1 Trypanotolerance in small ruminants of sub-Saharan Africa 2 3 4 Stanny Geerts1,2, Sabine Osaer3, Bart Goossens4, Déthié Faye5 5 6 7 8 9 1. Institute of Tropical Medicine, Animal Health Department, Nationalestraat 155, B-2000 Antwerp, Belgium 10 2. International Trypanotolerance Centre, PMB 14, Banjul, The Gambia 11 3. Boterbloemlaan 13, B-1933 Sterrebeek, Belgium 12 4. UNOPS, Endemic Livestock Project, P.O.Box 4060, Bakau, The Gambia. 13 5. BP 53, Butare, Rwanda 14 15 16 17 Corresponding author: Geerts, S. (sgeerts@itg.be) 18 1 1 Abstract 2 Although a lot of information is currently available on trypanotolerance in cattle, until 3 recently the trypanotolerant nature of small ruminants was not well known. 4 Trypanotolerance in small ruminants is less pronounced than in cattle and should 5 rather be considered as resilience than as resistance. West African Dwarf (WAD) 6 goats appear to be less trypanotolerant than Djallonke sheep. However, recent studies 7 have shown that there is an important introgression of genes of trypanosusceptible 8 breeds into WAD goat populations, which possibly explains the loss of 9 trypanotolerance in these animals. Measures need to be taken to safeguard and 10 upgrade the genetic purity of trypanotolerant goat and sheep breeds in Africa. 11 12 13 Trypanotolerant sheep and goat breeds Small ruminants play an important role in the rural economies of Africa. They 14 are kept mainly to generate income, as savings and for ceremonial purposes. In some 15 countries such as The Gambia, the use of goat milk for home consumption is also 16 important [1]. Sheep and goats are frequently managed by women and children. 17 Agyemang [2] estimates that about 32% of the sheep and 47% of the goats in West 18 and Central Africa are trypanotolerant. The breeds concerned are the Djallonke sheep 19 (Figure 1a) and the West African dwarf (WAD) goats (Figure 1b) . Both are dwarf 20 breeds with a height range of 30–50 cm for the goats, and 40–60 cm for the sheep. In 21 East Africa the Small East African (SEA) goats and the Red Maasai sheep are 22 considered as trypanotolerant [3;4]. They are slightly taller than the West African 23 breeds, and are not considered as dwarf breeds [5]. 24 Trypanotolerance is the ability to survive, reproduce and remain productive 25 under trypanosomiasis risk without the need for the use of chemicals to control the 2 1 vector or drugs to control the parasite [6]. It offers farmers an important option for 2 sustainable livestock production in areas at risk from trypanosomiasis. Contrary to the 3 widespread belief that these trypanotolerant breeds are less productive than others due 4 to their small size, it has been convincingly demonstrated that the productivity of 5 trypanotolerant cattle and small ruminants is similar or even better than that of 6 trypanosusceptible breeds when kept under similar conditions [7;8]. For example, the 7 productivity of Djallonke sheep and WAD goats kept under tsetse fly challenge was, 8 respectively, 42% and 68% higher than that of susceptible breeds, even when the 9 susceptible breeds (e.g. Sahelian) were maintained in tsetse free conditions [7]. 10 Over the past decade the trypanotolerant nature of small ruminants has been 11 studied in depth at the International Trypanotolerance Centre (ITC, Banjul, The 12 Gambia) and elsewhere. This review summarizes and critically analyses the available 13 information on trypanotolerant sheep and goats in sub-Saharan Africa. 14 15 The trypanotolerant character in small ruminants 16 17 Griffin and Allonby [3;4] were the first to show – both under experimental and 18 field conditions in Kenya - the trypanotolerant character of some indigenous sheep 19 and goat breeds. Comparison of indigenous SEA and Galla goat breeds with exotic 20 Saanen and Saanen x Galla crossbreed showed that SEA were more tolerant to 21 trypanosome infections than the Galla whereas the Saanen were highly susceptible 22 and the crossbreeds showed a tolerance intermediate between Saanen and Galla. 