Livestock Science 144 (2012) Contents lists available at SciVerse ScienceDirect. Livestock Science

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1 Livestock Science 144 (2012) Contents lists available at SciVerse ScienceDirect Livestock Science journal homepage: Influence of dietary supplementation with Acacia karroo on experimental haemonchosis in indigenous Xhosa lop-eared goats of South Africa U. Marume a, M. Chimonyo b,, K. Dzama c a Department of Livestock and Pasture Science, University of Fort Hare, P Bag X1314 Alice 5700, South Africa b Discipline of Animal and Poultry Science, University of KwaZulu-Natal, P. Bag X01, Scottsville 3209, Pietermaritzburg, South Africa c Department of Animal Sciences, Stellenbosch University, P Bag X1, Matieland 7602, South Africa article info abstract Article history: Received 25 May 2011 Received in revised form 9 November 2011 Accepted 11 November 2011 Keywords: Faecal larval counts Packed cell volume Worm counts Alkaline phosphatise Serum glutamic oxaloacetic transaminase The effect of dietary supplementation with fresh Acacia karroo on experimental haemonchosis was determined in 4-month-old castrated Xhosa lop-eared kids. The goats were randomly allotted into four treatment groups: non infected, non supplemented (N); infected, non supplemented (); infected, supplemented () and non infected, supplemented (N). Each treatment group contained six goats. Each goat in the infected groups was exposed to a single dose of 6000 freshly cultured L3 Haemonchus contortus larvae. The kids in the supplemented groups received 182 g/day (on DM basis) of fresh A. karroo leaves, constituting 50% of their diet. Blood samples were collected every two weeks to determine haematology and serum enzyme concentrations. At the end of the experiment at day 60, all the kids were humanely slaughtered to determine the worm burdens. A significant decrease in faecal larval counts (FLC) was observed in the group as infection progressed while it continually increased in the group after 4 of infection. At necropsy, the group had significantly (Pb0.05) higher worm counts than the group (225.5±43.75 versus 25±43.75). Highest packed cell volume (PCV) values were observed in kids in the N group. The goats had the lowest PCV values. Mean FAMACHA scores for the and groups were higher (Pb0.05) than that of the N. Alkaline phosphatase levels (ALP) in the group increased as the infection progressed. On the contrary, in all other groups, ALP levels decreased up to 4 post infection. Supplemented goats had a significantly (Pb0.05) lower mean serum glutamic pyruvic transaminase (SGPT) and serum glutamic oxaloacetic transaminase (SGOT) values than the non-supplemented group. Results from the current study suggested that consumption of fresh A. karroo leaves reduced infection level and subsequently the leakage of enzymes into blood. It can be concluded that consumption of fresh A. karroo leaves reduced the establishment of H. contortus in Xhosa lop-eared goats Elsevier B.V. All rights reserved. 1. Introduction Corresponding author. Tel.: address: Chimonyo@ukzn.ac.za (M. Chimonyo). Sustainable goat productivity cannot be guaranteed in many communal production systems without the effective control of gastrointestinal parasites, especially Haemonchus contortus. H. contortus is endemic and causes substantial economic losses to resource-poor goat farmers in Sub-Saharan Africa (Bath, 2000; Perry et al., 2002; Torres-Acosta and Hoste, 2008; Vatta et al., 2009). In monetary terms, for example, in Kenya and South Africa annual losses due to this parasite were estimated to be between US$ 26 million and US$ 45 million (Krecek and Waller, 2006). The depressive impact of H. contortus on goat productivity derives not only from death and sub-optimal performance of goats, but also from the high costs of drugs for prophylaxis and treatment. The development of anthelmintic resistance also exacerbates the limitations of the use of drugs in the control of H. contortus (Jackson and /$ see front matter 2011 Elsevier B.V. All rights reserved. doi: /j.livsci

