and its adaptation to a variety of conditions is remarkable.
However, the introduction of British breeds at the end of the nineteenth century confined the Criollo to the marginal regions of the country. This led to a reduction in the size of the breed’s population and drove it almost to extinction. In the 1960s, the National Institute of Agricultural Technology (INTA), a national research-extension organization, took on the task of rescuing and conserving the Criollo Argentino and promoted the creation of a breeders’ association, the Asociacin Argentina de Criadores de Ganado Bovino Criollo, which in a joint effort with INTA established a programme for the promotion of the breed. Subsequently, INTA and several universities have taken responsibility for conservation and characterization activities. In particular, INTA maintains a network of 12 active animal germplasm banks, five of which are devoted to the Criollo Argentino.
The primary method of choice for the banks is in vivo conservation, although one bank also employs cryoconservation. The institutional herds vary in size from 50 to 150 cows and are located in different agro-ecological regions across the country. INTA is planning to cooperate with the breeders’ association in genetic evaluation and characterization. The breeders’ association assists breeders in promotion and marketing.
Provided by Carlos Mezzadra.
CGRFA/WG-AnGR-7/12/Inf.6 Figure 2. Interactions among the possible stakeholders in a community-based breeding programme Community Community Community Exchange of animals • Extension • Performance and • AI and veterinary pedigree recording • Marketing assistance • Access to • Conservation of services • Microcredit • Training community animal genetic • Payment for ecosystem • AI and vet service members and resources • Microcredit • Environmental services animals • Social services • Data • Feed supplements services • Direct financial support • Data • Expertise and services Breeders’ associations and Government other NGOs • Infrastructure • Financial support As Figure 2 indicates, sometimes outside investment and assistance from the government or an NGO may be necessary in order to implement activities that help to ensure the sustainability of the targeted breed, especially in the initial phases of the programme. For example, construction of abattoirs or milk collection and processing facilities may increase livestock keepers’ access to markets and thereby increase their incomes and the economic sustainability their livestock-keeping activities. Outside assistance may be necessary to promote such opportunities among potential investors. Livestock keepers may also be willing to invest in technologies that improve productivity, such as AI, veterinary services or supplemental feeding, or in training to improve marketing (e.g. in cheese or yoghurt production), but may not have access to credit services through which to obtain investment funds.
Provision of such services may be a way of indirectly supporting in vivo conservation.
Action 1. Assist farmers in organizing a breeders’ association The animals belonging to breed population targeted for conservation will almost always be owned by a number of different people. Individual owners generally have the right to manage their animals as they see fit. However, breeders share some interests and some goals can be accomplished more easily as a group than by individuals. Therefore, formally organizing owners into a breeders’ association can yield benefits both for the individual owners and for the sustainability of the breed (see Box 23 for an example of the many possible benefits of a breeders’ association). It also creates a single entity with which the government or an NGO can work, which is likely to increase the efficiency of the conservation efforts. For long-term sustainability, it is imperative that breeders receive tangible benefits from being members of the association and that they drive the process. Detailed advice on establishing and monitoring breeders’ associations is presented later in this section.
CGRFA/WG-AnGR-7/12/Inf.6 Box The Banni Breeders’ Association – a case study from India The Banni Grassland in the Kachchh (Kutch) District of Gujarat in western India is the home of the Banni buffalo. The breed was developed by local pastoral communities, who employed their indigenous knowledge of breeding and the local ecosystem to create an animal genetic resource that is perfectly adapted to its unique environment. The Banni Grassland was once considered to be the finest and largest grassland in India, covering 2 400 km2. However, in the mid-1970s the Banni Grassland started to suffer degradation. First, salinity of the soil increased as rivers flowing through the grassland were dammed, restricting water flow and inhibiting the ability of the waters to flush salts away from the soil surface. Second, the alien plant Prosopis julifera was introduced in the area and became a highly invasive species.
In 2008, with support from Sahjeevan, an NGO working on environmental conservation and revival of traditional livelihoods, the pastoral communities of the Banni formed an organization called Banni Pashu Uchherak Maldhari Sangathan (Banni Breeders’ Association). The objectives of the association were to revive the livestock economy of the region, register the Banni Buffalo as a recognized breed, conserve the Banni Grassland and protect their customary grazing rights.
The breeders’ association, in collaboration with State Department of Animal Husbandry, Government of Gujarat, the NGO Sahjeevan and Sardarkrushinagar Dantiwada Agricultural University, carried out data recording, prepared a description of the breed, and sent a registration application to the National Bureau of Animal Genetic Resources, the body responsible for recognition of breeds in India. In 2010, the Banni Buffalo was successfully registered as the eleventh Buffalo breed of India. This was the first case in India in which a community that developed and conserved a breed was able to gain official recognition for what they have been doing for centuries.
In addition to preparing for the breed registration, in 2008, the breeders’ association started organizing an animal fair called “Banni Pashu Mela”, not only to promote the Banni breed, and its high profitability in the grassland ecosystem, but also to promote their unique pastoral way of life.
The association also successfully negotiated with two dairy processors to start an organized milk collection system in the Banni region, which now collects more than 100 000 litres of milk every day. More dairies are now showing willingness to start operations in the Banni region.
With more than 900 members from all 19 village governments in the Banni Grassland, the Banni Breeders’ Association is currently developing a participatory grassland conservation plan that draws on their traditional knowledge of grazing resources and understanding of sustainable resource utilization. The plan is designed to counter a standard fencing-based plan that was prepared by the governmental Forest Department without consulting the pastoral communities, despite the fact that these communities have been present in the Banni for more than 500 years and that they generate in excess of one billion rupees per year through the livestock production.
Provided by Devinder K. Sadana.
Action 2. Work with the community to establish a breeding programme The community may be willing to work with the government or other stakeholders to develop and adopt a breeding programme to improve the productivity and maintenance of genetic variability (see Section 7), but may require assistance in organizational and technical matters. Outside assistance can facilitate the adoption of breeding goals and the development, implementation and maintenance of breeding programmes. Box 24 summarizes the establishment of a community-based breeding programme for sheep in Ethiopia, where a government research institute provided technical assistance in developing a breeding programme.
CGRFA/WG-AnGR-7/12/Inf.6 Box A community-based breeding programme in Menz, Ethiopia Menz is an area of north-central Ethiopia where sheep farming is a primary economic activity. In an effort to improve the livelihoods of the local people, the national government targeted improvement of sheep productivity, including the adoption of community-based breeding programmes. The local sheep had several positive characteristics, including adaptation to the local feed sources, parasite resistance and tasty meat, but opportunities for genetic improvement were limited by a lack of organization among the farmers, poor knowledge of breeding programmes, the absence of data recording and negative selection through the slaughter of the fastest-growing males.
The project started in 20007 and was undertaken by researchers at the Debre Berhan Agricultural Research Center. A planning workshop determined the precise project sites. A questionnaire was used to gather information about ongoing breeding and husbandry practices and to characterize the breed and the production system. Follow-up studies addressed selection procedures and marketing and social aspects. In May 2009, the researchers proposed a new selection programme, based on selection of the top 10 percent of rams for live weight, lamb survival and fleece weight and the use of the selected rams for up to two years. The farmers agreed to the new system and the system, which was launched in June 2009 by recording data on the animals in the various flocks. Training was provided to the farmers and each was provided with a record book. Since then, data have been recorded on all newly born animals. Selection of rams is done twice a year – in February and June.
Rams are then distributed to breeding groups consisting of ewes managed together in communal grazing (eight rams per group). A farmer is nominated as the leader of each group and is responsible for rotating rams among the ewes and reporting on progress and any problems.
In addition to the breeding programme, several complementary initiatives were launched. To help generate interest among participants, at the time of selection, prizes are awarded to the owners of the best lambs in the programme and to the best farmers. In addition, technical assistance is provided on animal nutrition, diet formulation and disease control. Individual farmers are also selected and trained to provide basic community veterinary services. A cooperative is being developed for marketing and ongoing management of breeding activities.
The project is still in its initial stages and challenges remain, but many positive results are visible.
Because of their role in designing the programme, farmers are keenly interested in the project and regard it as a way to improve their livelihoods.
Source: Getachew and Gizaw (2010).
Working together as a breeders’ association, the community may, for example, identify the target traits in a selection objective within the breed. This is generally considered the most important issue in the genetic improvement of animals in a conservation programme. The objective should be clear enough to exploit the genetic potential existing in the population to the maximum extent possible. Genetic improvement will gradually increase the average genetic merit of the breed with respect to the chosen objective. The community may then participate in creating and promoting a brand for products derived from the breed. Such measures will strengthen the economic value of the animals and may ultimately lead to the breed becoming self-sustainable. In such cases, external support from the government can gradually be withdrawn and the breeders’ association can take over the running of the programme.
Details on establishing selection objectives are provided in the guidelines Breeding strategies for sustainable management of animal genetic resources in this series (FAO, 2010).
A breeders’ association can take responsibility for aspects of operating a breeding programme, such as animal identification, performance recording and genetic evaluation, with or without financial and technical assistance from the government. Capacity to undertake such activities is one of the major advantages of having a breeders’ association, because individual breeders will typically lack the time and technical capacity to perform them and because of the efficiency of having these activities performed by a central agency. Collaboration among different breeders’ associations may further CGRFA/WG-AnGR-7/12/Inf.6 increase the efficiency of these activities, as may the establishment of associations for transboundary breeds that are present in more than one country.