23 Similarly Red Maasai sheep exhibited a much higher tolerance to Trypanosoma 24 congolense than Blackhead Persian or Merino sheep. Although Whitelaw et al. [9] in 25 different breeds of goats in Kenya were not able to confirm the observations of Griffin 3 1 & Allonby [3;4], other studies strongly indicate that SEA goats [10;11] and Red 2 Maasai sheep [12;13] can be considered as trypanotolerant. Mutayoba et al.[14] 3 compared SEA goats from different areas of East Africa and concluded that there 4 might exist many varieties or types within the SEA goat population, some of them 5 being more trypanotolerant than others. 6 Several authors have shown that WAD goats and Djallonke sheep are more 7 resistant to trypanosome infections than other breeds (such as Sahelian or Red Sokoto 8 goats, Sahelian or Fulani sheep) or cross-breeds (F1 Sahelian-Djallonke) [15;16]. 9 Although, in some experiments, the resistance of trypanotolerant small ruminant 10 breeds was significantly better than trypano-susceptible breeds only for T. vivax, and 11 not for T. congolense [17]; in other experiments1, the opposite was true . These 12 differences might be explained by the trypanosome strain which was used, as it is now 13 well known that the pathogenicity of T. congolense varies widely from one subgroup 14 (savannah, forest, kilifi) to the other [18], or even within the savannah subgroup [19]. 15 Surprisingly, Cameroonian dwarf goats experimentally infected with T. brucei all 16 died, whereas no clinical signs developed after infection with T. congolense or T. 17 vivax [20]. Very severe anaemia and weight loss with no tendency to recovery were 18 also reported in Nigerian Djallonke ewes infected with T. brucei [21]. A similar 19 susceptibility of WAD goats to T. brucei was not confirmed in other experimental 20 infections [22] and was, to the best of our knowledge, never reported from the field 21 [23]. 22 1 Touré,S.M., Seye,M., Mbengue,M. and Dieye,T. (1983) Trypanotolerance: studies of comparartive pathology on dwarf Djallonke sheep and Sahelian Fulani sheep. In Proceedings 17th ISCTRC Conference (OAU-IBAR, ed), pp. 326-336, OAU 4 1 Genetic origin and mechanisms of trypanotolerance in small ruminants 2 The genetic origin of the trypanotolerance trait in East African Galla goats 3 was illustrated by the demonstration that Saanen x Galla crosses showed an 4 intermediate tolerance between Saanen and Galla [3]. Similarly, by comparing pure 5 bred with cross-bred animals, Kanyari et al. [10] and Goossens et al. [16] showed the 6 genetic basis of trypanotolerance in SEA goats and Djallonke sheep, respectively. By 7 contrast Faye et al. [24] were not able to show any difference in the degree of 8 parasitaemia, anaemia and antibody response between WAD goats and their F1 9 crosses (WAD x Sahelian). However, it is important to note that the WAD goats used 10 in this study were selected on the basis of their phenotype. This experiment should be 11 repeated using genetically characterised animals given the recent observations2 on an 12 important degree of introgression of Sahelian genes into the WAD goats. The same 13 remark can be made for most of the other studies since the animals were selected 14 purely on phenotypic criteria. 15 Contrary to cattle few research has been carried out to identify the 16 mechanisms of trypanotolerance in small ruminants. Very elegant studies using 17 haemopoietic chimaeras combining trypanotolerant (N’Dama) and susceptible 18 (Boran) genotypes have shown that trypanotolerance in cattle is a trait that involves 19 two at least partly independent mechanisms, i.e. the control of parasitaemia and the 20 control of anaemia. The control of parasitaemia does not depend on the haemopoietic 21 tissue and thus not on the immune system whereas the control of anaemia is 22 dependent on the haemopoietic tissue genotype [25]. There are indications that similar 2 Hoeven,E., Fidalis,M.N., Leak,S.G.A., Geerts,S., Hanotte,O. and Jianlin,H. (2007) Introgression of the Sahelian breed into West African dwarf goats. In Proceedings 28th ISCTRC Conference (Njogu,A.R., ed), pp. 622-626, AU 5 1 discrete mechanisms controlling anaemia and parasitaemia might be present in 2 Djallonke sheep [16], but this remains to be proven. Concerning the immune response 3 in trypanotolerant animals most of the available data are derived from experiments in 4 N’Dama cattle and show that