2 U. Marume et al. / Livestock Science 144 (2012) Coop, 2000; Nnadi et al., 2009). Moreover, use of chemotherapy has become a source of public concern due to the risk of accumulation of residues in meat products (Hoste et al., 2006). In severe infections, H. contortus causes anaemia manifested by low packed cell volume (PCV) (Yacob et al., 2008), hypoalbuminaemia and often depressed total blood protein content in goats (Ogunsanmi et al., 1994). The damage due to haemonchosis also result in the elevation of serum enzymes such as alkaline phosphatase (ALP), acid phosphatase (ACP), serum glutamic oxaloacetic transaminase (SGOT) and serum glutamic pyruvic transaminase (SGPT) (Sharma et al., 2001). Haematological values and serum biochemistry measures have often been used in predicting infections, monitoring goat health and provide information regarding immunological status, toxicant exposure, and other aspects of physiological functions in goats (Ogunsanmi et al., 1994; Stein et al., 1998; Vatta et al., 2001). A natural alternative to synthetic anthelmintics for controlling haemonchosis in goats is the use of tannin-rich plants (Min and Hart, 2003; Paolini et al., 2003; Terrill et al., 2009). Many rangelands in the resource poor areas of South Africa are characterized by large thickets of A. karroo shrubs. Although it is considered to be an invasive plant, A. karroo contains large amounts of polyphenols, especially, condensed tannins which have anthelmintic effects on H. contortus. Consumption of Acacia leaf meal has been reported to reduce the establishment and fecundity of H. contortus (Kahiya et al., 2003). The anthelmintic properties of A. karroo have also been established in cattle (Xhomfulana et al., 2009). In addition to its potential anthelmintic activity, A. karroo is a valuable protein source with crude protein of up to 230 g/kg (Nyamukanza and Scogings, 2008)andhas often been a major source of feed to browsing goats, especially during the dry season. Although they have been used in dried form in some experiments (Kahiya et al., 2003; Xhomfulana et al., 2009), fresh leaves from forages such as A. karroo are highly palatable and acceptable to browsing goats (Alonso-Diaz et al., 2008; Nyamukanza and Scogings, 2008). Nitrogen degradation is also higher in fresh forms of forages as compared with dry forms (Bonsi et al., 1994; Osuji and Odenyo, 1997). Although, the potential for manipulating host nutrition to favour the response to nematode infections has been pointed out (Hoste et al., 2006; Marume et al., 2010), and possible synergisms between various nutritional options for helminth control have been explored (Athanasiadou et al., 2009), the question that needs attention is are there any combinative effects of proteins and tannins in A. karroo on haemonchosis in browsing Xhosa lop-eared goats. The present study was, therefore, conducted to examine the effects of consumption of A. karroo leaves on live weight, body condition, haematological and biochemical changes in indigenous Xhosa lop-eared goats experimentally challenged with haemonchosis. The hypothesis tested was that nutritional supplementation with A. karroo reduces the establishment of H. contortus in Xhosa lop-eared goats. 2. Materials and methods 2.1. Study site, experimental design and treatments The study was conducted at the University of Fort Hare Farm, Eastern Cape Province, South Africa (32.8 S and 26.9 E). Twenty-four castrated 4-month old Xhosa lop-eared goats with an average body weights between 13.5±0.23 kg (mean±se) were used. The experiment was a 2 2 (infection level dietary level) factorial arrangement with two levels of nutrition (supplemented versus non supplemented) and two levels of Haemonchus infections (infected versus non infected). The kids were randomly allotted into four treatment groups: non infected non supplemented (N); infected, non supplemented (); infected, supplemented () and non infected supplemented (N). Each pen contained six goats. Supplemented kids were fed in individual cages Collection and analysis of A. karroo leaves Fresh A. karroo leaves were hand harvested each day to feed the goats for 60 days. The fresh leaves were given to the kids from day 0 (day of infection) up to day 60, at the end of the experiment. Harvested leaves were stored in plastic sacks to reduce moisture loss. The leaves were also sprinkled with water to maintain freshness. The goats were fed individually in feeding troughs in cages. A sample of the fresh leaves was obtained and dried for the determination of proximate composition and tannin levels in the leaves. The samples of feed were analyzed for DM, crude protein (CP), crude fibre (CF) and ether extract (EE) according to Association of Official Analytical Chemists (1990). The butanol-hcl assay as described by Giner-Chavez et al. (1997) was done to determine the condensed tannins (CT) while the Folin-Ciocalteau assays described by Terrill et al. (1992b) was performed to determine the total polyphenolic content of the dried A. karroo. The proximate analysis and tannin levels of A. karroo leaves are shown in Table Goat management From birth until weaning at approximately three months of age, the kids were raised indoors. At one month of age, the kids were separated from their dams late in the morning when the dams would