Action 3. Establish a nucleus herd for management of genetic improvement A nucleus herd of up to several hundred superior females and sufficient numbers of superior fertile males (about 1 for every 10 to 20 females) in the breed’s local region may be a key tool in managing the population. Within a nucleus herd, selection and mating decisions can be controlled more strictly, allowing the implementation of more complex and effective management approaches (see Sections and 7). An “open” nucleus design, in which relationships are established between private livestock keepers and institutional herds, with gene flow both directions and ongoing identification of superior animals in both populations, may be particularly attractive for local breeders. They gain a larger ownership stake in the programme and can benefit both by obtaining superior germplasm from the nucleus and by having the opportunity to sell their best animals to the nucleus at a premium price.
Furthermore, when managed correctly, open nucleus systems tend to reduce inbreeding relative to closed schemes. However, establishing and operating a nucleus herd requires significant technical and financial resources. Determining cost and benefits will be a critical first step in such an endeavour and support from the government or an NGO or from commercial organizations may be necessary.
Action 4. Provide incentives, including capacity building, and complementary institutions Providing specific incentives or other assistance may be a fundamental step in directing a breed at risk back on a sustainable course. For example, livestock keepers may be abandoning a given breed, or even animal production in general, because they cannot market the products at a price or quantity high enough to ensure a satisfactory livelihood. Assistance in improving productivity or in establishing a consistent market for the breed’s products may increase livestock keepers’ income, and thereby provide support to the livelihoods of the community and, help ensure the survival of the threatened breed. Livestock keepers have an exceptional amount of indigenous knowledge and may thus already be sufficiently skilled in managing their breed. However, the breed may be threatened by factors outside the control of the livestock keepers or by insufficiency in a particular capacity that can be overcome through training. Alternatively, livestock keepers may actually be providing a service that can be regarded as a common good that is not being properly valued by the market (e.g. maintenance of genetic diversity or ecosystem conservation). In such cases, providing payment for the services provided may be considered, as long as such payments are not market-distorting and respect international trade agreements. These options are discussed in more detail in Section 8.
The loss of a particular breed may also be related to wider-scale rural development problems that are causing breeders (and their children) to give up livestock production entirely and seek other livelihood opportunities. If this is the case, incentives such as those discussed above may not be sufficient. It is important to consider such wider-scale threats and how they might be addressed. Perhaps it may be necessary to establish non-agricultural services, such as improved educational opportunities for children, improved health care and local off-farm employment opportunities to help sustain the community in general.
Establishing a breeders’ association Rationale Breeders’ associations (also known as “breed societies” or “breed associations”) can be essential to the long-term success of conserving animal genetic resources, by playing many roles, including as serving as an effective monitor for threats to the breed. Most standardized and international transboundary breeds are well-served by breeders’ associations that have clearly-documented procedures and functions. In most cases the functions performed by the breed associations of standardized breeds have been assumed as a universal model for other classes of breeds. Many local breeds have less formal organization but have a very high priority for conservation. Accommodating the needs of such breeds will often mean that some details of the model for standardized breeds will often have to be adapted.
Organizing breeders presents numerous challenges. If herds have been isolated for long periods of time, each breeder is very likely to consider his or her own herd to be the only typical one, which may CGRFA/WG-AnGR-7/12/Inf.6 lead to fragmentation and difficulty in establishing common goals (see Box 25). The leading traditional breeders are often elderly, and may not have heirs interested in continuing to keep the family livestock. This threatens breed viability, as it can easily lead to the loss of the culture surrounding the breed. Getting beyond individual pride and self-interest is challenging, but is crucial for long-term success. Traditional breeders have assumptions and beliefs that vary, and that strongly shaped breed development and maintenance. Therefore, they may be reluctant to adopt substantial changes, even if they may seem to an outsider as necessary for the breed’s future survival.
Box Colonial Spanish Horse breeders’ associations in the United States of America Breeders of Colonial Spanish horses in the United States of America conserve a very fragmented and dispersed breed based on non-standardized populations and feral animals. The result of strong local attachment has left many breeders with a skewed appreciation of the overall breed, and an intense focus on the local resource. The result is 20 different breeders’ associations for this breed, which numbers at most 3 000 animals. Fragmentation makes long-term management and survival precarious. As a reaction to this, a few associations with an inclusive philosophy now also promote the breed. The fragmentation among the breeders is the result of strongly held opinions on breed purity, but at some point this leads only to a smaller and smaller gene pool that cannot avoid high levels of inbreeding in the long term.
Provided by Phil Sponenberg.
Breeders’ associations are generally democratic institutions. Members have to meet requirements for active membership and participation and to agree to a set of rules. They are able to participate in shared decision-making, are eligible to register livestock and can benefit from recording schemes and promotional efforts.
Objective: To create a well-functioning breeders’ association.
1. Genetic and demographic data on the breed;
2. List of breeders who keep animals of the breed;
3. Some knowledge of the history of the breed and its present functions;
and 4. Knowledge of the goals of the breeders and their history of cooperation.
• A well-functioning breeding organization with a description of:
requirements for membership registry protocols rules and by-laws dues and fees for members communication methods for education and training mechanisms for conflict prevention and resolution • Communication between the breeders’ association and national entities responsible for management of animal genetic resources.
Task 1. Assess the willingness of members of the community to establish a breeders’ association Action 1. Discuss the possibility of a breeders’ association during the initial participatory studies The success of a breeders’ association will largely depend upon the number of breeders participating and their willingness and enthusiasm. The level of interest should be evaluated as soon as possible, so as to avoid futile investment of time establishing an organization that is not sustainable. The community members should be made aware of the benefits of breeders’ associations and of the inputs required.
Action 2. Identify specific members of the community that may have a particular interest in joining a breeders’ association and serving in leadership roles CGRFA/WG-AnGR-7/12/Inf.6 A breeders’ association is a NGO and will only be successful if its leaders are well-respected in the community and committed to the association and to maintaining the breed. To ensure that breeders’ associations have sufficient grassroots support to ensure sustainability, some countries only provide governmental assistance after the associations are established and viable (see Box 26).
Box A two-step approach to supporting breeders’ associations in Latin America Some Latin American countries, such as Brazil and Colombia, have an interesting two-step approach to creating breeders’ associations. At the beginning, a group of breeders of a specific breed get together to create an organization that is considered a promotional association. The breeders then develop a set of breed descriptors and the by-laws, and invite new members to join the association to try to increase its membership. Once these phases are completed, the Ministry of Agriculture is contacted to examine the documents of the promotional association and to determine whether the population of animals referred to by the association warrants being considered as a distinct breed and whether the number of breeders is sufficient (e.g. in Colombia, there is a minimum requirement of ten breeders). If the Ministry of Agriculture considers that the promotional association adheres to all these requirements, it can be recognized, and becomes an official breeders’ association, with a mandate to operate the genealogical registration programme for the animals of that specific breed. Once the association is officially recognized, it is obliged to send an annual report of activities to the Ministry of Agriculture with copies of all registrations made during the year. The Ministry of Agriculture is then responsible for auditing the official associations. This procedure has proven to be successful and serves as an example that can be followed by countries that do not have a procedure for creating breeders’ associations.
Provided by German Martinez Correal and Arthur Mariante.
In some cases, the interest and dedication of a single person may be nearly sufficient to drive a breed’s conservation and development (see Box 27). However, even if a single individual is willing to sustain a breed, he or she may gladly accept outside assistance. This outside assistance may also decrease risks associated with having a single major breeder and stakeholder.
Box Conservation of Tharparkar cattle in India Conservation of a breed can be accomplished by the sheer vision, will and dedication of a highly motivated individual, as exemplified by the work undertaken by Mr Magraj Jain of Rajasthan in India.
It was during 1990s that he recognized that the productivity of the livestock in the region was deteriorating, and that the primary reason for this was the diminishing numbers of the local Tharparkar breed of cattle, which by the late 1990s was at risk of extinction. Through his NGO, named SURE (Society to Uplift Rural Economy, Barmer, Rajasthan) Mr Jain undertook the task of raising young pure-bred Tharparkar bulls and distributed them one-by-one in 34 villages. Villages were selected based on discussions with stakeholders and the interest expressed by the local people. Each bull was given to a family in the village who agreed to maintain it in exchange for earnings obtained through the use of the bull by other farmers for natural-service mating of their cows. The services were recorded. When the progeny became mature, their milk yield was also recorded. By 2007, more than 2 100 pure Tharparkar cows had been born to these bulls and the project continues to this day.
Provided by Devinder K. Sadana.
Task 2. Develop and implement a process, involving the relevant stakeholders, through which a breeding organization is described and established Action 1. Determine requirements for membership of the association Most breeders’ associations have different classes of membership. Full membership in many associations is limited to people that actively own and breed animals. Breeder-members have voting CGRFA/WG-AnGR-7/12/Inf.6 rights, which ensure that control of the breed’s future is determined by those most affected by decisions and who contribute to the conservation of the breed.