there is no difference between trypanosusceptible and 5 trypanotolerant cattle as far as antibody response to variable surface glycoproteins 6 (VSGs) of trypanosomes is concerned [26]. However, significant differences do exist 7 in antibody titres to non-surface exposed antigens (e.g. congopain) [26;27] which 8 might be involved in the neutralisation of toxic or pathogenic molecules of parasite 9 origin and thus contribute to trypanotolerance. To our knowledge similar experiments 10 to examine the type of antibody response in trypanotolerant small ruminants have not 11 yet been carried out, although significantly higher levels of anti-trypanosomal 12 antibody titres were reported in Djallonke sheep as compared to Djallonke-Sahelian 13 crossbreds [16]. 14 15 16 17 Trypanotolerance: resilience rather than resistance By contrast to trypanotolerant N’Dama cattle, which are able to control 18 parasitaemia quite well and which show a strong tendency to self-cure [12;28], the 19 nature of trypanotolerance seems to be different in small ruminants. Several studies, 20 both under experimental and field conditions, have shown that WAD goats and 21 Djallonke sheep which were infected with T. congolense or T. vivax were able to gain 22 weight, but showed a persistent parasitaemia [23;29;30]. In a long-term study using 23 WAD goats and Djallonke sheep experimentally infected with a virulent T. 24 congolense strain self-cure was only observed after 18 to 24 months [31]. This clearly 25 suggests that ‘resilience’ is a more appropriate term to characterise trypanotolerance 6 1 of sheep and goats than ‘resistance’. Resilience means that the animals are able to 2 ‘live with the trypanosome’ by limiting its pathological effects, the most prominent of 3 which is anaemia. Concerning the control of anaemia, the results of experimental 4 infections of small ruminants with T. congolense show that the hematocrit values in 5 Djallonke sheep and WAD goats are significantly less affected than in non- 6 trypanotolerant animals [15;17]. However, the erythropoietic response in Djallonke 7 sheep and certainly in WAD goats seems to be less efficient than in N’Dama cattle 8 [30]. 9 Concerning the genetic basis of resilience and the feasibility of breeding 10 programmes for increased resilience there are only few data available for 11 trypanotolerant small ruminants. Much research, however, has been done in the field 12 of resilience to nematode infections in sheep [32-34]. According to Woolaston and 13 Baker [32] breeding for resilience applies less selective pressure on the parasite than 14 breeding for resistance because the emphasis is on co-existence of the host and the 15 parasite . Nevertheless, Morris et al. [35;36] have shown that selection for resilience 16 to nematode parasite challenge in Romney sheep flocks is possible although very 17 time-consuming under commercial conditions. Compared with resistance, resilience 18 to internal parasites appears to be somewhat less heritable but at least as variable 19 [32;33]. In the field of trypanotolerance results are only available from the ITC’s 20 breeding programme. This programme is carried out under village circumstances and 21 aims to increase the efficiency of meat production and the resilience of WAD goats 22 and Djallonke sheep [37]. The heritability estimates for body weight and growth rate 23 traits for both species suggested that substantial improvement could be achieved by 24 selection. Estimates of heritability for weight at birth, at weaning and at yearling age, 25 pre-weaning and post-weaning growth rates were 0.5, 0.43, 0.30, 0.32 and 0.11 for 7 1 WAD goats and 0.39, 0.54, 0.21, 0.54 and 0.23 for Djallonke sheep, respectively [37]. 2 Unfortunately hematocrit measurements were not included in the latter study. This 3 would have allowed an even better estimate of the effectiveness of the breeding 4 programme for improved resilience. 