be turned out to the pastures for grazing, but were allowed to be with their dams late in the afternoon and overnight for them to feed. At weaning, the kids were grouped together and housed in an open sided barn for a period of 30 days after which they were allocated to one of the four treatments. During that time the kids were offered a basal diet of 460 g/head/day (DM) of Medicago sativa hay (CP, 203 g/kg: CF, 335 g/kg; Table 1) to meet their maintenance and growth requirements for protein and energy Table 1 Nutritional composition of Medicago sativa and fresh Acacia karroo leaves used in the study diets (% DM basis). Component Medicago sativa Acacia karroo Moisture content (fresh leaves) 47.6 Dry matter Crude protein Crude fibre Neutral detergent fibre Acid detergent fibre Ether extract Calcium Phosphorus Ash CT (Butanol-HCl assay) 2.1 Total phenolics (Folin Ciocalteu assay) 0.5

3 134 U. Marume et al. / Livestock Science 144 (2012) (80 g/day CP and 5.69 MJ/day ME), according to the National Research Council standards (NRC, 2007). The goats had ad libitum access to water. After they were allocated into the four different treatment groups, the kids in the supplemented groups received an additional 200 g per head per day of fresh A. karroo leaves collected each day for two months. The fresh A. karroo leaves were given to the goats individually in feeding troughs in two equal portions both in the morning and afternoon. The kids in the non supplemented groups continued to receive 500 g/day of lucerne. A state veterinarian examined the kids every week throughout the trial and also assured that the welfare standards were met Infection protocol Infective larvae (L3) of H. contortus (95% pure) were obtained from donor sheep infected with a pure strain of nematode at Onderstepoort Veterinary Institute (South Africa). On day 0 of the experiment, each of the goats in the infected groups was exposed to a single dose of 6000 L3 H. contortus larvae according to infection protocols used by Yacob et al. (2008) and Sharma et al. (2001). The experiment was conducted for 60 days Measurements Body weight and condition scores After every two weeks, the kids were weighed using the commercial goat weigh scale in the morning at 0800 h, faecal samples collected for faecal cultures and blood samples taken for haematological examination. Concurrently, body condition scores (BCS) were assessed by one researcher throughout the experiment using the 5 point scale (1 = very thin to 5 = obese) (Aumont et al., 1994) Faecal cultures, larval counts Faecal cultures were prepared for each kid in each treatment for faecal larval counts (FLC). Approximately 5 g of fresh faecal samples were obtained directly from the rectum of the kids at the beginning of the experiment and fortnightly thereafter. The faecal matter was spread into a plastic pan at less than 3 cm deep. Faecal matter was then covered with peat moss and misted with tap water to a moist consistency and covered with lid. Pans were incubated for 7 10 days at 27 C. The temperature was maintained throughout the experiment to reduce discrepancies. The faecal cultures were then placed into a Baermann apparatus to extract larvae according to the Baermann method (Sloss et al., 1994). Larvae were counted using a dissection scope at 7.5 magnification and averaged over three counts to determine the final concentration FAMACHA scores Parallel to the blood and faecal sample collection, the colour of the ocular mucous membranes of each of the kids was be examined and classified into one of five categories according to the FAMACHA eye colour chart: 1 = red, non-anaemic; 2 = red-pink, non-anaemic; 3 = pink, mildly-anaemic; 4 = pinkwhite, anaemic; 5 = white, severely anaemic (Bath, 2000) Haematology Packed cell volume was measured by the microhaematocrit method, while the differential leucocyte count (TLCA), mean corpuscular volumes (MCV) and mean cell haemoglobin concentrations (MCHC) were determined at the National Health Laboratory Services (NHLS), East London, South Africa Serum enzymes Quantitative estimation of alkaline phosphatase (ALP), serum glutamic pyruvic transaminase (SGPT) and serum glutamic oxaloacetic transaminase (SGOT) were done within 24 h of collection at the National Health Laboratory Services (NHLS) (South Africa). The enzymes were analyzed using a clinical chemistry analyzer (Gilford Impact, 404lE, Ciba Coming Diagnostic Corp., Gilford Systems, Oberlin, OH 44774). Enzyme assays were all performed on an UV vis spectrophotometer (SPECORD 50 PC, Analytik Jena AG) using respective commercial kits following the protocol described previously by Ogunsanmi et al. (1994) Slaughter procedures At day 60, all the goats