An important first step is to determine which animals and breeders are to be considered as representative of the traditional type of the breed. This process determines the foundation and forever shapes the descendant breed. The decisions regarding which breeders and animals to include within the breed usually occur simultaneously, because each affects the other. Outside entities, such as governmental organizations or NGOs, can help guide associations through this step. It is best to include animals that are pure-bred to the local population and to avoid any animals with known influence from outside breeds. However, if the number of verified pure-bred animals is too small to obtain a viable breeding population, standards may have to be relaxed somewhat to allow animals that are not pure-bred but have a high proportion of the desired breed to be included in the foundation stock (see Box 28). In this way a larger amount of genetic variation will be available immediately, which is likely to be useful for the future development of the breed.
Box Incorporating non-pure-bred animals into a breed founder population The concept of resemblance through common hereditary descent is a useful addition to any definition of a breed (see Box 1). Resemblance through common hereditary implies that, ideally, a breed has no exchange of genes with other breeds, i.e. that no introgression of genes from other populations takes place. A rule of thumb in practical breeding is that no more than 12.5 percent of outside (exogenous) genes (i.e. one of the eight great-parents is from another breed) should be accepted within a breed.
Following this rule of thumb means that if an animal’s proportion of exogenous-breed genes is greater than 12.5 percent, it is considered a member of a different “breed” and not allowed to be a member of a foundation population. Standardized herdbooks usually do not consider individuals with more than 12.5 percent exogenous genes to be pure-bred and most have much stricter rules for this percentage.
Provided by Phil Sponenberg.
In addition to a full regular membership category for active breeders, associations may consider also having other types of membership that may help to expand interest in the breed (and perhaps revenue for the association) but have limited influence over breeding policy. Such additional classes of membership may include associate membership for non-breeders or “junior” membership for non adults. Members belonging to such categories typically do not vote, but are entitled to all other benefits of membership. In breeds that don’t produce widely marketable products, such as many horse breeds, it may be necessary to include non-breeders as full members. In such breeds, non-breeders contribute to the breed through promotion and use, and their voices must be heard in decision-making.
Including new breeders in the association is essential, including breeders that are not members of the cultural group that originally kept the breed. Efforts to include new breeders can sometimes threaten traditional breeders, because some control over the future of the breed is usually surrendered.
Expanding the group of breeders involves cultural change that affects how the breed is selected and valued. Managing these tensions is challenging.
Where possible, an association should have special designations for long-term traditional breeders that help to ensure their continued participation. Special allowances for such breeders may include discounted costs in registration procedures or waiving or reducing membership dues.
Action 2. Establish registry protocols Most associations register animals and validate pedigrees. Procedures must be consistent and uniformly applied. A variety of software programs, each with strengths and weaknesses, are available.
The pursuit of complete accuracy comes at an economic cost. For example, DNA validation of pedigrees can provide high accuracy, but is unrealistic both for animals that are of low individual economic value and for animals raised in extensive situations.
The registry function of a breeders’ association for a non-standardized breed can often be similar to that for standardized breeds. For breeds raised in extensive production systems, especially in multi-sire CGRFA/WG-AnGR-7/12/Inf.6 herds and for species in which the animals have relatively low individual value (e.g. poultry, goats and sheep) other procedures are needed. One strategy is to register and monitor entire herds or flocks rather than individual animals. It is very important to ensure pure breeding. Procedures and validation must be tailored to fit each individual case, as they must reflect the realities of the local culture and local husbandry practices.
New and expanding breeders’ associations must develop methods for including candidate animals in the registered population. Some approaches can be applied on an animal-by-animal basis (see Box 29), whereas others apply to whole herds. Box 30 presents an example of an animal-by-animal procedure and Box 31 gives an example of herd-based registry. In economically developed countries, it is common for new breeders’ associations to have a short period in which foundation animals are registered, after which the registry is restricted to animals with registered parents and grandparents. This is a typical strategy for standardized breeds, but works poorly for non-standardized breeds because isolated pockets of pure-bred animals are likely to continue to be discovered for a long time. Procedures for inclusion of newly encountered animals must be developed, and must be applied uniformly and fairly.
Box Incorporating non-registered animals in a herdbook Inclusion of candidate animals should follow documentation of their origin and type. The history of the population (geography, foundation stock, length of genetic isolation, and the source and frequency of any additions of animals from other breeds) should be evaluated, as should the phenotypes of individual animals. Where possible, DNA analysis should be used to detect introgression from other breeds, but results must be interpreted carefully, as bona fide pure-bred animals from a long-isolated pocket of the breed will often have novel DNA variants with respect to the rest of the breed. These animals could be particularly valuable to the breed for their genetic distinctiveness, adding to genetic diversity. If desired, a progeny test can be done to validate animals’ ability to reproduce the traditional type. This latter step is less important if DNA validation is possible, but is useful in cases where candidates have novel phenotypic variants (e.g. coat colour, horns). Inclusion of newly discovered animals may also be important for some standardized breeds with long-established herdbooks. For example, registering new animals can be essential to long term conservation success of breeds with very small population sizes. Newly discovered animals must, however, enter the breed on equal footing with existing animals or their genetic contribution will be diluted out and any potential benefit from their inclusion in the herdbook will be forsaken.
Provided by Phil Sponenberg.
Performance and type classification recording schemes are important in many breeders’ associations, especially where breeders want to implement selection programmes. These data are useful for breed improvement schemes, and breeders’ associations are the logical place to maintain them.
CGRFA/WG-AnGR-7/12/Inf.6 Box Protocol for addition of non-registered animals in the herdbook of Huacaya alpacas in Peru Alpacas and llamas are among Peru’s most important animal genetic resources. To help improve their management, the Official Genealogical Registry for Alpacas and Llamas (OGRAL) was created in 1997 and a herdbook was established. One objective of OGRAL is to recognize and record the alpacas that conform to a desired phenotype, based on characteristics of their fleece and conformation, so that these animals can be added to the registry.
A committee composed of representatives of breeders’ associations, research institutions, NGOs and national universities established rules for registering alpacas of the Huacaya breed in a national herdbook. A visual scoring system was established with the following 100-point scale:
Fleece Fineness Length Density 10 Crimp Uniformity Conformation Head Height Fibre coverage of rear legs General appearance Assessments are undertaken by official technicians that have been trained by the Ministry of Agriculture. Alpacas that obtain at least 75 points are allowed to be registered in the OGRAL herdbook Only about 1 percent of the population achieves registration and such animals have high value as breeding stock.
Provided by Gustavo Gutierrez.
Box The protocols of the Coastal South Native Sheep Alliance in the United States of America The Coastal South Native Sheep Alliance in the United States of America has developed new protocols for dealing with landrace conservation. Organization and rules are minimal, but a commitment to pure-bred breeding is the main objective underpinning the group. The basic rules include:
• Sheep of other breeds are not to be kept on the same property as a Gulf Coast flock recognized by the Alliance. This prevents introgression from other breeds, especially where animals are not individually identified and multi-sire mating systems are used.
• Each flock submits a brief flock history, including original sources and foundation year.
• Additions to the flock are documented by source, date and sex. Ideally, additions only come from other flocks recognized as pure-bred by the Alliance.
• Each flock participant reports census figures annually, including the sources of any additions.
• If breeders maintain different family groups, each is tracked as a separate flock.
These rules protect the genetic integrity of the breed, while allowing it to persist as a traditional local resource. However, they require commitment on the part of the breeders, and even the low level of breed-specific activity involved (documentation of flocks, documentation of additions, annual census) can be enough of a change from tradition to result in non-compliance.
Provided by Phil Sponenberg.
CGRFA/WG-AnGR-7/12/Inf.6 Action 3. Establish by-laws Election procedures Well-defined procedures for decision-making can help prevent confusion and controversy. By-laws determine members’ eligibility, the mode of voting and the role of members in various sorts of decision. One extreme is a completely democratic form of organization in which members vote on every decision. This is generally limited to small associations. Larger associations usually have officers and/or a board of directors that makes decisions, with periodic elections to renew the board and ensure that it reflects the will of the membership.
It is important to establish election procedures, including the frequency of elections. Such rules encourage participation and develop loyalty and a sense of ownership of the association among the members. Participation of original breeders must be ensured and may require that their status is recognized in some way.
Board of directors In most associations, the members elect a board of directors, which enacts specific procedures and policies. The number of directors can vary, but to ensure continuity it is best to have staggered terms. For example, if a board has six members with three-year terms, then elections for two of the positions on the board could be held each year. This would allow the two new members to serve with two members that have served for four years and two that have served for two years. This ensures continuity, but also refreshes the leadership with new ideas and enthusiasm. To ensure this end, many associations limit the number of consecutive terms that any individual board member can serve. Most boards include a convening president, a vice president, a secretary, a treasurer and, in smaller associations, a registrar.
Action 4. Determine membership dues and fees Dues for membership and fees for services provided by the association, such as registration, are an important source of revenue that helps to meet the costs incurred by the association. In most situations, dues and fees are set by the membership or board of directors. Dues and fees must be set fairly and uniformly, and should have few changes over time. Dues and fees reinforce the message that registration and participation are valuable. Depending on the goals of the association and the variability in herd sizes, some breeders’ associations may charge only membership dues with no additional fees for registration of animals. This encourages the registration of all animals and promotes the participation of breeders with large herds.