5 6 Higher trypanotolerance in cattle than in small ruminants 7 Although comparative studies of the degree of trypanotolerance between 8 N’Dama and trypanotolerant small ruminants have not been carried out under 9 experimental conditions, a large amount of data collected in various regions of The 10 Gambia show that the prevalence of trypanosomiasis in N’Dama is quite similar to 11 that found in Djallonke sheep and WAD goats when exposed to a similar 12 trypanosomiasis risk [38]. However, it should be noted that small ruminants are less 13 exposed to tsetse challenge than cattle because of their different grazing pattern 14 (closer to the villages than cattle). This is supported by bloodmeal analysis of tsetse 15 flies which shows that tsetse feed significantly less on small ruminants than on cattle 16 [38]. Furthermore, Vale [39] showed that the feeding success of tsetse flies is also 17 much higher on cattle (35%) as compared to sheep and goats (1%) due to the anti- 18 feeding behaviour of the goats (including leg kicking and stamping, tail and ear flicks, 19 skin rippling). Consequently, a similar prevalence of trypanosomiasis in 20 trypanotolerant cattle and small ruminants living in the same environment strongly 21 indicates that sheep and goat are indeed less trypanotolerant than cattle and points to a 22 longer duration of the parasitaemia and a less efficient self-cure in the sheep and 23 goats. 24 25 WAD goats are less trypanotolerant than Djallonke sheep 8 1 Several experiments carried out at the ITC in The Gambia have shown that 2 WAD goats are less trypanotolerant than Djallonke sheep. After experimental 3 infection with two T. congolense strains with different pathogenicity the anaemia was 4 always more severe in WAD goats than in Djallonke sheep [29]. Furthermore, 5 Goossens et al. [30] found a better haemopoietic response and a significantly higher 6 antibody titre in Djallonke sheep than in WAD goats after experimental infection with 7 a virulent West African strain of T. congolense. Follow-up for two years after an 8 artificial infection with T. congolense showed that the productivity index of infected 9 Djallonke sheep and WAD goats was significantly lower than that of control animals 10 in the first year. However, infected Djallonke sheep became as productive as the non- 11 infected control animals in the second year, whereas the productivity of infected 12 WAD goats was only half of that of the control goats (Table 1), which again proves 13 the superior trypanotolerance of Djallonke sheep as compared to the WAD goats [31]. 14 These observations were confirmed during a large-scale two-year field study in two 15 areas, one with a high and the other one with a moderate tsetse challenge in The 16 Gambia [23]. Although the prevalence of trypanosomiasis (predominantly T. vivax, 17 followed by T. congolense and few T. brucei) was systematically higher in Djallonke 18 sheep than in WAD goats, the mortality and the abortion rates were higher in goats 19 than in sheep [23]. Both sheep and goats remained productive under moderate and 20 high levels of trypanosome challenge, but the WAD goats were more affected 21 (reduced haematocrit, weight loss) than the Djallonke sheep [23]. 22 The weak trypanotolerance of WAD goats from The Gambia was also 23 confirmed by Faye et al. [24] and Dhollander et al. [40], who compared pure WAD 24 goats with F1 crosses with Sahelian and Saanen goats, respectively. After 25 experimental infection with T. congolense there was no significant difference in the 9 1 levels of anaemia and parasitaemia between WAD and F1 crosses and the weight gain 2 was equal or even higher in the crosses than in the apparently (based on phenotype) 3 pure breeds. 4 5 6 Introgression of non-trypanotolerant genes in WAD goats All the previously mentioned studies were carried out with animals that were 7 not genetically characterised. Therefore, they have to be interpreted very cautiously in 8 the light of recent work on the genetic background of WAD goats . Hoeven et al. (see 9 footnote 2), using 15 microsatellites, examined 615 individuals from 17 indigenous 10 West-African goat populations from five West-African countries. Principal 11 component and admixture analysis indicated that there is a gradient of introgression of 12 Sahelian genes within the WAD goat populations consistent with the geographical 13 location and tsetse distribution. A first gradient was observed from north to south 14 (north to south Senegal and Guinea-Bissau), and a second gradient from east to west 15 (from Mali to Guinea) (Figure 2). Apparently, although the goats phenotypically 16 resembled dwarf goats, 31% to 53% of the genes of the four examined Gambian 17 populations were of Sahelian origin, which might explain the loss of trypanotolerance 18 in these WAD goats. Similar studies on the genetic background of other breeds of 19 trypanotolerant small ruminants would be very useful. 