were humanely slaughtered at a local abattoir for worm recovery from the abomasa. A day before slaughter, the goats were deprived of feed for 24 h. Clean water was provided ad libitum. The electrical stunner was used to stun the goats and then slaughtered using standard procedures. Skinning, evisceration and washing procedures were completed while the carcasses were on the overhead rail. The abomasa were removed and tied at both ends for worm recovery at the University laboratory Worm counts at necropsy At necropsy, the abdominal cavity was opened and abomasa were removed. The abomasa were opened and the contents emptied and washed with tap water into a bucket to make up 5 l. After thorough stirring, a 500 ml aliquot (10%) was collected into labelled plastic bottles and then examined to estimate the size of the worm populations. After settling for 5 6 h, approximately 50 ml was decanted and replaced with approximately 50 ml formalin as a preservative. The washed abomasa of each goat was soaked in water over night at room temperature and re-washed in 5 l of water. Subsequently, soak samples were collected for worm recovery, identification and enumeration (Fakae et al., 1999) Statistical analyses The differences in FLC, PCV, MCV, eosinophil counts, TLCA, MCHC, BCS, body weights and FAMACHA scores, ALP, SGOT and SGPT between the treatment groups were analysed using mixed model procedures for repeated measures of SAS (2003) according to Littell et al. (1996). Faecal larval counts and worm count data were log transformed [ln (χ+10)] and the resulting transformed variables were tested for normality using probability plots, skewness and kurtosis. The transformed data were reported as back-transformed means. Worm count data were analysed using the RANDOM statement in the mixed model procedure. Only and treatments

4 U. Marume et al. / Livestock Science 144 (2012) included in the FLC and worm count analyses, as the other two treatment groups tested negative for FLCs and worm counts. The linear model used was: Y ijk ¼ μ þ T i þ P j þ W k þ ðt P WÞ ijk þ E ijkl where: Y ijk observation (FLC, PCV, MCV, MCHC, TLCA, eosinophil counts, BCS and body weights) μ population mean constant common to all observations; Ti fixed effect of diet; P j fixed effect of infection; W k effect of week of sample collection (T P W) ijk diet x infection x week interaction; E ijkl random error term, assumed to be normally and independently distributed. Mean separations were determined using the PDIFF option in SAS (2003). supplemented kids maintained body weights between 0 and 6 post infection, sharply declining thereafter. In all groups, a gradually decline in BCS was observed up to the 4th week post infection coming to a constant thereafter. The decline in BCS was however, more pronounced in the group. The group had the lowest (Pb0.05) BCS than the other groups throughout the experiment Faecal larval counts Infection by H. contortus was expected to become patent in the infected groups three weeks post infection. Infection, diet and their interactions significantly (Pb0.05) affected faecal larval counts (FLC). An increase in FLC was observed in both infected groups up to 4 post infection (Fig. 1). The increase, however, was gradual in the treatment group but more pronounced in the group. The FLC decreased significantly (Pb0.05) in the treatment group after 4. On the contrary, FLC in the treatment group continued to increase until the end of the experiment. 3. Results 3.1. Body weights and body condition scores Changes in body weights and BCS are shown in Fig. 1.Infection, diet and their interaction significantly affected body weights. A sharp increase (Pb0.05) in body weight was observed in the N group between 0 and 2 post infection becoming constant, thereafter. In the and N groups, an increase in body weight was observed throughout the experimental period. The N group had the highest (Pb0.05) mean body weight at the end of the experiment. Infected but not 3.3. Worm counts At necropsy, the group had significantly higher (Pb0.05) worm counts than the group (225.5±43.75 versus 25±43.75). Mature worms constituted approximately 97% of the total worm counts at necropsy in the two infected groups FAMACHA scores Diet and diet infection interaction significantly (Pb0.05) affected FAMACHA scores. Throughout the study period, FAMA- CHA score for both the and groups were higher (Pb0.05) than that of the uninfected group (N) (Table 2). The N Faecal larval counts PCV (%) N 20 N 10 4 BWT (KG) N N 5 BCS N N 2 Fig. 1. Mean changes in faecal larval count (FLC), packed cell volume (PCV), body condition score (BCS) and body weight (BWT) of goats in different dietary and infection groups.