Box Promotion of the Leicester Longwool sheep in the United States of America The Leicester Longwool Breeders’ Association in the United States of America has come to play an important role as the guardian of the largest national flock of this breed. The breeders’ association discourages the usual competitive showing in favour of “card-grading” in which each sheep is individually evaluated by three judges for compliance with the breed standard. Following evaluation, each sheep is awarded a “card” determined by their relative quality: blue cards for superior breeding stock, red cards for good breeding stock, yellow cards for acceptable breeding stock, and white cards for those unacceptable as breeding stock. This process is educational because one of the judges speaks to the observers following the evaluation of each sheep, explaining the process and the results. This ensures effective education on breed type for both breeders and the general public.
Source: Sponenberg et al. (2009).
Action 5. Establish communication methods for educational and training programmes.
Communication within a breed association serves several different purposes, each of which will require specific mechanisms. Communication between breeders establishes a community among them.
The goal should be to foster a feeling of belonging and participation so that breeders feel involved in, and essential to, the future of the breed. Newsletters, meetings, field days, shows and fairs, web sites and electronic chat groups can all foster this.
CGRFA/WG-AnGR-7/12/Inf.6 Associations also need to educate breeders on effective breed maintenance. The goal should be to have an informed and committed membership that is knowledgeable about population dynamics and their importance to conservation. Breeders must also understand breed type as it relates to traditional use and value. Educational methods include field days or workshops, traditional newsletters and electronic forms of communication.
Breeders’ associations have an important role in promoting the breed and its products to an audience beyond the membership. Marketing and other promotional activities are essential for the long-term sustainability of an association (see Box 32).
Box Accounting for cultural differences among members of breeders’ associations Breeders’ associations must always reflect underlying cultural norms. This is especially challenging if a single breed is held by more than one cultural community. In the United States of America, the Navajo Churro sheep has centuries-long connections with both Din Navajo (indigenous) and Hispanic communities in New Mexico and Arizona, and more recently with Anglo breeders and enthusiasts.
Throughout its history, the Navajo-Churro population has oscillated widely, due in large part to various government interventions, including population displacement, subsidized cross-breeding and restrictions on grazing on public lands. Most recently, a large-scale culling programme in the mid-to-late 1900s pushed breed numbers to fewer than 500 head. In response, breeders and breeding programmes organized themselves to ensure that the breed did not drift to extinction. Breeders were fortunate in having key individuals in all three cultures that reached out to the other communities. These leaders built the Navajo-Churro Sheep Association with an organizational norm of cross-cultural appreciation and inclusion. Cultural diversity in the group is not ignored, but embraced and celebrated as an important aspect of breed conservation. This ensures that cultural diversity serves effective conservation rather than defeating it, by ensuring a strong commitment to the breed and its future.
Developing an inclusive association can be daunting, and although the Navajo-Churro Sheep Association has not yet fully integrated all breeders, especially among the indigenous communities, it applies an approach that considers the needs of the various ethnic groups. The Navajo-Churro registry depends on the inspection of each candidate sheep presented for inclusion in the flock book, even if both parents are registered. This model works best among the Anglo-American communities that are familiar with such procedures. These procedures are foreign to many of the Din and Hispanic breeders, whose families have been raising the breed for centuries. To improve integration, the organization has Navajo-speaking and Hispanic inspectors dispersed throughout the traditional range of the breed so that traditional breeders have relatively easy access to inspection of their sheep. Having inspections done by members of the same cultural group facilitates communication, encouragement, and participation.
Breed-specific promotions for Navajo-Churro wool and meat have linked participation in the breeders’ association with increased economic returns for these products. The Black Mesa Weavers for Life and Land, a non-profit enterprise assisting the Din community in particular, has provided services for marketing wool and woollen products on a “Fair Trade” basis. Linking participation, breed recognition and enhanced commercial opportunities has ensured a viable association. Participation in the association has become an important way for each of the various communities to emphasize its individual culture and its contribution to the breed, which has increased a sense of ownership of the breed. Although obstacles for both the breed and its communities of livestock keepers remain, cooperation among the communities has improved the outlook for the future of the breed.
Source: Sponenberg and Taylor (2009).
Action 6. Develop and adopt a procedure for conflict resolution Because breeders’ associations comprise individual breeders, conflicts among members are certain to arise. Most conflicts are between individual members or breeders, but some arise from fundamental differences in basic philosophy of animal breeding (e.g. the breeding goal), production and sustainable use. It is important to have mechanisms in place to spot conflict early and to resolve it fairly and quickly. The goal of conflict resolution must always take into account the breed and its need for CGRFA/WG-AnGR-7/12/Inf.6 engaged and involved breeders that communicate and share. Conflict resolution is especially important for new associations of local breeds because they may have more traditional and isolated breeders.
These conflicts must be judiciously solved if the breed is to expand rather than contract to the point of extinction. In some cases, conflicts can be foreseen and prevented or at least minimized. See Box for a specific example where a breeders’ association developed certain policies and practices accounting for cultural differences among members.
Ideally, it will be possible to reach a consensus and all members of the association will agree on a common strategy for breed conservation and development. However, in some cases the conflict may not be resolvable and the breeding association may split into factions, each of which having its own goals (see Box 34). In such cases population management is particularly important, because splitting the population will result in subgroups with smaller N e and fewer resources for association activities.
Box Introgression poorly accepted by some breeders of the Texas Longhorn in the United States of America In some instances, conflicts among breeders may not be easily resolved and drastic action must be taken. Such a circumstance confronted breeders of the Texas Longhorn cattle breed in the United States of America. Different groups of breeders had opposing opinions regarding the philosophical choice between selecting for improved production and strictly maintaining a traditional phenotype.
In particular, some of the breeders favoured using cross-breeding to improve production.
Traditionally minded breeders were apprehensive about possible changes in type that may result from cross-breeding. These breeders felt that it was safest to select for traditional type within the breed so as to avoid losing unique aspects of the breed, particularly its adaptation to hot, dry environments. This difference in opinion has split the breed into two groups, with the more traditionally minded breeders’ committed to pure breeding and resistant to changes in type and conformation. This effectively conserves the breed, but fixes conformation and production near current levels with little room for improvement. The long-term consequences of this decision are yet to be seen, although the chances of success are boosted by the fact that the traditional breed continues to excel in environmental adaptation and is sought as a maternal base in terminal cross breeding programmes.
Provided by Phil Sponenberg.
Auditing the breeders’ association and its activities Rationale Many breeders’ associations need support from either governments or from NGOs in order for them to function well in conserving animal genetic resources. This support may be justified by the fact that activities of the association contribute to public benefits such as maintenance of biodiversity, rural development and increased food security. Support may be financial or logistical. The provision of support justifies periodic assessment of the associations’ activities. Such evaluations help to ensure that the resources provided are being used effectively in conservation work.
Whether or not out outside support is being provided, a breeders’ association should still undertake periodic auto-evaluations, to ensure that its members are being properly served and its goals with respect to animal genetic resources management are being effectively achieved.
Breeders’ associations should actively monitor and appraise the role of the breed as a valued natural resource for the country. This task should include assessments of demand for the breed and its products and threats to its long term viability (FAO, 2010). Assessing and promotion of a breed’s value usually requires a strong breeders’ association, as weak or inactive associations may be overlooked by national efforts (FAO, 2009).
In some instances, particularly with rare breeds, the number of breeders is not large enough to perform all of the functions generally required from a breeders’ association. Even when associations are large CGRFA/WG-AnGR-7/12/Inf.6 enough to perform the tasks needed for their breed, maintaining separate sets of infrastructure for each breed may not be financially efficient. In such instances, a wise alternative may be to create an umbrella organization that performs various administrative functions for a number of otherwise independent breeders’ associations. An example of such a system is presented in Box 35.
Box Combined Flock Book – a multibreed society in the United Kingdom Small breeders’ associations frequently have limited resources. This shortage of resources may cause inefficient administration, and is likely to prevent the provision of comprehensive breeding and support services to breeders and members. The creation of a central facility by a larger organization covering several breeds can overcome these problems.
In the United Kingdom, the Combined Flock Book was established by Countrywide Livestock Ltd in 1974 as the breed society for breeds that were unable to maintain their own administrative offices, and it currently provides services for eight breeds of sheep. The primary objectives of the Combined Flock Book are to protect the genetic integrity (pure-breeding) of each breed and to promote the value the breeds.
The Combined Flock Book originally operated within the structure Countrywide Livestock Ltd and was administered by a committee comprising one representative from each breed under the chairmanship of the President of Countrywide Livestock Ltd. Each breed also was supported by its own group of breeders who were concerned primarily with promoting their breed. The Combined Flock Book was later transferred to the NGO Rare Breeds Survival Trust under a similar arrangement.
The services provided by Countrywide Livestock Ltd included:
• registration of all pure-bred animals with details of individual identity, sire, dam, date of birth and sex, plus information on colour and horns where appropriate;
• calculation of inbreeding, kinship, and individual founder contributions for each breed;
• realization of DNA profiles for parentage verification, product provenance and assignment to breed;
• provision of advice on breeding policy and individual mating plans;
• promotion of member breeds through a web site, literature, exhibitions, and livestock shows;
and • conflict resolution.