20 21 Trypanotolerance in small ruminants is a relative phenomenon 22 23 There are several factors that might have a negative impact on the 24 trypanotolerance trait of small ruminants, such as concurrent helminth infections, 25 inadequate nutrition level and the severity of the trypanosome infection. 10 1 In a controlled experiment, Djallonke sheep were able to withstand a single infection 2 with T. congolense, but when exposed to an additional Haemonchus contortus 3 infection, some animals lost their capacity to control the pathogenic effects of both 4 parasitic infections, which led, in some cases, to mortality because of their additive 5 effect [41]. The interaction between trypanosome and helminth infections was 6 evidenced by a higher eggs per gram of faeces (EPG) in Djallonke sheep infected with 7 both pathogens, confirming the immunodepressive effect of trypanosomiasis. This 8 effect was also demonstrated in WAD goats by Faye et al. [24], who observed a 9 significantly lower antibody response to H. contortus in T. congolense infected 10 animals than in otherwise non-infected animals. Osaer et al. [23;42] confirmed these 11 findings under natural field conditions with Djallonké sheep and WAD goats living 12 under high trypanosomiasis risk. 13 Dietary supplementation was acting independently but beneficially on packed 14 cell volume (PCV) level and weight gain. Hence, it diminished the severity of 15 anaemia, and reduced the weight loss caused by helminths and/or trypanosome 16 infection [41]. This was also observed in an on-farm intervention trial under high 17 trypanosomiasis risk [42]. Improved dietary levels in Djallonke ewes interacted 18 positively with trypanosome infection on some haematological and productivity 19 parameters, including reproductive performance [43]. Goossens et al. [30] further 20 showed that trypanosome-infected Djallonke sheep, when adequately fed, mounted a 21 better haematological response in the acute phase, as compared to infected WAD 22 goats. This was confirmed by Katunguka et al. [44] who also described a better 23 erythropoietic response in trypanosome-infected sheep fed with higher protein 24 supplements. This proves the beneficial effects of nutrition in the control of anaemia, 25 which is an important feature in trypanotolerance, as previously observed in N’Dama 11 1 cattle [45]. On the other hand, some studies carried out in goats and sheep showed that 2 trypanosome-infected animals under high protein level of diet had a higher 3 parasitaemia score than those under low level [44;46]. This was explained by an 4 enhancement of the growth and energy metabolism of trypanosomes due to the high 5 concentration of serum cholesterol and non-esterified fatty acids generated from the 6 diet [44]. 7 The severity of the trypanosome infection (virulent strain and/or natural high 8 trypanosomiasis challenge) may also affect the level of trypanotolerance in small 9 ruminants, as observed by Osaer et al. [29] and Goossens et al. [30]. Whitelaw et al. 10 [9] and Faye et al. [24] were not able to show any difference in susceptibility between 11 supposedly ‘susceptible’ and ‘resistant’ breeds of goats after infection with a highly 12 virulent trypanosome strain as determined by decrease in PCV. 13 The above studies indicate that all these factors (trypanosome infection, 14 species and strain; inadequate nutrition; helminth infection) act independently but all 15 contribute to the severity of the anaemia. Since the degree of trypanotolerance in 16 small ruminants seems to be linked to the activity of the haematopoietic response 17 [3;9;15;30;47] it is evident that all factors with a negative influence on the 18 erythropoietic response, will also have a negative outcome on the degree of 19 trypanotolerance. 