5 136 U. Marume et al. / Livestock Science 144 (2012) Table 2 Least square mean haematological values and FAMACHA scores of the goats in infected, non supplemented (); N, non infected, non supplemented (N), infected, supplemented () and N, non infected, supplemented (N) treatment groups. Parameter N N SEM Significance Mean corpuscular 63.8 c 56.9 b 53.1 a 58.9 b 2.19 * volume Total leucocyte 18.1 c 15.1 b 17.2 c 11.9 a 0.95 * count Eosinophils NS FAMACHA 2.10 ab 2.40 b 2.96 c 1.96 a 1.18 * abc Values in the same row with different superscripts are significantly different (P b0.05). *Pb0.05. group recorded the lowest FAMACHA score (1.96±1.18) while the highest (2.96±1.18) was in the group. No difference was observed in FAMACHA scores between the and the N groups Haematology All observed differences in PCV values between groups were significant (Fig. 1). Although there was an initial increase in PCV in the and the N groups between 0 and 2 post infection, it declined marginally in the N group. From 4th week onwards, however, the PCV for goats in the group were lower than those of the N group. A gradual decline in PCV was observed in the group throughout the experiment. While there were no differences (Pb0.05) observed between the N, and the N group, PCV values for these three groups were significantly higher than those of the group. Values for MCV, TLCA and eosinophil counts are shown in Table 2. Mean corpuscular volume values for the N and groups were initially lower than that of the group, but by 4 onwards, MCV values for the two groups had significantly (Pb0.05) increased to levels above that of the group. Highest meat MCV values were observed in the N group while the lowest was observed in the group. No differences in eosinophil counts were observed, while TLCA was, however, higher (Pb0.05) in the N and N groups Serum enzymes Infection, diet and their interaction significantly affected ALP. Variations in biochemical responses to infection and dietary supplementation are shown in Fig. 2. Alkaline phosphatase levels in the group increased as the infection progressed. On the contrary, in all other groups, ALP levels decreased up to 4 post infection. The ALP levels, however, gradually increased in the group after 4 post infection but remained constant in the N and N groups. Generally, ALP values of infected goats were significantly higher (Pb0.05) than those of the non infected groups (Table 2). In addition, goats in the had lower (Pb0.05) ALP values than those not supplemented (93.4±7.69 IU/l versus 156.3±7.69 IU/l). Infection had no effect on SGPT (Pb0.05). Supplemented goats, however, had a significantly lower mean SGPT value than the non supplemented group (20.3±1.25 IU/l versus 25.3±1.25 IU/l). Infection, diet and their interaction significantly (Pb0.05) affected SGOT values. The infected goats had significantly higher (Pb0.05) SGOT values than the non infected group (Fig. 2). The N groups had the lowest SGOT values throughout the experiment. In the group, a sharp increase in SGOT levels was observed while in the group, a gradual increase in SGOT levels was observed after 6 post infection. 4. Discussion The study intended to evaluate the efficacy of the use of fresh A. karroo leaves as an alternative approach in the control of haemonchosis in goats. One key element in the use of fresh Acacia leaves as both supplement and anthelmintic are their palatability and acceptability to goats. Goats do not prefer dried leaves (Kahiya et al., 2003). The goats used in the current study had no browsing experience. As such, the high consumption of the Acacia leaves throughout period confirmed the Acacia leaves as highly acceptable and palatable. Various studies also showed similar observation (e.g. Alonso-Diaz et al., 2008; Nyamukanza and Scogings, 2008). Acacia karroo is widespread in the drought-prone Eastern Cape province of South Africa and has been one of the major sources of feed for browsing goats. Xhosa lop-eared goats are more of browsers than grazers (Bakare and Chimonyo, 2010). It has higher body frame than other indigenous goats and long front legs making it possible to reach out to leaves of tree branches. Responses to fresh A. karroo leaves could be useful in making strategies on its management and control, as it is generally regarded as an undesirable plant that reduces grazing capacity of the veld. The marked decrease in FLC and low worm counts at necropsy in goats that consumed the Acacia leaves highlight that they possess anthelmintic properties. A. karroo leaves contain various types of secondary plant metabolites, including condensed tannins (Hoste et al., 2006; Kahiya et al., 2003). The ability of condensed tannins to bind to proline- and hydroxyproline-rich structures that form the body and internal organs