Provided by Lawrence Alderson.
Objective: Develop an auditing process for a breeders’ association and its breeding and conservation plan.
1. Laws and by-laws of the breeders’ association;
and 2. A description of the association’s activities including the breeding and conservation plan.
• Description of an auditing procedure for a breeders’ association and its activities.
Task 1. Evaluate participation and decision-making procedures Action 1. Describe and evaluate the mechanisms for participation Strong breeders’ associations encourage broad participation from their members and producers (FAO, 2010). All members should feel welcome to participate and to contribute. Major decisions should be taken in a democratic manner, avoiding domination by one or a few persons. Some associations get dragged into various controversies arising from the dissension of a single or small number of people and while some individuals may hold very strong opinions, in most cases the breed itself loses support and members. Ensuring that an association functions in a way that ensures a sense of community and benefits all members to a fair degree is an important objective.
CGRFA/WG-AnGR-7/12/Inf.6 Action 2. Assess the decision-making processes Breeders’ associations need shared and open decision-making procedures to foster broad ownership of the association and high levels of participation. The actual keepers and breeders must be consulted and considered if the association’s programmes are to be successful (FAO, 2010). Breeders and livestock keepers have intimate knowledge of the breed’s production system. Their participation must be ensured by respecting their opinions and attitudes.
Action 3. Evaluate the provision of benefits to association members Breeders’ associations should provide expert or technical support to their breeder members (FAO, 2010), including husbandry, health care, animal selection and animal breeding techniques, when appropriate, and strategies for the long-term viability of the breed.
Action 4. Evaluate the procedure for inclusion of new members The evaluation of mechanisms for including new breeders in the organization should include assessments of membership protocols and of the extent of participation by old and new breeders in the association and the registry. Breeders’ associations should be inclusive and welcome new breeders. This is especially important when new associations are being formed for non-standardized breeds. It is essential that the community of breeders for each breed is growing rather than shrinking. Strengthening cooperation among breeders is an important role for breeders’ associations (FAO, 2010).
Task 2. Appraise the genetic purity of the population As noted above, public support to a breeders’ association may be justified if the association is actively maintaining biodiversity by promoting maintenance of a valuable genetic resource. If this objective is not being reached, then continued support may not be warranted.
Action 1. Evaluate the level of pure-bred breeding as opposed to deliberate or casual introgression This evaluation needs to include detection of overt and fraudulent introgression by breeders, as well as avoiding the inclusion of cross-breeds through inattention. Breeders’ associations must insist on pure breeding and emphasize this to members. Commitment to pure-breeding should be openly stated as a core value of the association. Competitive activities such as animal shows or production award contests are a way to promote interest in the breed and reward active participation, but they can also provide an incentive for fraudulent cross-breeding. In some cases, awards in the show ring or production contest can go to animals with obvious introgression from other breeds if not prevented by the breeders’ association. This can send a very damaging signal to the breeder membership. In general, the association should insist that breeders recognize and appreciate the breed for what it is rather than trying to change it through crossing.
Action 2. Check accuracy of parental information Correctness in animal identification is important in order to ensure freedom from undesired cross breeding and precise evaluation of breeding values. As mentioned above, routine DNA tests will not be practical in many situations, but breeders’ associations may care to adopt a programme that randomly checks the parentage of a proportion of animals by using DNA from the individual and its putative parents. The government may wish to audit such a programme, especially if support is being provided for identification and performance recording.
Task 3. Evaluate the status of the animal genetic resources under management The breeders’ association needs to maintain its identity and its sense of ownership of the breed. These aspects will be largely influenced by the autonomy of the association with respect to the breeding programme, with more autonomy leading to a greater sense of ownership. However, if outside financial support is being provided, then some reasonable conditions may be applied to this autonomy.
Occasional auditing by the funding agency of the population and breeding programme may help to evaluate the breed’s prospects for sustainability and identify threats to its viability.
Action 1. Analyse the population structure and its use in the breeding programme Procedures for designating strains or families within the overall breed, mechanisms for census of the breed, and evaluations of population structure should be appraised. Effective breeders’ associations periodically monitor the population structure of the breed. Breeders should be educated against CGRFA/WG-AnGR-7/12/Inf.6 temporary fads in breeding so as to ensure a broad representation in the next generation. The overuse a few well-known animals will result in a bottleneck. This is damaging in common breeds and disastrous in breeds that are at risk of extinction. Breeders’ associations should educate breeders on healthy population structure. Recognition of the value of different strains or sire families within the breed can be very helpful, especially in non-standardized breeds where variation in different families can be relatively large. Breeders’ associations should monitor the breed’s population size, including the details of animals with large average genetic relationship with the rest of the breed.
Action 2. Evaluate the breeding and conservation programme The plans for the management of the genetic variability of the population should be appraised, including periodic exchanges of breeding animals among breeding herds, and any cryoconservation activities, such as targeted freezing of gametes from under-represented animals. If the breeding and conservation plan involves genetic improvement, then the genetic and phenotypic trends of the population should be checked to gauge the effectiveness of these activities. Breeders’ associations should be alert to herds and individual animals that are of high genetic diversity and importance to the future of the breed (FAO, 2005). Breeders’ associations can actively promote rules and protocols that serve to prevent genetic erosion by encouraging genetic exchanges among member herds (FAO, 2010). However, such efforts need to be broadly based so as to ensure that no single herd swamps the breed by providing a disproportionate share of breeding animals.
Centralized ex situ conservation on institutional farms Rationale In situ conservation is usually the preferred option for in vivo conservation of animal genetic resources, due to the advantages discussed in Section 3. However, in some situations, ex situ conservation of live animals is a more practical option. For example, a breed may have reached a population size that is too small to be raised by a group of farmers. Alternatively, a breed may be valued primarily for its option and existence values (see Box 12), and therefore not profitable enough to be maintained by livestock keepers, but may still need to be immediately accessible in its live animal form. Perhaps tight central control of breeding is required and such control can only be achieved via ex situ conservation on a single farm. Such programmes are typically operated by the government or NGOs rather than by the commercial sector.
In many countries, institutional farms are owned by government and NGOs that are dedicated to research, teaching and development. Most of these farms maintain economically important breeds of various animal species, and are used as demonstration centres as well as for the production and dissemination of superior germplasm. For example, India has a well-developed system of institutional farms. Establishing organized breeding farms is important, especially for genetically eroded breeds that have very small populations and are sparsely scattered in their home regions.
Ex situ conservation of livestock breeds by establishing breed-specific farms involves a substantial investment in infrastructure and other resources. For these reasons, ex situ conservation programmes are usually limited to a few very unique breeds and maintain relatively small populations. As explained in Section 2, the genetic constitution of a small population can change rapidly through genetic drift, possibly resulting in the loss of genetic peculiarities and reduction in genetic variability, so the most important challenge in managing the population in ex situ conservation is to sustain genetic variability.
Objective: Establish and maintain populations of important animal genetic resources in centralized breeding herds.
1. Lists and of local breeds that are candidates for ex situ conservation and descriptions of their characteristics;
and 2. Knowledge of the location of individual animals from these breeds including those kept by CGRFA/WG-AnGR-7/12/Inf.6 private breeders and on existing institutional farms.
• Institutional herds for at-risk breeds, with active programmes for maintaining their genetic variability.
Task 1. Undertake the necessary preliminary planning, including feasibility studies, and secure access to facilities and funding Action 1. Assess the available institutional breeding farms An ex situ in vivo conservation programme will be more financially feasible if existing facilities and personnel can be used. These facilities may include governmental or non-governmental farms.
Action 2. Determine the breeds to be targeted by the conservation programme Following the procedures set out in Section 3, establish the targets regarding which breeds to include in breeding farms, paying close attention to breeds that have not already been targeted by existing programmes.
Action 3. Perform a feasibility study As noted above, ex situ conservation of animal genetic resources in vivo is an expensive undertaking and requires substantial planning. The population is unlikely to be financially self-sufficient, so the agencies (public or private) supporting the conservation activities have to be convinced of their value.
A feasibility study must be undertaken to determine the costs of establishing and maintaining the conservation programme. The assessment of costs must consider the initial acquisition of the animals, their maintenance, the acquisition or refurbishing of facilities, the maintenance of the facilities and personnel costs. Any revenues expected to be generated by the herd should also be accounted for.
Action 4. Identify possible donors In addition to the government, NGOs with an interest in conserving the diversity of agricultural genetic resources should be considered.
Action 5. Prepare and present proposals for the conservation plan to government officials and/or donor agencies A strong argument will be needed to convince donors to provide support for the conservation activities. The value of the animal genetic resources to be conserved must be stressed, including the opportunity costs of its loss. A well-done feasibility study (Action 3) will aid in preparing the proposal.
Task 2. Establish and operate the conservation programme Assuming Task 1 has been completed successfully, the real work of establishing and operating the ex situ conservation programme can get underway.