20 21 22 Resistance to ticks and helminths Many studies have been carried out to examine the genetic resistance of sheep 23 and goat breeds to gastro-intestinal (GI) nematodes. This review, however, focuses 24 only on trypanotolerant breeds. Both under experimental and field conditions, it has 25 been shown that there exists a marked individual variability in response to infections 12 1 with H. contortus with a large percentage (about two thirds) of Nigerian WAD goats 2 being resistant and a small proportion (less than 10%) being susceptible to infection 3 [48-50]. Although a genetic basis for ‘haemonchotolerance’ has not yet been 4 established, the observations strongly point to a genetic resistance to H. contortus. 5 Similarly in a multifactorial field trial Goossens et al. [16] showed significantly 6 higher nematode egg counts in Djallonke-Sahelian crossbreeds than in pure Djallonke 7 sheep, which indicates an innate resistance and/or a more efficient immune response 8 to nematodes. 9 Thirty years ago, in East Africa, Preston and Allonby [51;52] showed that Red 10 Maasai sheep are more resistant to H. contortus than other sheep breeds, which was 11 later confirmed and examined in more detail by Baker et al. [34;53]. Similarly, it was 12 shown that SEA does and kids were more resistant to GI nematodes than were Galla 13 does and kids [54;55] . 14 Although resistance to certain tick species (Amblyomma variegatum and 15 Hyalomma spp., but not Boophilus spp.) has been clearly demonstrated in N’Dama 16 cattle [56] almost no research has been carried out to study this aspect in 17 trypanotolerant small ruminants. In a longitudinal study over a period of five years 18 Odo [57] was not able to find any significant differences in tick loads between WAD, 19 Red Sokoto goats and their crossbreds. Unfortunately, Odo did not give any 20 information on the tick species involved. 21 22 23 Genetic purity of trypanotolerant sheep and goats Many countries have adopted livestock-improvement programmes based on 24 crossbreeding of endemic with exotic breeds. Furthermore, as mentioned above, there 25 is a tendency to increase the small size of the trypanotolerant small ruminant breeds 13 1 by crossbreeding with larger breeds resulting in an important introgression of genes 2 from trypano-susceptible breeds. Both approaches put the indigenous livestock breeds 3 at risk. Fortunately, in 1995, the ITC started a small ruminant breeding programme in 4 The Gambia, with the aim of improving the efficiency of meat production and the 5 level of trypanotolerance of WAD goats and Djallonke sheep. Analysis of the results 6 over a seven-year period (1995–2002) showed that the programme which was run 7 under low-input village conditions was very successful and resulted in clear genetic 8 improvement for body weight and growth rate [37]. 9 In order to further promote the use of endemic livestock as a sustainable 10 approach for livestock breeding in rural areas in West Africa a new regional project 11 has recently started (see Box 1). In addition to conservation of the local breeds and to 12 increase their productivity, the project takes in equal measure account of conservation 13 of their environment. This long-term multi- donor project aims at conserving and 14 promoting N’Dama cattle, Djallonke sheep and WAD goats and their unique genetic 15 traits, as well as critical habitat for these breeds in their regions of origin in West 16 Africa. By ensuring sustainable populations of targeted endemic ruminant livestock 17 breeds in four West African countries it will support human food production and 18 improve rural economies. It will be implemented by establishing models for 19 community-based management of endemic ruminant livestock and their habitat at 20 pilot sites, and strengthen production, marketing, and policy environments in support 21 of these breeds. The project aims to integrate relations between animal genetic 22 resources, ecosystems, production systems and human welfare. 