of the nematode and to change their physical and chemical properties makes them potential inhibitors of Haemonchus infections (Anthony et al., 2005; Brunet et al., 2007; Kahiya et al., 2003). Inthecurrentstudy,thereductionintheFLCcouldhavebeen associated with the tannin-induced reduction in fecundity of the worms. Condensed tannins may interfere directly with the biology of various nematode stages or indirectly, by improving the host nutrition through protection of the diet proteins from ruminal degradations thereby modulating worm biology (Paolini et al., 2003; 2005). Contact with tannin extracts affects the establishment of third-stage larvae, either by disturbing the exsheathment (Brunet et al., 2007) or the association of parasite with the mucosae (Brunet et al., 2008). In addition to the widely held view that Xhosa lop-eared goats are resistant to haemonchosis (Marume et al., 2010), further studies need to be conducted to establish the interaction of genes for resistance and effect of tannins. The subdued increase in initial body weights in the N and groups compared to the N during the first 2 weeks of the trial was expected. Consumption of high concentrations of condensed tannins (>7% of dry matter) has often been associated with a reduction in feed intake, growth inhibition and interference with the morphology and the proteolytic activity of microbes in the rumen (Min et al., 2003; Waghorn and McNabb,

6 U. Marume et al. / Livestock Science 144 (2012) ALP (IU/L) N N SGOT (IU/L) N N 40 SGPT (IU/L) N N 10 Fig. 2. Least square mean faecal larval counts, alkaline phosphatase (ALP), serum glutamic oxaloacetic transaminase (SGOT), serum glutamic pyruvic phosphatase (SGPT) levels of infected and non infected Xhosa goats. 2003). Consumption of moderate or low concentrations of condensed tannins (b6% of DM), however, has been observed to be increase growth and milk production in browsers (Rochfort et al., 2008). In the current study, as the experiment progressed, both groups of goats that received dietary supplementation gained more weight than those on the basal diet and this can be explained in two ways. Firstly, as explained above, condensed tannins in A. karroo leaves may have reduced the establishment and effects of haemonchosis resulting in normal body function and hence the expected weight gain. Secondly, A. karroo leaves contain considerable amounts of protein (>23%/DM) compared to other forage legumes (Nyamukanza and Scogings, 2008). Its potential as a protein source in browsing goats has been reported earlier (Aganga et al., 2000; Goromela et al., 1997). Similar reports have been made for cattle (Mapiye et al., 2010). Therefore, the suppression of infection elicited by the Acacia leaves suggests a combinative effect of tannins and proteins that resulted in increased growth of the goats. The decrease in PCV in the infected groups from 4 onwards was expected. This period coincided with the full establishment of the infection. However the initial increase in PCV in the group is difficult to explain. It is probable that the infection had not been established, indicating that the goats had no worm eggs at the commencement of the experiment. The haematophagous nature of H. contortus is associated with anaemia and, hence, reduction in PCV. In animals, PCV changes are linked to the intensity of H. contortus infection (Marume et al., 2010; Yacob et al., 2008). Goats in the group maintained PCV levels, probably due to the suppression of the infection by the condensed tannins in the Acacia leaves. To support the impact of tanniferous forages, goats fed on sericea lespedeza hay showed lower FEC and a sustained PCV after 35 days of infection (Moore et al., 2008; Shaik et al., 2006). High PCV values observed in the N group could be due to the effect of the A. karroo leaves on both boosting the immune system and providing additional proteins to the goats. The high PCV values in this group are also consistent with the observed high body weights compared to the other groups. High MCV values obtained in the, N and N are consistent with the absence of infection, in the case of N and N group and the suppression of the infection by the A. karroo dietin thecaseof thegroup.changes in the levels of MCV, MCH and MCHC are expected to indicate fluctuations in erythropoietic activities due to infection. The infection is manifested by increased blood loss and low levels of haemoglobin in erythrocytes. The leucocyte differential aids in the assessment of the leucocytosis. Leucocytes have an important role in the resistance against nematode infection. In the current study, the TLCA in the group were higher than that of the uninfected groups (N and N). FAMACHA scores, PCV, body weight changes and body condition have been used to evaluate