Action 1. Establish the herds at the institutional farm A general consensus suggests that an N e of 50 is a reasonable goal for conserved populations of animal genetic resources (FAO, 1984;
FAO, 1998). This need not be an immediate goal to be achieved at the start, but rather a goal to be aimed for over time through breeding management and continual acquisitions. In addition to stock that may already be available in institutional herds, animals can be purchased from livestock keepers in the breed’s breeding tract. Although reducing the size of the base population for a conservation programme will save money, it may introduce a founder effect (a type of genetic drift resulting in loss of genetic variability when a new population is established by a very small number of individuals selected from a larger population). The allelic frequencies of the small sample of animals may differ from that of the larger population and some alleles may be lost completely.
Animals purchased from outside should be free from disease or any noticeable defects, conform closely to the desired breed characteristics and be as unrelated as possible. If possible, they should have above-average performance for traits of economic importance (production and adaptive traits) so as to promote the economic self-sustainability of the farm. The best available breeding males (at least one for every ten females) should be procured.
Action 2. Develop breeding and husbandry strategies for the institutional herd CGRFA/WG-AnGR-7/12/Inf.6 Given that keeping the animals in an institutional situation allows for central control over breeding decisions, the most advanced systems for controlling genetic variability (i.e. minimal coancestry or optimal contribution theory – see Section 6) should be applied. Given the value of the animal genetic resources and substantial investment that the government or private donors will have made in establishing the conservation farm, exceptional effort must be taken to minimize the risk of diseases, accidents, genetic drift, inbreeding and contamination from other breeds. If artificial selection is being practised (or if natural selection may occur due to environmental differences between the natural and controlled environments), and because of the potential for genetic drift, it is possible that a genetic gap will develop between the conservation herd and the original breed in its native region and the population in the conservation programme. The potential for effects of this kind should be assessed and monitored. This “adaptation to captivity” is generally considered more important for wild animal species (e.g. Frankham, 2008), but may be relevant for livestock, especially those that come from a particularly harsh production environment.
Action 3. Establish a gene bank (i.e. in vitro cryobank) for periodic storage of germplasm from animals in the ex situ in vivo programme Maintaining genetic material from animals in the original population will allow recovery of the genetic variability if the population drifts from its original base and will facilitate reconstitution of the breed if a catastrophic event (e.g. disease, fire or natural disaster) wipes out a significant portion of the live population. As noted in Section 4, ex situ in vivo conservation (especially on a single farm) carries risks, as all the live animals may be threatened if a catastrophe occurs.
Action 4. Organize participation by livestock keepers and institutions in the production and use of male animals Ideally, it will be possible to use the ex situ population as a resource in the active management and improvement of the in situ population, for example, by making superior young males from the herd available for use in the general population.
Additional advice on ex situ in vivo management of animal genetic resources with very small populations can be found in guidelines produced by the European Livestock Breeds Ark and Rescue Net (ELBARN, 2009).
Dispersed ex situ conservation involving institutional herds and livestock keepers’ herds As noted above, the establishment and operation of an institutional farm for ex situ in vivo conservation of animal genetic resources can require a significant investment. One feasible means of overcoming this constraint is to expand the population through the use of a dispersed or decentralized model in which animals already available at established government farms are combined with those kept by NGOs and by private individuals who are willing to keep the animals on a commercial or hobby basis.
Many existing institutional facilities are already involved in important conservation activities. Nearly all could easily play a more significant role in conservation with little or no adverse effect on their other roles in development and animal breeding. The institutional farms could be associated with AI centres, nucleus herds and/or in vivo germplasm repositories. Livestock keepers would be both users and suppliers of germplasm. Facilities for producing frozen semen can be developed on such farms in order to disseminate the germplasm from pure-bred males typical of the breed to collaborating farmers. A network of several breed-specific herds can provide a basis for integrated breed conservation and systematic genetic improvement. The basic design of the model is shown in Figure 3.
An example of how certain institutional farms in India could be used for conservation is presented in Box 36.
CGRFA/WG-AnGR-7/12/Inf.6 Figure 3. An example of a decentralized programme for ex situ conservation of a breed by employing institutional herds belonging to private breeders and farmers Genetic repository Semen and/or embryos Tested males or Central nucleus semen Private Private •male test station breeder breeder •AI centre Candidate males •research/teaching and replacement facility females Private Private breeder breeder Private breeder Objective: Establish and maintain populations of important genetic resources in one or more institutional nucleus and decentralized breeding herds.
1. Lists of breeds that are candidates for ex situ conservation and descriptions of their characteristics;
2. Knowledge of the location of individual animals from these breeds including those belonging to private breeders and existing institutional farms;
and 3. Secured support from the government or a development agency.
• A network of institutional and local livestock keepers’ herds with an active programme for sustainable management of an at-risk breed of livestock.
Task 1. Establish the conserved population Action. Identify the base animals At least 25, and preferably more than 50, female animals should be identified from among the institutional herd and livestock keepers’ herds on the basis of breed characteristics and performance traits (production and adaptation). The animals should be tagged and recorded. Male animals (preferably at least one for every ten females) should also be identified and tagged and raised by their owners and made available to other livestock keepers for breeding purposes. The owners of candidate males may be provided with an incentive per female per year for rearing the animals and retaining pure-bred offspring.
CGRFA/WG-AnGR-7/12/Inf.6 Box The role of Gaushalas in conservation in India The Gaushalas of India are institutional self-contained cow shelters with their own land and housing facilities. They are usually supported by a combination of donations and government assistance.
There are currently more than 4 000 Gaushalas across India. Most of these Gaushalas primarily cater to the needs of non-lactating, weak, unproductive and stray cattle, but it is estimated that more than a quarter of them have the potential to be used for in vivo conservation (Sadana, 2007). Many Gaushalas in India maintain pure-bred animals of different local breeds, often in greater concentrations than can be found in surrounding herds belonging to local livestock keepers.
A few progressive Gaushalas are repositories of well-described local cattle breeds and produce quality males. They thereby contribute directly to the conservation and improvement of these breeds. However, they do not have sufficient resources and technical support to conserve and improve these breeds in the most effective manner. Undertaking the following actions could allow these progressive Gaushalas to be utilized more effectively for in vivo conservation:
• Identify the Gaushalas with a large number of pure-bred animals belonging to breeds at risk.
• Support the development of the infrastructure necessary to transform these Gaushalas from rehabilitation centres into genetic resource centres.
• Within each Gaushala, group the pure-bred and non-pure-bred animals and house them separately – choose the pure-bred animals selectively if population sizes permit.
• Implement identification, performance recording and breeding programmes to improve the pure-bred stock through selective breeding.
• Distribute excess pure-bred stock to the local community, targeting farmers that are willing to continue pure-breeding of the animals.
An agreement should be made with the participating Gaushalas not to resort to cross-breeding or other such practices that may dilute the genetic purity of the breed. In return, the Gaushalas could be provided with scientific and technical support and, if necessary, with financial assistance. The Gaushalas should be encouraged and supported in identifying unique and value-added products of the local breeds so as to increase their economic value.
Provided by Devinder K. Sadana.
Task 2. Manage the conserved population Various approaches can be used to manage the conserved population. Described below is a programme that is based on the distribution of male animals and/or semen.
Action 1. Mating of base animals to produce new males The base females should be mated or inseminated with a male or semen of the same breed. The livestock keepers can be contracted to rear the resulting male progeny for up to six months. A large number unrelated young males should be selected for inclusion in the conserved population (around one for every ten to twenty females). Hereafter, there are two approaches that can be followed, depending upon whether the animals are to be raised by livestock keepers or by a development agency or institutional farm.
Action 2. Manage the selection and distribution of males Option 1: Development agency or institutional farm If AI technology is available, the selected young males are purchased by the development agency or institutional farm. The agency rears the males until maturity and then trains them for semen production. At maturity, a total of at least 25 males are selected on the basis of growth, semen quality and freezability. As many as 3 000 doses of semen from each male may be collected, cryopreserved and used in for breed improvement and conservation (more prolific species will requires fewer doses).
The surplus breeding males are distributed to the farmers for natural mating in the breeding tract.
CGRFA/WG-AnGR-7/12/Inf.6 Option 2: Private livestock keepers Individual livestock keepers rear the selected males and receive an incentive payment for maintenance costs. When the males reach breeding age, the livestock keepers maintain them and provide breeding services to the female animals in the local area through natural mating. The farmers charge a fee for the breeding services in order meet the expenses of further maintaining the breeding males. At this point the incentive payments stop and the livestock keepers are expected to maintain the males solely on the basis of the breeding fees.
Action 3. Design breeding and mating strategies Selection of males and distribution of semen and males for mating should be implemented with the objective of maximizing genetic diversity according to the general theories described in Section 6 and in line with the conditions and technical capacity of the country. When technical capacity for applying complex breeding programmes is low, sire use should be uniform as much as possible (i.e. to achieve approximately equal numbers of offspring per sire). When technical capacity is greater, optimum contribution theory can be used in bull selection.
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CGRFA/WG-AnGR-7/12/Inf.6 FAO. 2011b. Surveying and monitoring of animal genetic resources. FAO Animal Production and Health Guidelines. No. 7. Rome (available at www.fao.org/docrep/014/ba0055e/ba0055e00.pdf).