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Small Ruminant Res. 50, 203-207 32 33 34 19 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Glossary Trypanotolerance is the ability to survive, reproduce and remain productive under trypanosomiasis risk without the need for using chemicals to control the vector or drugs to control the parasite [6]. As explained in the text trypanotolerance can either be innate resistance or resilience to trypanosome infections. Resistance is the ability of a host to initiate and maintain responses to suppress the establishment of certain parasites, in casu trypanosomes and/or eliminate the parasite load [32]. Since self-cure often occurs in bovines resistance is the appropriate term to characterise trypanotolerance in N’dama cattle. Resilience is the ability of the host to maintain a relatively undepressed production level under parasite challenge [32]. Djallonke sheep are able to gain weight and to reproduce even in the presence of a persistent parasitaemia. Therefore resilience is the most appropriate term to describe the phenomenon of trypanotolerance in sheep. Box 1. Project ‘Sustainable management of endemic livestock in West Africa’ 20 Project area: The Gambia, Mali, Senegal, Guinea Funding: African Development Bank, United Nations Development Programme, Global Environment Facility, the Governments concerned, FAO, CIRDES, ILRI Duration: 2008–2013 Some of the expected outcomes o Production and productivity of endemic ruminant livestock sustainably improved o Commercialisation and marketing systems of endemic ruminant livestock and livestock products strengthened o Natural resources in project pilot sites conserved, and sustainable managed for the benefit of endemic ruminant livestock, ecosystem services and human livelihoods o Legal, policy and institutional frameworks established at the local, national, and sub-regional level for in-situ conservation of endemic ruminant livestock o A sub-regional networking system established for cooperation, information exchange and coordinated support for the conservation of endemic livestock Some of expected realizations o Rehabilitation and equipment of five zoo-technical research stations with a view to re-launching genetic improvement programmes in the participating countries o Establishment of multiplication systems within 200 village herds in order to ease access by mixed farmers to selected breeding stock o Training of 15,000 livestock farmers including 8000 women in different domains (animal feeding, breeding management and 20 improvement of the habitat of small ruminants o Development and equipment of 17 slaughter areas, two sub-regional livestock markets and 17 local livestock markets o Construction and equipment of mini-dairies of a capacity of 200 to 600 l per day for women’s associations More info: Dr. B. Goossens (BartG@unops.org) 1 2 21 1 2 Figure 1. Trypanotolerant small ruminants at the International 3 Trypanotolerance Centre in The Gambia. (a) A flock of Djallonke sheep (photo by 4 Andreas Schönefeld©). 5 6 (b) West African Dwarf (WAD) goats (photo by Abdou Fall©) 7 22 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Figure 2. Population structures of indigenous goat populations in West Africa (red= WAD, green = Sahelian) (source: Hoeven et al., 2007-see footnote 2; Hoeven, E. 2004. Genetic characterisation of WAD goats, unpublished report, ITC, 17 pp.) Legend: geographic location of sampled West-African goat populations; for Senegal (SE): SESL=StLouis, SEF=Fatick, SEC=Casamance (Kolda-Sédhiou); for The Gambia (GA): GABG=BrikamaGunjur, GAK=Keneba; GAN=Niamina, GAS=Sololo; for Guinea-Bissau: GB=Gabù-Bafata; for Guinea (GU): sites A, B, C, D and E; for Mali (MA): MAM=Manankoro; MAS= Sagabory; MAT=Tousséguéla 72% Mauritania 31% 69% SAH SESL Table 1. Productivity indexa of Djallonke sheep and West African Dwarf (WAD) 41% 59% Senegal goats under an experimental infection with T. congolenseb 16 SEF 31% 31% 28% 53% Year 1 47% Control GAN Year 2 Mali Infected 57% 43% GAK 69% GABG 69% Control Infected 15.5 16.5 15.6 7.5 GAS Djallonke 27% 73% sheep WADSECgoats 72% 28% 13.3 4.9 11.1 1.2 58% MAS 42% GB Gambia 17 Productivity index : weight (kg) of 5 month old offspring produced per female a 33% 18 maintained per year 19 b 20 Data from Ref. [31] Guinea-Bissau 24% 67% GUA 76% GUB 81% 58% 42% 80% Guinea 31% 20% GUC 69% MAM GUE 57% Sierra Leone GUD Liberia 23 43% Ivory Coast MAT 19%