subclinical haemonchosis and also resistance of goats to parasites (Baker et al., 1998; Van Wyk and Bath, 2002). In the current study, the FAMACHA scores for the uninfected goats were lower than in infected groups, as expected. However, no significant difference was observed between the and N groups. On the contrary, the FAMACHA scores for the were significantly higher than those of the other three treatment groups. High FAMA- CHA scores in the revealed the effects of the infection. In addition, the high FAMACHA scores observed in this group are consistent with the high FLC, worm counts at necropsy, low body weight gain and low PCV observed in this group. The FAMACHA system has been extensively tested in South Africa with some excellent results and has been used in identifying infected animals for treatment (Kaplan et al., 2004; Van Wyk and Bath, 2002). Its use in goats has been validated by Vatta et al. (2001). Serum biochemistry measures have often been used in predicting infections, monitoring goat health and provide information regarding immunological status, toxicant exposure, and other aspects of physiological functions in goats (Ogunsanmi et al., 1994; Stein et al., 1998; Vatta et al., 2001). Serum enzyme

7 138 U. Marume et al. / Livestock Science 144 (2012) levels are likely to change in different clinical conditions (Sharma et al., 2001). Information obtained from serum biochemistry measures in infected and non infected goats can therefore provide excellent basis for judgement with respect to the nature of the disease and the extent of tissue and organ damage (Tibbo et al., 2008). Although serum enzymes are of value in detecting liver diseases, they can also be used to establish some disruptive activities of haemonchosis and other parasites in the organs of their origin or altered membrane permeability through traumatic damage to the lining of the abomasal mucosa (Al-Zubaidy et al., 1987). The low ALP values in the groups compared to the NSI goats suggest the role played by A. karroo supplementation in reducing the damage caused by H. contortus. The ALP values for the group were comparable to the N and the N goats. Generally, the haemolytic activity of H. contortus culminates in the invasion and damage to the abomasal mucosa, causing the enzymes to be leaked into blood. This was confirmed by the high values for ALP observed in the group. Similar observations were made by Sharma et al. (2001) in Barbari goats, and in parasitized West African Dwarf goats by Mbuh and Mbwaye (2005). Although protein supplementation of parasitised goats has been shown to reduce the level of haemonchosis (Hoste et al., 2006; Phengvichith and Ledin, 2007), the effect of condensed tannins in forage legumes has received little attention. The ALP in the, N and N were lower than those reported by Mbuh and Mbwaye (2005), but comparable to those reported by Sharma et al. (2001). Elevated levels of these enzymes normally indicated an increase in haemolytic activities of H. contortus in infected goats (Sharma et al., 2001). The effect of A. karroo on serum biochemical response to haemonchosis is also highlighted by the low SGOT values obtained in the group compared with the group. Siddiqqua et al. (1990), Chakraborty and Lodh (1994) and Sharma et al. (2001) reported an increase in SGOT levels in goats infected with haemonchosis. In the current study, A. karroo reduced the haemolytic activity of H. contortus resulting in low SGOT levels, as also observed with the ALP in the group. Serum glutamic oxaloacetic transaminase levels in the infected and non infected goats were also comparable to those reported elsewhere (Mbuh and Mbwaye, 2005; Sharma et al., 2001). Together with acid phosphates, ALP, SGOT and SGPT are some of the enzymes that are used as tools in diagnosis of clinical disease condition. 5. Conclusions Consumption of fresh A. karroo leaves reduced infection establishment, resulting in an increase in PCV and a reduction in enzyme leakage into blood Xhosa lop-eared goats supplemented with fresh A. karroo leaves. Therefore, A. karroo can be regarded as an important feed resource that mitigates the effects of haemonchosis on goat productivity. Conflict of interest statement All authors have ready through the document and had no objections to the publication of the manuscript. References Aganga, A.A., Adogla-Bessa, T., Omphile, U.J., Tshireletso, K., Significance of browses in the nutrition of Tswana goats. Arch. Zootec. 49, Alonso-Diaz, M.A., Torres-Acosta, J.F.J., Sandoval-Castro, C.A., Hoste, H., Aguilar-Caballero, A.J., Capetillo-Leal, C.M., Is goats' preference of forage trees affected by their tannin or fiber content when offered in cafeteria experiments? Anim. Feed. Sci. 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