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CGRFA/WG-AnGR-7/12/Inf.6 VI. DESIGNING THE CONSERVATION PROGRAMME – MAINTAINING GENETIC VARIABILITY As described in the preceding sections, a population’s degree of risk and consequently its probability of survival are greatly dependent on the level of genetic diversity it harbours. High levels of genetic variability allow the population to adapt to changes in the environment or the production system and will prevent the rise of inbreeding and its deleterious consequences. Small populations, such as those that are likely to be targeted by conservation programmes, are more prone to lose genetic information.
Therefore, implementing management strategies that maintain genetic diversity is crucial to the success of conservation programmes.
Two strategies can be used to manage the genetic diversity of small populations. The first of these is to maintain or increase the genetic variability within the population (this section):
1. adopt a general breeding strategy to maintain the conserved breed, based on a clear understanding of options for maintenance of the genetic variability;
2. consider adopting a mating strategy to decrease inbreeding;
and 3. incorporate cryoconservation in the management of genetic variation in the in vivo programme.
The second potential strategy is to optimize selection response and genetic variability within the populations (Section 7) 1. adopt a general breeding strategy to maintain the conserved breed;
and 2. design a breeding programme that generates genetic improvement while maintaining genetic variability.
In most cases, the choice between the two strategies will depend upon the population size – either census size or N e. Genetic improvement runs counter to the objective of maintaining of genetic variability, so maintenance of variability will take precedence in small populations. As described in Section 2 (Task 3), maintaining variability should usually be emphasized for breeds with critical or endangered risk status, especially for the former. For vulnerable populations, the genetic improvement strategy becomes more realistic. However, the maintenance of genetic variation should always be an essential part of a breeding programme.
Maintaining genetic variability within small populations Rationale Loss of genetic information In livestock populations of a stable size, the loss of genetic information (alternative alleles contributing to the genetic variability in the population) is usually caused by selection and genetic drift. In a very general sense, natural selection acts to eliminate deleterious alleles, while artificial selection tends to fix alleles that improve the phenotype of carriers. In small populations not subject to an explicit process of artificial selection, the impact of selection (either natural or artificial) is small (most of the genetic variants are neutral with respect to phenotype) and the fate of an allele (i.e. its eventual loss or fixation) is mainly driven by genetic drift, which is a random process. Genetic drift is the fluctuation of the frequencies of alleles (i.e. the number of copies of them in the population) due to the finite and random sampling of gametes to generate offspring (Falconer and Mackay, 1996). This random sampling operates at two levels. First, if the number of offspring is small and reproduction is not controlled (i.e. mating is random, resulting in variable numbers of offspring per individual) some members of the population may not contribute offspring to the next generation. The unique genetic information of these animals will be lost, while other animals will contribute multiple copies of their genetic information to the next generation. Second, to generate an offspring, each individual contributes a gamete carrying just one of the two alleles at each position in its genome. If the parent is heterozygous (i.e. it carries two different alleles at a given locus) only one of the two variants will be CGRFA/WG-AnGR-7/12/Inf.6 transmitted to a given offspring. This may result in the loss of genetic information even if all individuals leave offspring.
Consequences of the loss of genetic information The consequences of the genetic drift process for small populations are:
• the probability increases that none of the copies of a particular allele will be transmitted to the next generation (i.e. genetic information will be lost);
and • the probability of mating between relatives (i.e. inbreeding) increases, and so does the probability that animals will inherit identical copies of an allele in any gene.
An increased proportion of mating between relatives occurs in small populations even when mating is random, and the probability of intra-family mating increases as the size of the population decreases.
Of course, inbreeding arises in populations of any size if relatives are mated deliberately. The probability of homozygosity by descent is greater the higher the average degree of relationship between individuals in the population. Increased homozygosity reduces the expression of fitness related or productive traits, compromising the survival of the population, a phenomenon known as “inbreeding depression” (see Box 37).
Box Inbreeding depression Livestock species are diploid, which means that each individual carries two copies of the genetic information (two alleles) at each position (locus) in its genome. At a particular locus, the two alleles can be the same (homozygous) or different (heterozygous). The performance of an individual for a given trait will depend on the type of alleles it carries. Sometimes an allele gives a visible effect only if it is in homozygosity (i.e. when both alleles are the same). Such alleles are called recessive. If the allele has deleterious effects (confers inferior expression of the trait) heterozygous individuals will have a normal performance, but they will be carriers of the deleterious allele. Genetic drift promotes an increase in the number of homozygotes. The deleterious alleles that were undetected because of the compensatory effects of heterozygosity become more frequently exposed and the mean value for the trait decreases. This decline in mean performance of the population is called inbreeding depression. If the allele has effects on fitness-related traits, the consequence is lower probability of survival. If the trait controlled by the locus involves productivity, the mean performance of the population will decrease and the profitability of the breed will be compromised. In either of these situations (decreased fitness or decreased profitability), the risk of extinction will be increased. Thus, small population sizes lead to both genetic and demographic consequences (see Table 6).
Inbreeding coefficient The inbreeding coefficient (F) is a measure of diversity that ranges from 0 (non-inbred) to 1 (a completely inbred and homozygous individual, with two exact copies of all chromosomes). Inbreeding is unavoidable in small populations and if the pedigree can be traced far enough back, common ancestors will invariably be found, demonstrating that all animals are related. Therefore, the average F of a population is dependent on the definition of a reference/base population in which all individuals are assumed to be non-inbred and unrelated (e.g. founder individuals when pedigree recording started).
Consequently, populations with many generations of genealogy will tend to have high average F, whereas populations with shallow pedigrees will have low F.
Because of this bias caused by differences in the depth of pedigree data, a more informative parameter is the rate of inbreeding (F), defined as the change of inbreeding per generation relative to the amount of inbreeding that can still occur (i.e. 1 - F). The advantage of this concept is that it is independent of a reference population, allowing comparison between populations whose histories are known to different degrees. Under several assumptions, F can be calculated using simple formulae.
Calculating F helps in predicting the future performance of a population, determining the minimum CGRFA/WG-AnGR-7/12/Inf.6 composition of a viable population or designing management strategies. Further details of these methods are provided later in this section.
Rate of inbreeding F is also a useful parameter for describing the present situation of a population. It allows detection of events that occurred in the history of the population (e.g. bottlenecks, or periods when the population consisted of a reduced number of individuals) and also helps to determine the risk status of the breed (see Section 2). Additional genealogical analyses can also be undertaken to study the history of a population from the genetic perspective. These analyses yield parameters such as effective number of founders, founder genome equivalents and founder representation (Caballero and Toro, 2000) that help to determine how much variation the population had at its origin and how well the population has been managed in the past.
Effective population size As explained in Section 2, effective population size (Ne ) is a parameter that is commonly used to evaluate the genetic variability of a population, based on the assumption that larger populations will be less subject to random drift and will thus have more genetic variability. To review, Ne is the size of an idealized population that has the same rate of inbreeding (F) as the real population. This idealized population has equal number of males and females, all of which have an equal opportunity to produce offspring. It is a theoretical concept – such a population would never really exist in a livestock breed. However, it serves as a basis for comparison. Various types of departure from the characteristics of the idealized population will affect the calculation of N e. This increases the difference between the number of animals in the population (the census size) and the N e (see Box 7 for a discussion of factors that affect the determination of N e ).
There is a close relationship between F and Ne (F = 1/2Ne ) and, consequently, both parameters describe the same concept. For most situations, the loss of genetic variability (as measured by the parameter genetic variance) is also proportional to F, which indicates that this parameter is a useful measure of the ability of a conservation programme to maintain genetic variability.
Relationship and coancestry coefficient The relationship between individuals (measured using the coancestry coefficient, f, i.e. the probability of sampling alleles identical by descent from two individuals) is another helpful parameter because of its connection with classical measures of diversity. The global coancestry of a population reflects the extent to which the genetic information found in the different individuals is redundant. Mathematically, the average population coancestry is = (1 - H e ) where H e is the expected heterozygosity. As described in Section 3, H e is a common measure of genetic diversity. When the numbers of males and females in the population are different, the global coancestry must be calculated as of the mean coancestry between pairs of males, plus of mean coancestry between females, plus of the mean coancestry between every possible pair of sire and dam. The coancestry coefficient is also related to inbreeding because the F of the offspring is the f of the parents;
an individual cannot carry alleles that are identical by descent if its parents did not share the allele, as each of the alleles at a locus comes from a different parent.
Rate of inbreeding and rate of coancestry Rate of inbreeding (F) is related to the probability of homozygosity by descent, and rate of coancestry (f) is related to loss of diversity. Under random mating, both parameters equalize in a few generations. However, if the population is subdivided or if assortative mating is practised (i.e. if mating of relatives is promoted or avoided), F and f can diverge (Falconer and Mackay, 1996).
Consequently, to determine the genetic endangerment of a population, N e should be calculated from both F and f. A population subdivided into several genetically isolated lines (e.g. through geographic isolation or deliberate breeding within families) may harbour high levels of population wide diversity (low levels of global f), but will suffer from the harmful effects of large F within in each subpopulation.
Both F and f can be easily calculated from genealogies. Consequently, it is highly recommended that, when the production system and physiology of the species allow, the pedigree of the population be CGRFA/WG-AnGR-7/12/Inf.6 traced through the generations by recording the sire and the dam of every individual. Pedigree recording will allow for the implementation of very effective management strategies (see later in this section and in Section 7) and will avoid the need to resort to more expensive methodologies such as molecular marker genotyping. Several computational methods have been developed to calculate F and f in any pedigree, including some that can deal efficiently with large genealogies. Free software is also available to perform such calculations. Two examples are ENDOG 8 (Gutirrez and Goyache, 2005) and POPREP 9 (Groeneveld et al., 2009), but many others have been developed (some of which are listed in Boettcher et al., 2009). Such programs tend to provide more informative and reliable results (and greater estimates of F) when pedigrees include more generations of data.
Minimum effective population size The minimum acceptable N e has been defined as the N e of a population that is safeguarded from becoming extinct because of the effects of inbreeding depression (or other threats related to reduced genetic variability). In general, 50 has been established as an acceptable N e, at least to guarantee the survival of the population in the short and medium term. Consequently, the desired F per generation should not exceed 1 percent (F = 1/). This can be achieved with different population structures in terms of combinations of males and females. For example, with no selection (i.e. random number of offspring per parent) 25 breeding males and 25 females yield the desired value, but a decrease in the number of males has to be compensated for by increasing the number of females. For example, males and 34 females, and 14 males and 116 females, also yield an N e of 50 (for information on the computation of N e see Box 7). The management procedures implemented in the population will affect the numbers of breeding animals needed for an N e of 50. The required numbers will be lower when only conservation strategies are applied and higher when selection for a particular trait is applied (see Section 7). Of course, the population sizes described here are minimum sizes. Larger numbers of animals are always to be preferred for long-term survival of a population.
Direction of management strategies From the above discussion it can be concluded that management strategies should aim to minimize genetic drift effects by minimizing F (f) or maximizing N e. The knowledge of the factors affecting N e will aid in the design of effective conservation strategies. When the N e of a population drops significantly lower than 50, it may reach a level where it cannot be sustained because of the negative effects of a lack of genetic diversity on fitness and fecundity and the accumulation of genetic defects.
Such a situation is referred to as an extinction vortex, because population numbers are expected to decrease uncontrollably in each successive generation (like water draining from a sink or bathtub). For such populations, a more radical strategy of “genetic rescue” must be applied (see Box 38).
Objective: Understand the factors associated with genetic drift and develop strategies to minimize its occurrence.
Knowledge of the following characteristics of the breed to be conserved:
1. Size of the population;
2. Reproductive capacity of the species;
and 3. Possibilities for exchange of genetic material between herds or flocks belonging to different stakeholders.
1. A general breeding plan that will minimize genetic drift and maintain genetic diversity.
http://www.ucm.es/info/prodanim/html/JP_Web.htm http://popreport.tzv.fal.de CGRFA/WG-AnGR-7/12/Inf.6 Box Genetic rescue When a population is not fit enough to reproduce itself and, thus, the number of breeding animals is irrevocably decreasing in every generation, the population has become trapped in an extinction vortex. Such a situation often arises because of excessive inbreeding in the past (a bottleneck), a great drop in genetic variability levels and the accumulation of genetic defects. If a breed enters extinction vortex, two strategies can be implemented.
One option is to change the environment of the population (e.g. by establishing an in vivo ex situ programme) to a more favourable one where the animals can receive greater veterinary care and the fitness of individuals is high enough to survive and reproduce. Attention must be paid to ensuring that the population does not become adapted to the new, more favourable, conditions, as this may preclude the reintroduction of the population to its former production environment.
A second alternative is limited crossing with another breed that is adapted to a similar environment and ideally has specific adaptive traits that are similar to those of the at-risk breed. This process is known as genetic rescue. The number of introduced individuals should be kept to a minimum, but even a small amount of foreign (i.e. from a different breed, not necessarily from a different country) genetic material can have a large positive effect. For example, if a proportion p of foreign alleles is introduced, the proportional reduction in inbreeding is 1 - (1 - p)2. With the introduction of 10 percent foreign alleles, the inbreeding in the population is reduced by nearly 20 percent, depending on the initial F. For example, if F = 0.30, an introduction of 10 percent of outside alleles leads to a population with F = 0.24. The whole process has to be very carefully controlled so as to avoid excessive introgression of foreign genetic information. Among the cross-bred offspring, those showing the original phenotype should be selected to create the next generation by backcrossing with pure-bred individuals from the at-risk breed, until most of the foreign genetic information has been removed. Molecular markers can be used to increase the accuracy of selection decisions aimed at purging foreign alleles.
Task 1. Adopt a general breeding strategy to maintain the conserved breed, based on a clear understanding of options for maintaining the genetic variability of the population Logistic and financial capabilities vary from one herds or population to another. Therefore, not all the strategies presented in the following pages will be feasible in every situation. The simplest actions are generally presented first, followed by more technical and sophisticated options. Breeders and other managers of animal genetic resources should decide which options are most appropriate for their particular situations. The first three actions are likely to be applicable in most conservation programmes.
Action 1. Include as many animals as possible from the start, as the extent of genetic drift depends on the number of individuals available Animals should be healthy and, as far as possible, non-inbred and unrelated. However, related animals should not be eliminated if the capacity and financial resources of the programme allow them to be maintained. Efforts should be made to involve all the herds of the breed in the programme. Doing so will enable the programme to begin by exploiting the maximum possible variability and opportunities to diminish the action of genetic drift. In addition, females that have previously or routinely been crossed to males of other breeds should be recovered and used exclusively for within-breed matings.
Clearly, if conditions permit, the real population size should be increased as quickly as possible to reduce the risk of extinction due to demographic stochasticity and to increase the N e.
Action 2. Increase the number of breeding males Half of the genetic information is transmitted by each sex. Therefore, the less-represented sex (usually the males) will be the deciding factor in decreasing N e, irrespective of the number of individuals of the other sex that there are in the population. For example, assuming random numbers of offspring across males and females, a population with 2 males and 1 000 females has the same N e as a population with 4 males and 4 females. With just one male, all the descendants will be at least half-sibs and average F will increase from zero to more than 0.125 in just one generation.
CGRFA/WG-AnGR-7/12/Inf.6 Action 3. Prolong the generation interval The F is defined per generation, as genetic drift occurs due to the random sampling of alleles when gametes are produced. However, if the generation interval (i.e. how long it takes to replenish a set of parents – see Box 39) is longer, the same F per generation has to be divided among more years, diminishing the loss of genetic diversity per year. Generation interval can be increased by keeping individuals as long as they are able to reproduce and by extending their period of use by using their cryopreserved genetic material. As long as an individual or its genetic material is available for breeding, its genetic information is not lost and can contribute to enhancing the genetic variability in the breed.
Box Generation interval The generation interval (L) is the genetic unit of time for a population. It is defined as the average age of parents at the time of the birth of their straight-bred replacement. In many cases this parameter will be approximated by the average age of the parents at the birth of all the offspring, but this need not to be so. Due to differences in reproductive life spans, the generation interval may be different for males and females, and should be calculated separately. Because half of the alleles are contributed to the population by males and half by females, the generation interval is the average generation interval of the breeding males and the breeding females. For example, if offspring are born when the sires are one year old, and 60 percent and 40 percent of the offspring are born when dams are one and two years old respectively, L s = 1 and L d = 0.6*1 + 0.4*2 = 1.4 years (s = sire and d = dam). Calculating the average gives a figure of 1.2 years for the L of the population. The longer individuals are used as breeding animals the greater L will be.
However, it is important to note that when a programme of selection and genetic improvement is implemented, an increased generation interval will reduce the response to selection per year. Thus, once a breed’s population has increased to a size that is sufficiently large to allow selection (e.g. females and N e > 50), generation interval has to be balanced against the other factors influencing genetic gain and maintenance of variability. This topic is addressed in Section 7.
Action 4. Balance the contribution of each individual The rationale of this action is to provide the same opportunity to all animals to transmit their genetic information to the next generation. In a simple situation where the numbers of males and females are equal and no selection is practised, the effective population size is approximately N e = 4N / (2 + S2 k ), where N is the census size and S2 k is the variance in the number of offspring contributed by each individual. If individuals’ contributions are equalized, S2 k becomes zero and the N e is twice the population census size (the largest possible N e ). In simple terms, equalizing contributions can be realized by obtaining one son from each male and one daughter from each female. However, such planning can be realized only under highly controlled conditions.
Regular selection and mating systems When conditions permit, Action 4 can be applied in a “regular” hierarchical system. In such a system, an equal number of females are mated to each male in every generation. The general idea is still to equalize contributions to the next generation and maximize the probability that every individual transmits its genetic information. The strategy works by performing a type of within-family selection (i.e. selecting the best of the sibs from each family) so that one male is obtained from each male family and one female from each female family (Gowe et al., 1959). Under this procedure, the formula for the rate of inbreeding is F = 3/(32N M ) + 1/(32 N F ) (where N M and N F are the numbers of breeding males and females respectively) which is less than the F obtained with random contributions:
F = 1/(8 N M ) + 1/(8 N F ).
This strategy can be refined so as to get an even smaller F by controlling not only the numbers of offspring per parent but also the contribution of each individual to its descendants across generations (Snchez-Rodrguez et al., 2003 – see Table 8).